Pelagic Amphipods of the

Genus Hyperia and

Closely Related Genera

(Hyperiidea: Hyperiidac)

THOMAS E. BOWMAN

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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 13 6

Pelagic Amphipods of the

Genus Hyperia and

Closely Related Genera

(Hyperiidea: Hyperiidae)

Thomas E. Bowman

SMITHSONIAN INSTITUTION PRESS

City of Washington

1973
ABSTRACT

Bowman, Thomas E. Pelagic Amphipods of the Genus Hyperia and Closely

Related Genera (Hyperiidea: Hyperiidae). Smithsonian Contributions to

Zoology, number 136, 76 pages, 1973.—The genus Hyperia is restricted to large

species with no fusion of pereonites, mandibular palp present in both sexes, and

coxae not fused with pereonites. Eight species are recognized, of which two are

new, and one, Hyperia macronyx, is transferred to Hyperiella. The small species

of Hyperia, with some pereonites fused, mandibular palp absent in female, and

coxae fused to pereonites, are assigned to Hyperioides (1 species); Lestrigonus

Milne Edwards, restored from synonymy (6 species, 1 new); Hyperietta, new

genus (5 species, 3 new); Themistella (1 species); and Hyperionyx, new genus

(1 species). Keys are given to the species of each genus. Available information

on distribution is summarized for each species.

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

in the Institution's annual report, Smithsonian Year.

Library of Congress Cataloging in Publication Data

Bowman, Thomas E.

Pelagic amphipods of the genus Hyperia and closely related genera (Hyperiidea: Hyperiidae)

(Smithsonian contributions to zoology, no. 136)

Bibliography: p.

1. Hyperia. 2. Hyperiidae. I. Title. II. Series: Smithsonian Institution. (Smithsonian contributions to zoology, no. 136)

QL1.S54 no. 136 [QL444.A5] 591.08s [595'.371] 72-8907

For tale by the Superintendent of Document*, U.S. Government Printing Office

Washington, D.C. 20402 - Price $2 (paper cover)
Pelagic Amphipods of the

Genus Hyperia and

Closely Related Genera

(Hyperiidea: Hyperiidae)

Thomas E. Bowman

Introduction

A comprehensive taxonomic treatment of the

amphipod family Hyperiidae has not appeared

since the 1889 monograph of Bovallius. Important

contributions to the taxonomy of this family have

been made by Vosseler (1901), Stephensen (1924),

Barnard (1930, 1932), and Yang (I960), but the

need for an up-to-date revisionary study remains

unsatisfied. In my studies of the Hyperiidae

collected during the California Cooperative Oceanographic Fisheries Investigations (CalCOFI), I

encountered several undescribed species as well as

taxonomic problems that I felt unable to resolve

without making comparative studies of hyperiids

from other parts of the world. I therefore published

only the results of my investigations of the genus

Parathemisto (Bowman, 1960) and have delayed

until now the presentation of a revision of Hyperia

and closely related genera.

The present work considers the genus Hyperia

and the species now assigned to it. Some of these

species are transferred herein to currently recognized genera (Hyperiella, Hyperioides), to genera

now considered synonyms of Hyperia (Lestrigonus,

Themistella), and to new genera {Hyperietta,

Thomas E. Boxvman, Department of Invertebrate Zoology,

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

Hyperionyx); all these genera are reviewed in this

paper. The other genera of Hyperiidae include

three monotypic genera (Bougisia, Pegohyperia,

Phronimopsis), Iulopis with two species, and

Hyperoche. Of these only Hyperoche presents

obvious taxonomic problems, and a monographic

treatment of it will be offered later.

MATERIAL.—For this investigation I have had

available the amphipod collections of the Division

of Crustacea, Smithsonian Institution, obtained

from many sources. Their careful curation and arrangement for maximum convenience by the late

Clarence R. Shoemaker has facilitated my work

considerably. Several recent major collections of

hyperiids used extensively in this study and now

deposited in the Division of Crustacea are from the

following:

CalCOFI Cruises 1, 5, 9, 20, off the Pacific coast

of the United States from the latitude of the

Columbia River to that of Punta Eugenia in middle

Baja California, made available to me by Dr.

Martin W. Johnson, Scripps Institution of Oceanography.

Cruises of the University of Washington R/V

Brown Bear in the northeastern Pacific, sent to me

by Dr. William I. Aron.

Cruises 12, 13, 14 of R/V Ombango in the Gulf

of Guinea, sent to me by Dr. Alain Crosnier, Office

de le Recherche Scientifique et Technique Outre

Mer (ORSTOM) Republique du Congo.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Collections of Hyperiidae by ships participating

in the International Indian Ocean Expedition,

1962-65 (HOE), sorted from plankton samples and

sent to me by Mr. Chandrasekharan Nair, Indian

Ocean Biological Centre.

I am most grateful to the persons listed above, as

well as to the following, who have made it possible

for me to examine specimens from their museums:

Drs. R. W. Ingle and Roger Lincoln, British

Museum (Natural History); Dr. H.-E. Gruner,

Zoologisches Museum, Berlin; Dr. Charlotte

Holmquist, Naturhistoriska Riksmuseet, Stockholm;

Dr. Torben Wolff, Universitets Zoologiske Museum,

Copenhagen.

ZOOGEOGRAPHY.—The above collections have enabled me to extend the known distribution of a

number of species and to give some idea of the

overall pattern of distribution of other species. The

principal aim of this study, however, has been to

improve our understanding of the taxonomy of

the Hyperiidae, and the comments on species

distribution will perhaps be of most service in

indicating the gross inadequacy of our current

knowledge of the distribution of pelagic amphipods.

For some of the species occurring in the CalCOFI

area, I have tentatively extrapolated from distributions there to overall Pacific distribution patterns

found by Brinton (1962) to be characteristic of

species of euphausiids.

TAXONOMICALLY USEFUL CHARACTERS.—TO assist

the reader, a diagram of a typical <? hyperiid, with

the parts labeled, is given in Figure 1.

-PEREON

elson

FIGURE 1.—Diagram of a $ hyperiid am phi pod.
NUMBER 1S6

The following abbreviations are used throughout

this paper:

Al, A2 = antenna 1 and 2

Md = mandible

Mxl, Mx2 = maxilla 1 and 2

Mxp — maxilliped

P1-P7 = pereopods 1-7

sl-s7 = segments 1 (coxa) to 7 (dactyl) of pereopods

Upl-Up3 = uropods 1-3

In arranging the species of Hyperia, Bovallius

(1889) separated them initially according to the

degree of fusion of the pereonites. His scheme,

which was followed by subsequent workers, remains

basically valid and is used herein. Bovallius believed that the amount of pereonite fusion in a

species did not vary with sex or age. He reached

this conclusion after examining young specimens of

Hyperia faberi (=Hyperietta vosseleri in this

paper), one of several species comprising the new

genus Hyperietta, in which pereonites 1-2 are

fused in both sexes and in all growth stages. As

shown by Yang (1960) and Laval (1968), the amount

of fusion varies with age and sex in certain other

species (those included here in Lestrigonus). This

variation will be discussed in the accounts of the

individual species, but the following summary may

be helpful:

TABLE I.—Fusion of Pereonites in the Family Hyperiidae

None 1-2 9 $ 1-3 9 $ 1-5 9 1-3, $ 1-2 9 1-4, $ 1-2 9 1-5, $ 1-4

Hyperia

Hyperiella

Hyperoche

Iulopis

Parathemisto

Pegohyperia

Bougisia

Hyperietta

Hyperioides

Phronimopsis

Hyperionyx Themistella Lestrigonus crucipes L. latissima L. bengalensis

L. schizogeneios L. macrophthalma

L. shoemaker!

Other characters emphasized by previous workers

are the form of the head, the armature of the

pereopods and uropods, the size and shape of the

telson, and to a limited extent the mouthparts,

especially the presence or absence of the Md palp

in the 9. The ventral process" of the head has

received some attention, but it has not been emphasized that this process is actually the gland cone

bearing the orifice of the antennal gland. In

gammaridean amphipods it can be easily seen that

the gland cone opens on the second of the 5 free

peduncular segments of A2. In the Hyperiidae the

gland cone and the segment on which it opens are

fused to the surface of the head. In 9 Hyperia and

Hyperiella the gland cone segment is followed by 2

short free peduncular segments and a longer

1-merous flagellum (Figures 8a, 18c, 20c, 22b). In

Hyperioides there is only 1 free peduncular segment

(Figure 24d), and in Lestrigonus, Hyperietta,

Themistella, and Hyperionyx the entire A2 is reduced to a single rudimentary free segment. Evidently the first 2 antennal segments are always

fused with the head in $ Hyperiidae, but the pattern of fusion among the other segments is not

known.

The size and shape of the gland cone are helpful

taxonomic characters, especially in the genus

Lestrigonus. The extent to which it overlaps or is

separated by a gap from the buccal mass, when

viewed laterally, is also useful.

Among the mouthparts I have found the Mxp

to be especially valuable taxonomically. The outer

lobes are separate and elongate in Hyperia,

Hyperiella, and Lestrigonus; fused medially and

rather broad in Hyperietta; separate and obovate

in Hyperionyx. The inner lobe is greatly reduced

in Hyperietta and Themistella. The most convincing evidence that Lestrigonus fabrei Milne Edwards

is not the same species as that called H. fabrei by

subsequent authors (=H. vosseleri Stebbing) lies

in the entirely different form of the Mxp of the

two species (compare Figures 27c and 41/). Curiously, this difference has not been pointed out

previously, not even by Stebbing, who established

H. vosseleri because he thought it too small to fit

into Milne Edwards' species.

The armature of the dactyls and distal margin of

the propus of P5-7 offer characters of generic

significance. The propal margin may be produced

into a spine {Lestrigonus; Figures 29e; 33fc,/; S5p,
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

r; 36A) or a spinulose lobe (Hyperietta; Figures

44/J; 47d, e) that overlaps the dactyl medially; in

Lestrigonus crucipes (Figure 31a) the dentate spine

immediately identifies the species. The presence or

absence of spinules on the proximal part of the

occluding margin of the dactyl is also a dependable

character.

Family HYPERIIDAE Dana, 1852

Head usually globular; eyes almost always

occupying most of lateral surface of head. A1-2

short in $, with 1-segmented flagellum; long in

cf, with slender, multisegmented flagellum. Md

palp present in cf, often absent in 9 • Outer lobes

of Mxp free (most genera), or fused medially

(Hyperietta). PI simple, subchelate, or chelate;

P2 chelate. P3-4 similar, sometimes prehensile.

P5-7 similar, sometimes elongate, usually not

prehensile. Gills on pereonites 2-6. Urosomites

2-3 fused. Up 1-3 with 2 rami. Type genus:

Hyperia Latreille.

Sexual dimorphism is marked in the Hyperiidae,

and at one time males of the genus Hyperia were

placed in a separate genus, Lestrigonus Milne

Edwards. In addition to the differences in Al-2,

the female has a shorter and plumper body. The

male body is more elongate, owing to the greater

development of the pleon and pleopods, and the

male is obviously the more active sex. Although

nothing is known about reproductive behavior in

these amphipods, the greater development of

sensory and locomotor structures in the male indicates that he actively seeks out the female. The

great preponderance of males in collections made

under a light at night supports this proposition.

Key to Genera of the Family Hyperiidae

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

At least pereonites 1-2 fused. Coxae fused with pereonites (except in Bougisid) 7

2 Eyes without facets. Head produced anteriorly into acute process between Al and A2

Pegohyperia Barnard

Eyes normal, with facets. Head not produced between Al and A2 3

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

lulopis Bovallius

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

4. Pl-2, carpal process laterally compressed, knife-shaped, without spines Hyperoche Bovallius

Pl-2, carpal process spoon-shaped, with somewhat concave center and raised margins

armed with spines 5

5. P3-4 prehensile, dilated s5 closing against s6. Parathemisto Boeck

P3-4, s5 not dilated 6

6. P5 or P5-6 longer than P3-4 II. Hyperiella Bovallius

P3-4 longer than P3-6 1. Hyperia Latreille

7. Eye small. Head produced anteriorly into acute process between Al and A2. Coxae

distinct Bougisia Laval

Eye covering most of head surface. Head not produced between Al and A2. Coxae fused

with pereonites 8

8. First 2 pereonites fused in adult $ and 9 11

First 3, 4, or 5 pereonites fused in adult 9; first 2, 3, or 4 pereonites fused in adult $ .... 9

9. P5-7 subequal; P6 only slightly longer than P5 and P7. Telson of moderate size

IV. Lestrigonus Milne Edwards

P5 much shorter or distinctly longer than P6-7. Telson very short 10

10. Pereonites 1-3 fused. P5 much shorter than P6-7. VII. Hyperionyx, new genus

Pereonites 1-5 fused. P5 much longer than P6-7 VI. Themistella Bovallius

11. P2 chelate; s7 closing against process of robust s6; P3-7 prehensile Phronimopsis Claus

P2 chelate, s6 closing against process of s5. P3-7 nonprehensile 12

12. 9 A2 moderately long, shape sinuous in profile. Md incisor with serrate apex. Mxp with

well-developed inner lobe. Exopods of Upl-3 with notched lateral margins

III. Hyperioides Chevreux

9 A2 rudimentary. Md incisor with smooth apex. Mxp with rudimentary inner lobe.

Exopods of Upl-3 without notches V. Hyperietta, new genus
NUMBER 136

I. Hyperia Latreille, in Desmarest, 1823

DIAGNOSIS.—Large species, 10-30 mm in length.

Head deeper than long. Pereon more or less dilated

in 9 > aH pereonites free. Coxae not fused to

pereonites. Al peduncle 3-merous; $ flagellum

1-merous. A2 peduncle 3-merous; 1st segment

fused with gland cone. Md with serrate molar;

palp present in both sexes. Mxl, outer lobe with

5 apical spines; palp with robust spine at mediodistal corner. Inner lobe of Mxp well developed,

usually with 2 terminal spines. PI subchelate or

barely chelate, carpal process only slightly developed. P2 chelate; carpal process spoon-shaped,

bearing spines along edges of spoon. S5-6 of Pl-2

with many spines. P5-7 shorter than P3-4. Inner

ramus of o* Up3 broad.

DERIVATION OF NAME.—From Hyperia, a fountain

at Pherae in Thessaly. Gender feminine.

TYPE-SPECIES.—By objective synonymy, Cancer

medusarum O. F. Mtiller, 1776. In his original

diagnosis of Hyperia, Latreille (1823) included a

single species, H. sueurii Latreille, n. sp., based on

his figures of Phronima published a few years

earlier (Latreille, 1818; pi. 328: figs. 17-18). These

figures were copies of much earlier illustrations by

St^rm (1762, pi. 1: figs. 12-13) of his nonbinominal

species "Pulex cancriformis antennis brevissimus,

corpore latiore." Since Miiller's Cancer medusarum

is also based entirely on St0rm's animal, it is a

senior objective synonym of Hyperia sueurii and

must be cited as the type-species of Hyperia.

Species of Hyperia spend at least part of their

lives in association with scyphomedusans (Hollowday, 1947; Dahl, 1959a, 1959b; Bowman, Meyers,

and Hicks 1963; Metz, 1967). They inhabit cool

water, living either at high latitudes or at depths

where cool water occurs in low latitudes.

The status of several of the nominal species of

Hyperia (as restricted above) has remained unsettled right up to the present. There has been

general agreement that H. latreillei Milne Edwards

and H. hystrix Bovallius are junior synonyms of

H. galba Montagu and H. medusarum (Miiller)

respectively, but beyond this opinions have varied

widely. H. gaudichaudii Milne Edwards has been

considered a junior synonym of H. galba by some

authors (Walker, 1904, 1907; Stephensen, 1924;

Barnard, 1930, 1932; Vinogradov, 1956); others

have recognized it as a distinct species (Stebbing,

1888; Bovallius, 1889; Walker, 1903; Chilton, 1912;

Barnard, 1916). H. spinigera has been synonymized

with H. galba (Stephensen, 1924; Reid, 1955;

Hurley, 1960a), and H. antarctica with H. galba

(Vinogradov, 1956) and H. spinigera (Barnard,

1932). Both are recognizable species, in my opinion.

Key to the Species of Hyperia

1. Posterior margin of s5-6 of P5-4 completely unarmed, even when viewed with high

magnification .5. H. antarctica

Posterior margin of s5-6 of P3-4 armed with setae which may be very short or as long as

width of segment and which may be very closely shaped or widely scattered 2

2. Setae on posterior margins of s5-6 of PS—4 uniformly short and closely spaced. Posterior

margin of pleonite 3 very convex 6. H. spinigera

Setae on posterior margins of s5-6 of P3-4 longer and not uniform in length, less numerous

and more scattered, sometimes missing from s6 S

3. UpS protopod less than half as wide as long 7. H. leptura

Up3 protopod half or more than half as wide as long 4

4. Coxa of P4 pointed and projecting laterad in adult. Limited to Antarctic (including South

Georgia) 4. H. macrocephala

Coxa of P4 not pointed or projecting laterad in adult. Not limited to Antarctic 5

5. UpS endopod distinctly more than half as wide as long 8. H. crassa

UpS endopod half or less than half as wide as long 6

6. Posterior margin of s6 of Pl-2 not overlapped by spines 2. H. galba

Posterior margin of s6 of Pl-2 overlapped by spines 7

7. Spines on distal part of s6 of Pl-2 reaching or extending beyond apex of s7

1. H. medusarum, medusarum form

These spines not reaching apex of s7 8

8. Length 10-15 mm. North Pacific 1. H. medusarum, hystrix form

Length more than 20 mm. South Atlantic and Pacific 3. H. gaudichaudii
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1. Hyperia tnedusarum (O.F. Miiller)

FIGURES 2-6

Cancer tnedusarum O. F. Mjiller, 1776:196.

Hyperia tnedusarum (O. F. MiiHer) .-Murdock, 1885:143.-

Bovallius, 1887b: 16; 1889:147-159, pi. 9: figs. 1-21 [literature, synonymy] .-Holmes, 1908:490.-Stephenson, 1923:15-

17, chart 2 [synonymy, distribution]; 1924:80-81.—Dunbar,

1942:37; 1954:782-783; 1963:3 [distributk>n].-Bousfield,

1951; 138-139; I956:144.-Shoemaker, 1955:71-72.-Vinogradov, 1956:210-21 l.-Oldevig, 1959:125.

Hyperia latreillei H. Milne Edwards, 1830:388, pi. 11: figs.

l-7.-Bovallius, 1889:164-175, pi. 9: figs. 31-43, pi. 10: figs.

1-17.

Hyperia hystrix Bovallius, 1889:159-163, pi. 9: figs. 22-30.

Hyperia galba (Montagu).-Brusca. I967a:388; 1967b:542-543

[misidentification].

DERIVATION OF NAME.—Not specifically stated,

but presumably refers to association of species with

scyphomedusae reported by Str0m (1762).

TYPE-LOCALITY.—S0ndm0r, Norway.

DIAGNOSIS.—Head length (lateral view) subequal

to length of pereonites 1-2 combined. Interantennal

lobe prominent (medusarum form) to moderate

(hystrix form). Female Al flagellum falcate; A2

flagellum very slender. Relative length of segments

of Md palp, 1:1.7:2.1; segment 2 gently arched;

segment 3 nearly straight. Outer lobe of Mxp much

longer than inner lobe; seta on anterior surface of

inner lobe long, numerous, and dense. S6 of PI and

P2 with many spines on medial and lateral surfaces,

more numerous and longer in older and larger

specimens. In medusarum form (Figures 4c,d,fg)

some spines reach beyond apex of s7; in hystrix

form (Figures 4a,b,e) spines are shorter and dactyl

is relatively longer. Posterior margins of s6 and s7

serrate; serrations present in medusarum form but

obscured by spines and twisting of segment, hence

not readily apparent and margin appears notched.

S4-s5 of P3-P4 with series of long and short spines

on anterior margin, long spines about equal

in length to width of segment; spines more

numerous in medusarum form (Figure 3a), less

numerous in <?» where they may be limited

to s5. S6 minutely serrate, anterior margin usually

unarmed, but may bear a few spines. P5-P7 subequal in length in 9 ; in tf P5>P6=P7, usually

unarmed except for cluster of spines at anterodistal

corner of s2 of P7. Additional setae occasionally

present, especially in young of medusarum form;

variation in setation shown in Figure 3. Posterolateral corners of pleonal epimera ending in points,

most pronounced in pleonite 3.

VARIATIONS.—This species, the most common

along the Pacific coast of North America, is rather

variable. Specimens from Pt. Barrow, Alaska, are

about 20 mm in length and closely resemble USNM

specimens from Frenchman's Bay, Maine. This is

the typical H. medusarum described by Bovallius

and Sars which occurs along the east coast of North

America as far south as Connecticut and which is

commonly found on scyphomedusae in Alaskan

and Canadian Pacific waters. (Bowman, Meyers,

and Hicks, 1963). Adults collected at Friday

Harbor, Washington, in may cases from medusae,

resemble those from Pt. Barrow, but are smaller

and may have shorter spines on the gnathopods.

Specimens taken in plankton nets and midwater

trawls between the latitudes of Vancouver Island

and San Diego are smaller (9-15 mm), although

fully mature, and the setal armature of the

pereopods is less well developed. These specimens

may belong to the form given specific status

(Hyperia hystrix) by Bovallius (1889) and reduced

to a synonym of H. medusarum by Stephensen

(1924) after examining Bovallius' syntypes. After

studying carefully the material in the Smithsonian

Institution I am inclined to accept Stephensen's

position, at least for the present. Although the

majority of North Pacific specimens can be assigned

with reasonable confidence to either H. medusarum

(sensu stricto) or H. hystrix, the presence of seemingly intermediate specimens at Friday Harbor

suggests the possibility of intergradation between

the two forms. A definitive study based on more

material than is available to me now is needed to

resolve the question.

DISTRIBUTION.—The hystrix form was taken in

limited numbers on the CalCOFI cruises, and

Figure 6 shows its combined occurrences on Cruises

1, 5, and 9. Most of the positive stations were north

of Point Conception, reflecting an affinity for cooler

water. The medusarum form was not found in the

CalCOFI samples, perhaps because of its closer

association with scyphomedusae in coastal waters,

or because it is limited to cooler water than that in

the area of the CalCOFI cruises.

The distribution of H. medusarum in the Atlantic Ocean will be considered with the discussion of

the distribution of H. galba.
NUMBER 1S6

FIGURE 2.—Hyperia medusarum (a-g, $) : a, hystrix form, Puget Sound, Washington; b, head,

anterior, hystrix form, North Pacific (4O°34'N, 147°55'W); c, head, medusarum form, Pt. Barrow,

Alaska; d, pleon, hystrix form, off San Francisco; e, pleon, medusarum form, Pt. Barrow; f-g,

Al-2, hystrix form, off San Francisco; h, Md, $ hystrix form, off San Francisco; i, Mx2 outer

lobe, 9 hystrix form, off San Francisco; /, Mxp, lateral, 9 medusarum form, Pt. Barrow; k, outer

lobe of same, posterior surface; 1-m, urosomes of $ and 9 , Pt. Barrow; n-o, urosomes of

3 and 9 , off San Francisco.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE S.—Hyperia medusarum: a, P3, medusarum form 9 , Pt. Barrow, Alaska; fc, PS, hystrix

form $ , off San Francisco; c, P3, medusarum form $ , Puget Sound, Washington; d, P3,

medusarum form £ , Friday Harbor, Washington; e, P4, hystrix form 9 , off San Francisco; /, P4,

medusarum form <$, Friday Harbor; g, P4, medial surface, medusarum form $ , Kuleet Bay,

Vancouver Island; h-j, P5-7, hystrix form 9 , off San Francisco; k-m, P5-7, hystrix form 9 ,

Monterey Bay, California; n-p, P5-7, medusarum form, 9 , Pt. Barrow.
NUMBER 1S6

FIGURE A.—Hyperia medusarum 9: a-b, Pl-2, hystrix form, off San Francisco; c-d, Pl-2 medusarum form, Pt. Barrow, Alaska; e, PI, hystrix form, Monterey Bay, California; /, PI, medusarum

form, Friday Harbor, Washington; g, PI, medusarum form, Frenchman's Bay, Maine.

FIGURE b.—Hyperia medusarum, P7: a, 9, 16.5 mm, Monterey Bay, California; b, $, 21 mm,

Bering Sea; c, juvenile, locality unknown; (d-j, from Auke Bay, Alaska) d, $ 14 mm; e,

$ 10.5 mm; /, $ 9.5 mm; g, 9 10.0 mm; h, 9 7.5 mm; i, 9 5.0 mm; /, 9 3.7 mm.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Distribution of Hyperia medusarum, hystrix form,

CalCOFI Cruises 1, 5, and 9 combined. Dashed line shows

approximate limits of area sampled during cruises.

2. Hyperia galba (Montagu)

FIGURES 7, 15p

Cancer gammarus galba Montagu, 1813:4, pi. 2; fig. 2.

Hyperia galba (Montagu) -Guerin, 1825:771.-Bovallius,

1889:180-188, pi. 10: figs. 25-32 [literature, synonyray].-

Stephensen, 1923:17-19, chart 3 [distribution]; 1924:81-83,

chart 11; 1944:9.-Shoemaker, 1920:24E; 1926:3; 1955:71.-

Chevreux and Fage, 1925:401-402, fig. 401 .-Behning, 1939:

354.-Dunbar, 1942:37; 1954:783 [distribution]; 1963:3 [distribution].-Schellenberg, 1942:241-242, fig. 202.-Hollowday, 1946:passim [association with medusae].—Bousfield,

1951:138; 1956:144.-Certain, 1953 [chromosomes]; 1960

[chromosomes].-Alvarado, 1955:219-220.-el Maghraby and

Perkins, 1956:494.-Vinogradov, 1956:210.-Dahl, 1959a;

1959b [evidence for ectoparasitism].-Oldevig, 1959:125.-

Bowman. Meyers, and Hicks, 1963:140-146, figs. 1, 2e-h.—

Bulycheva, 1964:317-318.-Metz, 1967 [association with medusae].

DERIVATION OF NAME.-Presumably from the

Latin "galbus" [=yellow], referring to color of the

animal.

TvPE-LocALiTY.-South coast of Devonshire,

England.

DIAGNOSIS.—Head shorter than pereonites 1-2

combined; gland cone rather pointed. Posterolateral

corner of pleonal epimeron 3 ending in small

point; posterior margin moderately convex. Outer

lobe of Mxp with a few short setae along inner

margin; inner lobe about y4 as long as outer lobe.

Surfaces of s6 and s7 of P1-P2 with relatively few

setae, those of s6 not extending past serrate

posterior margin of base of dactyl. S5 of P3-P4

with a few short setae; posterior margin of s6 finely

serrate, without setae. S2 of P5-P7 rather narrow;

P5 without setae, P6-P7 with cluster of setae at

distal corner of s2. Length of adult o* and ° ,

10-24 mm.

DISTRIBUTION.-AS Bulycheva (1964) has pointed

out, Hyperia galba is limited to the Arctic and the

cooler regions of the northern Atlantic and Pacific

Oceans. It has been reported from the South

Atlantic, Antarctic, and Indian Oceans, but none

of these records is supported by illustrations or

descriptions. Some have resulted from the authors

considering H. spinigera, a valid and distinct

species, to be a junior synonym of H. galba. I list

below the published records of Hyperia galba that

I consider unacceptable. Bulycheva (1964) also

considers the Antarctic and Indian Ocean records

of H. galba to be erroneous.

Doubtful and Erroneous Records of Hyperia galba

Author

Caiman, 1898

Walker, 1904

Stewart, 1913

Spandl, 1924

Spandl, 1927

Barnard, 1930

Barnard, 1932

Ruffo, 1949

Reid, 1955

Hurley, 1960a

Siegfried, 1963

Brusca, 1967a,b

White and Bone, 1972

Locality

Puget Sound

Gulf of Manaar, Ceylon

off Madeira

northern Red Sea

north of Cape Verde Is.

west of Falkland Is.

Melbourne Habor, Australia

west of Falkland Is.

.South Africa

Chile?, Argentine?, Antarctic

(locality uncertain)

Gulf of Guinea

Indian sector of Antarctic

Atlantic coast of southern Africa

southern California

South Orkney Is.

H. galba has a circumpolar distribution in the

Arctic Ocean (Bulycheva, 1964). In the western

Atlantic it extends south to Chesapeake Bay (Bowman, Myers, and Hicks, 1963), and in the eastern

Atlantic it reaches the coasts of England, France,
NUMBER 136

11

FIGURE 7.—Hyperia galba (a-h, $ from Newfoundland): a, anterior end, lateral; b-c, Mxl outer

lobe of adult and 4.9 mm juvenile; d, Mxl inner lobe; e, Mx2, adult, setae omitted; /, Mx2

outer lobe, juvenile; g, Mxp lateral; h, Mxp outer lobe, posterior; (i-j, pleonal epimera) i, $

from Nova Scotia; ;, $ from Long Island Sound; (k-s, from Ireland) A, coxae 1-7, 9 ; ', $ PI,

medial; m, $ P3; n-r, $ PS-7; 5, telson and Up3.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and Spain (Stephensen, 1923, 1924; Alvarado, 1955).

It is said to have been collected from numerous

localities in the areas surrounding the Azores by

Prince Albert Ier of Monaco (Chevreux, 1900,

1935; Pirlot, 1939); these identifications require

confirmation, since they are at or beyond the

southern boundary for this species. It is the only

hyperiid occurring in the Baltic. It is apparently

absent from the Mediterranean (Stephensen, 1924),

although some authors (e.g., Chevreux and Fage,

1925; Spandl, 1927; Pirlot, 1929) include the

Mediterranean within its range.

Records of H. galba from the Pacific are much

fewer than those from the Atlantic. Derjavin (1927)

recorded it from Avachinska Inlet (at Petropavlovsk) on the Kamchatka Peninsula. Behning (1939)

reported it from the Bering, Okhotsk, and Japan

Seas, without giving specific localities. Buycheva

(1964) considers the Japan Sea record erroneous.

Vinogradov (1956) found it in the western Bering

Sea, the Gulf of Anadir, the Kamchatka region, and

the waters of the Kurile-Kamchatka Trench. Yoo

(1971) found it off Japan north of 37°50'N. None

of these western Pacific records is supported by

descriptions or illustrations.

In the eastern Pacific H. galba was reported from

Puget Sound by Caiman (1898), and from southern

California by Brusca (1967a, 1967b), but it is

likely that these authors actually had the hystrix

form of H. medusarum. The USNM has specimens of H. galba from four localities near the

Alaska Peninsula: Kodiak; Unalaska; St. George I.,

Pribilof Is.; and in the Bering Sea east of the

Pribilof Is. (Albatross sta. 3540, 56°27'N, lee^'W).

I have not seen specimens of H. galba from elsewhere in the Pacific in spite of examing numerous

specimens of Hyperia collected by towing nets and

trawls and by removing them from medusae. Apparently it does not occur off the Pacific coast of

Canada or the contiguous United States.

Compared with H. medusarum, H. galba is

abundant in the Atlantic and scarce in the Pacific.

The distribution of the two species in the Atlantic

is considered below.

Comparison of Atlantic Distribution of

Hyperia medusarum and H. galba

Both H. galba and H. medusarum are ArcticSubarctic-Boreal species associated with scyphomedusae. Is there competition between the two

species or is this avoided by niche differentiation?

So little is known of the habits and ecological

requirements of the two species that I can only

summarize what I have gleaned from published

works.

H. galba seems to be by far the more abundant

species in the Atlantic. Stephensen (1924) reports

that the Thor obtained 615 specimens of H. galba

from 40 stations, mostly southwest of Ireland, and

only 2 specimens of H. medusarum from 2 stations.

The Ingolf (Stephensen, 1923) collected about 20

specimens of H. galba from 6 stations, mostly north

of 60°N, but failed to collect H. medusarum. From

plankton collections in Ungava Bay, Dunbar (1954)

reported 12 specimens of H. galba from 4 stations

and 2 specimens of H. medusarum from 2 stations.

Collections in Belle Isle Strait, reported on by

Bousfield (1951), yielded IS specimens of H. galba

from 4 stations and only 1 specimen of H. medusarum. Collecting from Cyanea capillata, Hicks

obtained 120 H. galba and 1 H. medusarum in

Narragansett Bay, and 57 H. galba and 3 H. medusarum in the Niantic River, Connecticut (Bowman,

Meyers, and Hicks, 1963).

The host preferences of the two species are not

known. Both have been found associated with

Amelia, Chrysaora, Cyanea, and Rhizostoma. In

addition, H. medusarum has been taken from

Thaumantias (Stephensen, 1923), and young individuals of H. galba may be found on the hydromedusan Melicertidium octocostatum (Schellenberg, 1942). Occasionally H. galba may be found

on the ctenophore, Beroe (Stephensen, 1923; Schellenberg, 1942). Dales (1966) states that H. medusarum is associated with Rhizostoma pulmo, whereas H. galba occurs with Rhizostoma octopus and

Chrysaora hyoscella. Dales does not document this

statement nor say whether it is his own observation.

Experimental studies of host preference might aid

us to understand the differences between these

two species of Hyperia.

Comparing the occurrence of the two species off

the west coast of Ireland, Tattersall (1906) suggested that H. medusarum was a more distinctly

oceanic form than H. galba. According to Schellenberg, H. galba penetrates farther into the Baltic,

reaching Warnemiind, whereas H. medusarum only

reaches the Great Belt (Store Baelt). Metz (1967)

reports that H. galba is abundant in the Isefjord
NUMBER 136 13

Area of Denmark (S 17.6-21.8%c), but does not

mention H. medusarum.

H. galba extends farther south in the Atlantic

than H. medusarum, reaching at least to the latitude of the coast of Spain (Alvarado, 1955). According to Schellenberg (1942) H. medusarum does

not penetrate the inner North Sea, and its absence

from Chevreux and Fage's (1925) volume on the

amphipods in the Faune de France series indicates

that it does not occur on the Atlantic coast of

France. In the eastern Atlantic the presence of H.

galba and the absence of H. medusarum from

Cheasapeake Bay (Bowman, Meyers, and Hicks,

1963) also suggests a tolerance for higher temperature in H. galba.

3. Hyperia gaudichaudii H. Milne Edwards

FIGURES 8-10

Hyperia gaudichaudii H. Milne Edwards, 1840:77.—Nicolet,

1849:245.-Bovallius, 1887b: 16; 1889:175-179; pi. 10: figs. 18-

24.-Stcbbing, 1888:1394-1398. pi. 169; 1914:374.-Walker,

1903:40; 1907:7.-Chilton, 1912:513.-Barnard, 1916:25-

286.-Dick, 1970:55-56.

Lestrigonus gaudichaudii.—Bile, 1862:289, pi. 48: fig. 3.

Hyperia galba (Montagu) -Barnard, 1930:411-412; 1932:

273.-Siegfried, 1963:8.-Hurley, 1969:33.

DERIVATION OF NAME.—Presumably after the

French botanist Charles Gaudichaud-Beaupre", 1789—

1854, who made important collections during his

voyages on the Uranis, L'Herminie, and La Bonite.

DIAGNOSIS.—Very similar to H. medusarum, hystrix form, but generally larger (10-14 mm). Confined to the southern hemisphere in antiboreal and

perhaps Antarctic zones.

RELATIONSHIPS.—Illustrations are given here of

specimens from the west coast of southern Africa

reported as H. galba by Siegfried (1963), sent to

me by R. I. Dick, University of Cape Town. A few

figures are also given of a female and a male from

Challenger station 312 in the Straits of Magellan,

loaned to me by R. W. Ingle, British Museum

(Natural History). The similarity to the hysterix

form of H. medusarum is obvious, and maintaining

H. gaudichaudii as a separate species is not easily

defended. My reasons for doing so are largely

zoogeographical. Whereas hystrix has a boreal distribution, that of gaudichaudii is antiboreal, and

there is no possibility of gene flow between them.

Biantitropical species among epiplanktonic Crustacea are rare and are becoming rarer as their

taxonomy is examined more closely. Of the few

biantitropical hyperiids the best known example

is Parathemisto gaudichaudii, whose distribution is

reviewed in detail by Kane (1966). The vast majority of coldwater hyperiids, however, are confined

to one hemisphere; consequently I prefer to give

full specific status to Hyperia gaudichaudii rather

than to maintain that H. medusarum is a biantitropical species.

DISTRIBUTION.—//, gaudichaudii is known to occur

along the southern coasts of Africa, Australia, and

South America (Figure 10). Walker's two records

from the Ross Sea (1903, 1907) seem questionable

since they are the only reported occurrences south

of the Antarctic Convergence. I suspect that

Walker's specimen's were immature Hyperia macrocephala in which the coxae of P4 had not yet

developed the pointed and splayed form so characteristic of the adult of that species. Chilton's (1912)

record from Scotia station 541 (37°41'N, 29°25'W)

is also unacceptable; it lies well outside the range

of H. gaudichaudii but within that of H. galba and

may well refer to the latter species.

4. Hyperia macrocephala (Dana)

FIGURES 11, 12

Tauria macrocephala Dana, 1853:988-989.—Bovallius, 1885:

16-17; 1887b: 19; 1889:81-82, figs. 1-4 [from Dana].-Chevreux, 1913:86 [in list].-Shoemaker, 1914:76.-Spandl, 1927:

156-158, fig. Sag.

Taura macrocephala.—-Dana, 1855; pi. 68: fig. 2 [typographical error].

Hyperia macrocephala (Dana) .-Bate, 1862:296, pi. 49: fig. 2

[from Dana].—Shoemaker, 1945a:291-293, fig. 2A-B.—Emison, 1968:202, fig. 11.

Hyperia galba (Montagu) .—White and Bone, 1972: passim.

DERIVATION OF NAME.—From the Latin, meaning

"large head."

TYPE-LOCALITY.—Antarctic Ocean, off Oates Coast

(ca. 66°S, 157°E).

DIAGNOSIS.—Head about as long as pereonites 1-2

combined; gland cone rather short. Coxa of P4

of large mature specimens pointed, projecting

laterally. Outer lobe of Mxp armed with short

setae only; inner lobe shorter than outer, with

dense covering of setae on anterior surface and 2

terminal spines. Pl-2 with very spinose distal segments. P3-4 with numerous short setae on s5-6;
14

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8.—Hyperia gaudichaudii, $ from west coast of southern Africa: a, A2, lateral; b, labrum;

c, incisor of right Md; d, incisor and lacinia of left Md; e, labium; /, Mxl outer lobe; g, Mxl

inner lobe; h, Mx2 inner lobe; i, Mx2 outer lobe; /, Mxp outer lobe; k, PI medial; /, P2 medial;

m, P3; n-p, P5-7; q, telson and Up3.
FIGURE 9.—Hyperia gaudichaudii, from Challenger station 312, Straits of Magellan (a-f, $): a,

head, lateral; b, pleon, lateral; c, PI, lateral; d, PI coxa; e, P4 coxa; /, P7 coxa; (g-q, 9) g,

anterior end, lateral; h, head, anterior; i, ventral part of head, lateral; ;, P2, lateral; k-q, coxae

of Pl-7.

FICURE 10.—Distribution records for Hyperia gaudichaudii. Stippling indicates areas where exact

localities were not given. Walker's Ross Sea records, considered questionable, are indicated by

open circles.
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 11.—Hyperia tnacrocephala: a, head and pereon, lateral, young $ (drawing by C. R.

Shoemaker, 1945b, fig. 2A); b, 9 head and pereon, dorsal; c, 9 head; d, 9 pleon; e, 9 labrum; /, incisor of 9 right Md; g, incisor of 9 left Md; h, $ lacinia mobilis; i, $ Mxl outer

lobe; ;, 9 Mxl inner lobe; k, 9 Mx2; /, 9 Mxp outer lobes; m, PI medial, young $; n, P2

medial, young $; o, P3 medial, young $; p, 9 urosome; q, $ telson and Up3.
NUMBER 136 17

FIGURE 12.—Hyperia macrocephala, distribution records (circles). Hyperia antarctica Spandl,

type-locality (triangle).

anterior margin of s6 and distal part of anterior

margin of s5 minutely serrate. P5-7 naked except

for cluster of setae at anterodistal corner of s2 of

P6-7. Pleonites with sharply pointed posterolateral

corners; posterior margin of epimera of pleonite 3

rather strongly convex. Uropods rather slender.

Telson as long as width at base (cf) or slightly

shorter (9), about 0.4 as long as peduncle of Up3.

Length up to 29 mm.

RELATIONSHIPS.—Mature individuals of this distinctive species, the largest of the genus, are easily

recognized by the winglike coxa 4. Immature specimens have normal coxae. Thus far only 8 specimens

have been reported, 7 from 3 localities adjacent

to Antarctica and 1 from South Georgia. With the

exception of Spandl's (1927) specimen, from the

stomach of an AdeTie penquin, all were collected

from medusae.

DISTRIBUTION.—As a result of the increased biological investigations in the Antarctic in recent

years, additional specimens of H. macrocephala

have come to the Smithsonian Institution, all from

the McMurdo Sound area of the Ross Sea. William

Emison contributed 12 specimens collected from

the stomachs of Ad£lie penguins (Emison, 1968).

Carleton Ray donated two large collections from

medusae, one containing 59 females and 42 males,

the other consisting of 28 females and 21 males.

The first collection also contained a female Hyperiella dilatate Stebbing.

The known distribution of H. macrocephala

(Figure 12) shows that it occurs around the entire
18

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Antarctic continent. Thus far the only record outside of Antarctic coastal waters is one from South

Georgia (Shoemaker, 1914). From the large number,

101, found on a single medusa by Ray, it appears

that H. macrocephala may be rather abundant in

Antarctic waters. Curiously, it is not included by

Hurley (1969) in his distribution maps of Antarctic

hyperiids, although it, rather than H. antarctica,

despite the latter's specific name, appears to be the

most common species of Hyperia in Antarctic

waters. H. antarctica lives at greater depths and

unlike H. macrocephala is not restricted to the

Antarctic but has a nearly worldwide distribution.

In my opinion, Antarctic records of species of

Hyperia other than H. macrocephala and H. antarctica are highly questionable.

5. Hyperia antarctica Spandl

FIGURES 12-14

Hyperia antarctica Spandl, 1927:153-155, fig. 2a-g.—Hurley,

1969:33.

Hyperia spinigera Bovallius.—Shoemaker, 1945:238 [females

only; misidentification].

DERIVATION OF NAME.—Not stated, but obviously

refers to type-locality.

TYPE-LOCALITY.—Antarctic Ocean, off Leopold

and Astrid Coast (68°18'S, 80°27'E).

DIAGNOSIS.—Female. General appearance of body

more narrow and elongate than in $ of other

species of Hyperia, and more nearly resembles c?

of these species. Pereonites relatively long. Posterior

corners of pleonite 1 rounded, of pleonites 2-3

with small points. Al flagellum digitiform, in contrast with usual slender shape in Hyperia, with

cluster of apical setae and moderate number of

esthetes on concave medial surface. Md with shorter

marginal setae on molar than in H. medusarum;

teeth of incisor limited to molar half; segments of

palp subequal, distal segment markedly falcate.

Inner margin of outer lobe of Mxp with a number

of short setae and a long seta inserted at about

y3 the distance from base; inner lobe longer than

outer lobe, strongly recurved, with 2 robust spines

at apex. Posterior parts of distal margins of s2-4

of Pl-2 with long setae; distal margin of s6 of PI

rounded, not produced into carpal process, with

very long setae, some nearly as long as s6; posterior

margin of s6 with long setae, notched where setae

are inserted, but not serrate. S5 of P2 with very

short setose carpal process, medial setae very long;

posterior margin of S6 notched but not serrate,

with long setae. P3-4 similar, unarmed except P3

has cluster of setae at posterior distal corner of s2

and 1-2 setae at posterior distal corner of s3. P5-7

robust; unarmed except for cluster of setae at anterior distal corner of s2 in P7; in length P7>P6=

P5. Uropods very similar to those of H. medusarum.

Telson slightly longer than width at base, slightly

more than half as long as peduncle of Up3. Length

of adult 12-28 mm.

MALE.—The male of H. antarctica has not been

reported previously. The immature male from off

Vancouver Island is very similar to the female.

Except for Al and A2 there is no obvious difference

in the male at this stage of maturity.

RELATIONSHIPS.—K. Barnard (1932) considered

H. antarctica to be identical with H. spinigera,

but there are many significant differences between

the two species: e.g., the proportions of the Md

palp segments, the length of the Mxp inner lobe,

the armature of Pl-4, and the relative width of

the Up3 inner ramus.

Because of the incompleteness of Spandl's account (he described and illustrated only the pereopods) I was at first uncertain of the correctness of

my identification of H. antarctica. Fortunately I

was able to compare my specimens with the holotype, borrowed from the Zoologisches Museum der

Humboldt Universitat, Berlin, through the kindness of Dr. H.-E. Gruner, and to confirm their

identity. Spandl gave the length of the holotype

as about 10 mm, but this measurement was apparently only approximate and made with the

pleon bent under the pereon. With the pleon

unfolded and held in a naturally curved position

the holotype measures about 24 mm. It is a fully

mature ? with well-developed oostegites.

Of the specimens recorded as Hyperia spinigera

from the Bermuda area by Shoemaker (1945b),

three are in the Smithsonian Institution. One of

these, a 14.5 mm °. with fully developed oostegites,

collected in a net towed at 1463 m, is a typical

specimen of H. antarctica.

DISTRIBUTION.—The only published record for

H. antarctica is the original one of Spandl (1927)

from a 200 m vertical tow in the Indian Ocean

sector of the Antarctic (68°18'S, 8O°27'E; see Figure
NUMBER 136 19

FIGURE 1$.—Hyperta antarctica (a-b, $): a, anterior end, lateral; b, urosome; (c-q, $) c, lateral

view; d, pleon lateral; t, Al lateral; /, incisor of right Md; g, Md palp; h, Mxl; i, Mx2; j , Mxp,

posterior; k, Mxp, lateral; I, PI; m, P2; n, P3; o, P5; p, P6; q, P7.
20

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 14.—Hyperia antarctica, distribution records. The type-locality is shown on Figure 12.

12). I have found it in samples from 14 additional

widely scattered localities shown in Figure 14. The

depths of sampling ranged from 200 m to about

2000 m with most of the tows being made at 400-

600 m. Even from such limited data it is evident

that H. antarctica has a very wide distribution at

mesopelagic depths. Presumably it is associated

with medusae, like other species of Hyperia, but

there is no hint at present as to what medusae may

be involved.

6. Hyperia spinigera Bovallius

FIGURES 15-16

Hyperia spinigera Bovallius, 1889:191-194, pi. 10: figs. 33-

39.-Vosseler, 1901:58.-Tattersall, l906:22.-Barnard, 1932:

273-274, fig. 160-Thorsteinson, 1941:87-88, pi. 8: figs.

79-82.-Dunbar, 1942:37 [spingera!]; 1963:3.-Shoemaker,

1945a:238, text-fig. 35A-D[£ specimens only].-Hurley,

1955:140-143, figs. 83-95; [?] 1965:15.-Oldevig, 1959:125.-

Brusca, 1967a:388; 1967b:452.

Hyperia galba (Montagu) .—Norman, 1900:128 [partim].—

Stephensen, 1924:81-83 [partim].-Barnard, 1930:412.-Hurley, 1960a: 111-112; 1969:33.

DERIVATION OF NAME.—Not given, presumably

refers to the armature of the gnathopods.

TYPE-LOCALITY.—Spitzbergen; off northern coast

of Norway; off south coast of England.

DIAGNOSIS.—Body moderately robust, length up to

21 mm. Posterior margin of pleonal epimeron 3

strongly convex. Segments of Md palp subequal;

incisor with about 15 teeth. Outer lobe of Mxp

longer than inner lobe; setae on inner and outer

lobes shorter and sparser than in H. medusarum.

Setae of PI and P2 short compared with those of

other species of Hyperia. P3-4, posterior margins

of s4, s5, and s6 densely covered with short slender

spinules. S2 of P5-7 broad, with group of spines at

anterodistal corner in P7; anterior margins of

s4-5 with dense covering of short spinules in P5,

bare in P6-7. Inner ramus of Up2 and Up3 broad.

RELATIONSHIPS.—The collections of the Copenhagen Museum contain 6 vials labeled "Hyperia

spinosa" by Bovallius and stated by Stephensen

(1924) to be syntypes of H. spinigera. Stephensen

considered these syntypes to be aged specimens of

H. galba, and therefore placed H. spinigera in

synonymy with H. galba.

These same 6 vials were kindly loaned to me by

Dr. Torben Wolff. The 17 mm male from the north

side of Iceland labeled "Borch ded. 1859" is a
NUMBER 136 21

FIGURE 15.—Hyperia spinigera, $: a, lateral view; b, incisor of right Md; c, Md palp; d, Mxl,

inner lobe; e, Mxp, oblique view; /, Mxp, posterior; g, PI; h, P2; i, P3; /', P5; k, P6; /, urosome;

(m-o, pleonal epimera of specimens in Copenhagen Museum labeled "Hyperia spinosa" by

Bovallius) m, "61 °N, 20°W, Olrich ded. 1867"; n, "18°S, 2°W, Andrea ded. 1864"; o, "North

side of Iceland, Borch ded. 1859" [=.H. medusarum, hystrix form]. Hyperia galba, $ : p, s6 of P3.
22

specimen of Hyperia medusarum, hystrix form; the

other specimens are clearly distinct from H. galba,

and as pointed out by Stephensen must constitute

at least part of the type-series of H. spinigera,

although, curiously, none are from localities listed

by Bovallius in his original account of the species.

The specimens of the type-series, as well as the

other specimens that I have examined, exhibit very

little variation. Characters useful in distinguishing

them from H. galba are the following:

1. Some of the spines on s6 of PI and P2 extend

across the posterior margin in H. spinigera; this

margin is devoid of spines in H. galba.

2. The posterior margins of s5 and s6 of PS and

P4 have longer spinules in H. spinigera (compare

Figures 15» and 15p). S5 usually has several spines

in H. galba, none in H. spinigera.

3. The epimera of pleonite 3 are more strongly

produced posteriorly in H. spinigera than in H.

galba (compare Figures \ba,m,n and li,j). Although

Tattersall (1906) stated that the posterolateral

corners of pleonite 3 are sharply pointed in H.

galba and rounded in H. spinigera, Shoemaker

(1945b) showed clearly that the corners are pointed

in H. spinigera. The difference between the epimera

of the two species is not in the development of the

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

points but in the more strongly convex posterior

margin in H. spinigera, shown by Shoemaker

(1945b, fig. 35c) and Hurley (1955, fig. 94).

The female of H. spinigera has not been described. Several authors (Norman, 1900; Barnard,

1932; Shoemaker, 1945b; Hurley, 1956) have recorded females, unsupported by descriptions or

illustrations, except for Barnard's figure of the

urosome. None of these records can be confirmed.

Norman (1900) considered H. spinigera a. synonym of H. galba, but stated, "Should other authors

disagree with my views in this matter, the [2] female

specimens of H. galba taken by me at Birturbuy

Bay and the one taken off Valentia [both localities

off Ireland] by the 'Porcupine' are, from the character of the gnathopods, to be referred to H.

spinigera." I have examined Norman's 3 females,

loaned to me by the British Museum (Natural

History) through the courtesy of R. W. Ingle; in

my opinion they are typical H. galba.

Barnard's female from Discovery station 298 in

the North Atlantic (13o01'N, 21°34'W) was also

loaned to me by the British Museum (Natural

History). It is identical with specimens from the

Gulf of Guinea assigned in this paper to a new

species, Hyperia crassa. My reasons for proposing

FIGURE 16.—Hyperia spinigera, distribution records,

records verified by examination of specimens Q unverified records from the literature
NUMBER 136 23

a new species for these specimens rather than placing them in H. spinigera are discussed under H.

crassa.

I have examined Shoemaker's female specimens

from Bermuda, which are deposited in the Smithsonian Institution; they are typical specimens of

H. antarctica.

DISTRIBUTION.—Figure 16 shows the occurrences

that I have compiled from the literature and from

specimens in the Smithsonian Institution. H. spinigera is obviously a widespread species and the

present compilation gives only a fragmentary

picture of its distribution. Most of the collections

were made at considerable depths.

7. Hyperia leptura, new species

FIGURE 17

DERIVATION OF NAME.—From the Greek "leptos"

[=slender] + "oura" [=tail], referring to the

slender uropods.

TYPE.—Holotype and only specimen, 9 w'

1

^

embryos in marsupium, USNM 111244, from CalCOFI Cruise 20, station 130.35, off Bahfa Ballenas,

Baja California (26°19.5'N, 113°49/W), in 140-0

m net tow, 14 November 1950.

DIAGNOSIS.—Head longer than pereonites 1—2

combined; gland cone rounded. Outer lobe of Mxp

with several long setae near inner margin; inner

lobe about half as long as outer lobe, anterior

surface heavily setose, apex with single strong spine.

S2 of Pl-2 relatively narrow, about 2.5 times as

long as wide, with a number of short setae on

proximal part of anterior margin; s5-6 long and

narrow, armed with numerous long spines. P3-4

long; posterior margins of s3-6 spinose; posterior

margin of s6 finely serrate. Setae on P5-7 limited

to a few on anterior margins of s2 and s5 (Figure

17/-A). Epimera of pleonite 3 not produced posteriorly. Upl-3 long and slender; rami lanceolate,

those of Upl especially long. Telson slightly shorter

than width at base.

RELATIONSHIPS.—Only one specimen of this

species has been discovered, but its characters are

so distinctive that I have no hesitation in establishing a new species for it. It appears to be closest

to H. macrocephala, but in addition to its much

smaller size, it differs in the form of coxa 4, pleonal

epimera 3, the Mxp, and the narrower peduncle of

Up3.

8. Hyperia crassa, new species

FICURES 18-19

Hyperia spinigera Bovallius.-Barnard, 1932:275-4 [$ only],

fig. 160b.

DERIVATION OF NAME.—The specific name, from

the Latin "crassus" [=thick, fat], refers to the

broad protopods and endopods of the uropods.

TYPES.—All from Ombango Cruise 12 in the

Gulf of Guinea. Holotype $, USNM 137502, station 298 (Grand Schmidt 2), 3°0rS, 9°25'E, 300

meters of wire, 1 May I960, 1040 hrs. Allotype d1.

USNM 137503, station 302 (Grand Schmidt 19),

4°47'S, 10°42'E, 1100 meters of wire, 4 May 1960,

0450 hrs. Paratypes, 3 $, USNM 137504, station 300

(Grand Schmidt 11), 3°48'S, 19°08'E, 200 meters of

wire, 2 May 1960, 1800 hrs.

DIAGNOSIS.—Body rather compact and heavyset,

length 12-15 mm. Head unusually large, longer

than pereonites 1-3 combined, in lateral view much

longer dorsally than ventrally. Pereonites with distinctive dorsal profile shown in Figure 18a; pereonite 2 highest, pereonite 7 concave anteriorly, convex posteriorly; pereonites 2, 3, and 4 with transverse folds. Pleonal epimera with pointed posteroventral corners; posterolateral margin of pleonite

3 only weakly convex. Proximal segment of Md

palp much shorter than distal segments; incisor

with 9 or 10 teeth. Mxp inner lobe long, with long

apical spines. Segments of Pl-2 with well-developed

spines; s2 of PI with very convex anterior margin.

P3-4 with very few spines on posterior margins of

s4-5. P5-7 unarmed, with rather narrow s2. Up2-3

with very broad protopods and endopods.

RELATIONSHIPS.—Barnard (1932) called attention

to the very broad uropods of a female Hyperia

which he identified as H. spinigera. In his discussion he gave the impression that he had some

reservations about this identification, and these

reservations have proved to be entirely justified.

This female, loaned to me by the British Museum

(Natural History), closely resembles the Ombango

females of H. crassa, as can be seen by comparing

Figures 18 and 19. The long head and broad uropods on both Ombango and Discovery females,
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 17.—Hyperia leptura, new species, $ holotype: a, lateral view; b, head, c, pleonal epimera;

d, coxae; e, Mxp, posterior; /, Mxp, lateral; g, PI; h, P2; i, P3; ;, P5; k, P6; /, urosome, dorsal.
NUMBER 1S6 25

FIGURE 18.—Hyperia crassa, new species (as, 9): a» lateral view; b, Al; c, A2; d, incisor of right

Md; e, Mxl inner lobe; /, Mx2 inner lobe; g, Mx2 outer lobe; h, Mxp posterior; i, Mxp lateral;

/, PI; k, PI dactyl, medial; /, P2; m, P3; n, PS, part of s6 and s7, medial; o, P4; p, P5; q, P6,

r, P7; 5, urosome; t, $ head, lateral; u, ^ Md palp.
26

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Hyperia crassa, new species, $ from Discovery

station 298: a, head and pereonites 1-3, lateral; b, pleonite 3,

lateral; c, P2, distal segments; d, P4; e, P5; /, P6; g, P7; h,

urosome, dorsal.

unique features of H. crassa, clearly establish their

specific identity.

DISTRIBUTION.—West coast of Africa, 3-5°S (Ombango stations). SE of Cape Verde Islands, 13°01'N,

21°34'W, Discovery station 298 (Barnard, 1932).

II. Hyperiella Bovallius, 1887b

DIAGNOSIS.—Body small to moderate sized; pereon

rather plump; pereonites all separate. $ Al and

A2 4-merous. Md palp present in 9 > incisor serrate. Mxl, inner lobe with 5 terminal spines. Mx2

with 1 terminal spine on inner lobe, 2-3 on outer

lobe. Mxp, inner lobe well developed, ending in 2

spines; outer lobes separate. PI subchelate; P2

chelate; posterior margin of s6 serrate in both PI

and P2. P3-4, posterior margins of s5 and s6 serrulate, that of s5 bearing a few slender spines. P5

(or P5-6) much longer than P3-4; P7 (or P6-7)

subequal to P3-4. S7 of at least P3-5 very long.

Up3 elongate, except in H. macronyx.

DERIVATION OF NAME.—Not given, presumably

diminutive of Hyperia. Gender feminine.

TYPE-SPECIES.—Designated by Bovallius, 1889,

Hyperiella antarctica Bovallius, 1887b.

REMARKS.—The foregoing diagnosis is modified

from the original by Bovallius in order to accommodate Hyperia macronyx Walker (1906). H. macronyx differs from the two other members of the

genus in that both P5 and P6 are elongate and

Up3 is shorter, but in other respects it is more

similar to Hyperiella antarctica and H. dilatata

than to any of the species of Hyperia as restricted

herein. The alternative to transferring H. macronyx

to Hyperiella would be to propose a monotypic

genus for Walker's species. The latter course seems

less desirable, since it would not call attention to

the similarity of H. macronyx to H. antarctica and

H. dilatata.

Hyperiella is close to Hyperia, but is smaller than

any species of Hyperia, and P5 (or P5-6) are relatively longer. In some respects it resembles Parathemisto; P3-4 approach the prehensile condition

of these legs in Parathemisto, and elongate uropods

and characteristic of Parathemisto.

Key to the Species of Hyperiella

1. P5-6 subequal, longer than other pereopods. Up3 peduncle about twice as long as telson

11. H. macronyx

P5 much longer than P6. Up3 peduncle distinctly more than twice as long as telson 2

2. Anterodistal corners of P6-7 produced into sharp triangular processes. Up3 $, exopod

2/3 or more as wide as endopod 10. H. dilatata

Anterodistal corners of P6-7 produced into blunt processes. Up3 $, exopod less than 2/3

as wide as endopod 9. H. antarctica
NUMBER 136 27

9. Hyperiella antarctica Bovallius

FIGURES 20n, 21g i

Hyperiella antarctica Bovallius, 1887a:20; 1887b:566, pi. 45:

hgs. 72-80; 1889:242-246, text-fig, p. 242, pi. 11: figs. 42-

51.-Stewart, 1913:256.-Barnard, 1930:414; 1932:275, fig. 161

[distribution].-Kane, 1962:301-302.-Vinogradov, 1962:25.—

Hurley, 1969:32, map 5 [distribution].

DERIVATION OF NAME.—Not given; presumably

geographical.

TYPE-LOCALiTY.-Drake Passage, 58°43'S, 76°W.

DIAGNOSIS.—Length 6-8 mm. Posteroventral

corners of pleonal epimera blunter and less prominent than in H. dilatata. P5—7, anterodistal corners

of s2-4 produced into more or less pointed processes, but processes weaker and blunter than in

H. dilatata. Up3 cf, exopod less than

 2/3 as wide

as endopod.

RELATIONSHIPS.—See discussion under H. dilatata.

DISTRIBUTION.—According to Hurley (1969), H.

antarctica inhabits the region between the Subtropical Convergence and the edge of the pack ice.

10. Hyperiella dilatata Stebbing

FIGURES 20a-m, 21a-/

Hyperiella dilatata Stebbing, 1888:1403-1404, pi. I7l.-Bovallius, 1889:247.-Walker, 1907:8.-Spandl, 1927:162-164,

fig. 5a-h.-Barnard, 1930:413-414; 1932:274-275, fig. 161

[distribution]; 1937:4.-Stephenson, 1947:76.—Hurley, 1961:

600; 1969:33, sheet 2, map 5[distribution].—Vinogradov,

1962:25.—Emison, 1968:passim [food of Adelie penguin].

DERIVATION OF NAME.—Refers to the inflated

pereon.

TYPE-LOCALITY.—Indian sector of Antarctic, 63°

30'S, 88°57'E.

DIAGNOSIS.—Length 6-8 mm. Posteroventral

corners of pleonal epimera sharper and more prominent than in H. antarctica. P5-7, anterodistal

corners of s2-4 produced into conspicuous triangular processes. Up3 tf, exopod % or more as

wide as endopod.

RELATIONSHIPS.—The differences that previous

workers have used to distinguish between H. antarctica and H. dilatata are not altogether convincing, but the material at my disposal has not allowed

me to shed much light on the problem. My specimens of H. dilatata, collected from the stomachs

of Ad&ie penguins by William B. Emison, are

somewhat deformed and partly digested, but were

sufficiently intact to permit illustration of most of

the significant details. With regard to H. antarctica,

I have examined only 3 males from the collections

of the Copenhagen Museum, identified by Bovallius

and believed to be syntypes. The original label for

these specimens is lost, and a label in K. Stephensen's handwriting reads (translated from the Danish

by Torben Wolff), "Amongst the 'Icebergs' of the

South Polar Sea. Wessell." In his original description Bovallius (1887b) gave the locality as "Antarctic Seas, around Cape Horn." In 1889 Bovallius

cited the type-locality more precisely: "The American Antarctic region: Lat. 58°43/

; Long. 76°W."

This locality lies in the western part of the Drake

passage, south of the northern limit of drift ice,

hence is compatible with the syntype label.

Accepting the 3 males as syntypes leaves unresolved the question of whether H. antarctica and

H. dilatata are distinct species, since the condition

of the specimens is only fair. A brief history of the

taxonomy of the two species may help to show the

nature of the problem.

The original description of each species was

based on only one sex, the male for H. antarctica

and the female for H. dilatata. Bovallius (1889)

gave a key to the two species that distinguished

them by 3 characters: (1) The pleonal epimera,

with posterolateral corners rounded in H. antarctica

and pointed in H. dilatata. (2) The anterodistal

corners of s2-4 of P5-7, rectangular in H. antarctica

and acutely produced in H. dilatata. (3) The endopod of Up3, "ovate" in H. antarctica, "narrowly

elongate and sharp-pointed" in H. dilatata.

The usefulness of the first character became

doubtful when Stewart (1913) noted pointed pleonal epimera on a cf H. antarctica. This finding

was verified by Barnard (1930, 1932), although he

stated that the points were not as well developed

as in H. dilatata. In the specimens seen by me the

points are somewhat stronger in H. dilatata (compare Figures 20m and 20n), but it is risky to rely

on such a purely quantitative character when nothing is known about its variability. Moreover, the

appearance of the points may be affected by the

action of the preservative; inflation of the epimera,

obscuring the nature of the margins, is frequently

seen in preserved hy peri ids.

That Bovallius' second difference is also a matter

of degree was noted by Barnard (1930, 1932), who
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 20.—Hyperiella dilatata (a-k, ?) : a, Al; b, A2; c-d, incisor and palp of right Md; e-f,

outer and inner lobes of Mxl; g, Mx2 (setae omitted); h-i, posterior and oblique views of Mxp;

j , Mxp, inner lobe (setae omitted); k, PI; /, PI dactyl; m, pleonal epirnera $. Hyperiella

antarctica $ : n, pleon, lateral.
NUMBER 1S6 29

FICURE 21.—Hyperiella dilatata: a, 9 urosome, dorsal; b, $ urosome, dorsal; c, 9 PS, d,

9 P5; e, $ P6; /, $ P7. Hyperiella antarctica: g, Up3; h, P6; i, P7.
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

found pointed anterodistal corners on s2-4 of P5-7

in H. antarctica less prominent than those on H.

dilatata. In specimens seen by me the points are

distinctly more prominent in H. dilatata, but I

cannot rule out the possibility that the points have

been abraded in the H. antarctica syntypes.

The third difference in Bovallius' key concerned

a sexually dimorphic character. The broader

"ovate" endopod of Up3 of H. antarctica was that

of a male Hyperiella, and the "narrowly elongate"

endopod of H. dilatata was an attribute of a female Hyperiella (compare Figures 21a and 216 of

the cT and 9 uropods of my specimens of H.

dilatata). As Barnard (1930) observed, Spandl

(1927) added to the problem by describing the

uropods of an immature male of H. dilatata; uropods of young males are narrow, like those of females. But Barnard himself compounded the

confusion by stating that the endopods of Up2-3

of the male H. antarctica are lanceolate and slightly

narrower than those of the female H. antarctica.

My drawing of the uropods of the o" H. dilatata

(the first for this species) shows the endopod of

Up3 ovate and only slightly narrower than that of

the male H. antarctica. Figures of the uropods of

the female H. antarctica have not yet been published, and I am unable to evaluate Barnard's statement concerning them.

In summary, none of the characters said to be

useful in distinguishing between H. antarctica and

H. dilatata are of proven reliability. Both species

(if they are not conspecific) are in need of detailed

redescriptions, including studies of their variability.

DISTRIBUTION.—From the Antarctic Convergence

to the Antarctic continent (Hurley, 1969).

11. Hyperiella macronyx (Walker), new

combination

FIGURES 22-23

Hyperia macronyx Walker, 1906:452; 1907:7-8, pi. 1: fig. 1.-

Barnard, 1930:412-413.-Emison, 1968:203-206 [food of

Adelie penguin].—Hurley, 1969:33, map 5 [distribution].

DERIVATION OF NAME.—Not given; presumably

from the Greek "macro" [=large] -|- "onyx"

[=claw], referring to the long dactyls on P3-7.

TYPE-LOCALITY.—Two locations in the Ross Sea,

Antarctica: (1) Coulman Island; (2) Discovery winter quarters, on tip of Cape Armitage, south side

of Ross Island, McMurdo Sound.

DIAGNOSIS.—Length, up to 13 mm. Pereon not

so plump as in other species of Hyperiella. Pleonite 3 with point on posterolateral margin. Mx2

with 3 spines at apex of outer lobe. Mxp without

row of spines on distal margin of basal plate. P6

slightly longer than P5, both considerably longer

than P7, which is subequal to P3-4. S7 of P3-7

elongate. Telson slightly longer than wide, about

half as long as peduncle of Up3.

RELATIONSHIPS.—Easily distinguished from its

congeners by the characters given in the key.

DISTRIBUTION.—Known only from the Ross Sea.

III. Hyperioides Chevreux

Hyperioides Chevreux, 1900:143.

Parahyperia Vosseler, 1901:56 [conditional name].

DIAGNOSIS.—Small species with body rather compressed laterally. Head globular, produced' anteriorly above insertion of Al; eyes occupying most of

its surface or limited entirely to dorsal surface.

Pereonites 1-2 fused in both sexes. Coxae fused

with pereonites. Posterior elevation of o* pleonite

1 pronounced. Telson rather short. 9 Al 2-3-

merous. 9 A2 1-merous, moderately long, with

characteristic sinuous shape in lateral view; gland

cone rounded below. Md with dentate incisor;

palp absent in 9- Mx 1 outer lobe with 3 large

terminal spines and a smaller subterminal spine.

Mx 2 outer lobe with 1 terminal and 1 subterminal

spine; inner lobe with 1 terminal spine. Mxp inner

lobe well developed, with 2 terminal spines. PI—2

chelate, with spoon-shaped carpal process bearing

marginal spines. P5-6 distinctly longer than P3-4

and P7; dactyls of P5-7 long, with comb of fine

setules on proximal 3rd of anterior margin. Outer

rami of uropods notched.

DERIVATION OF NAME.—Not given; presumably

from Hyperia -f- the Greek "oides" [=resembling].

TYPE-SPECIES.—By monotypy, Hyperioides longipes Chevreux, 1900. Gender feminine.

REMARKS.—The above diagnosis modifies that of

Chevreux (1900) to include characters which this

study has convinced me are significant at the generic level. Until now Hyperioides has been monotypic, but I am transferring to it a second species,
NUMBER 136 31

FIGURE 22.—Hyperiella macronyx: a, $ lateral; b, $ Al, medial; c, $ A2, lateral; d, left

$ Md; e, Mxl, outer lobe; /, Mxl, inner lobe, apex, setae omitted; g, $ Mx2, setae omitted; h,

Mxp, lateral; i, $ Mxp, outer lobe, posterior; ;, $ Mxp, outer lobe, posterior.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 23.-Hyperiella macronyx: a, $ PI, medial; b, $ P2, medial; c, $ P3, lateral; d, 9 P3,

juncture of s&-7, medial, e, $ P6, lateral; /, $ P7, lateral; g, $ urosome.
NUMBER 136

Hyperia sibaginis Stebbing (1888). This action was

foreshadowed by Vosseler's (1901) misidentification

of H. longipes as H. sibaginis; Vosseler was of

course unaware of Chevreux's description of H.

longipes. Chevreux's diagnosis of Hyperioides accommodates H. sibaginis except for his statement

that the eyes are limited to the upper part of the

head.

Key to the Species of Hyperioides

Eyes limited to upper part of head 12. H. longipes

Eyes occupying most of head surface IS. H. sibaginis

12. Hyperioides longipes Chevreux

FIGURES 24. 25

Hyperioides longipes Chevreux, 1900:145-145, pi. 17: fig. 2;

1913:6; 1935:189-191.-Lo Bianco, 1902:422, 447; 1904:43,

pi. 22: fig. 66.-Walker, 1903:229-230, pi. 19: figs. 7-13.-

Stebbing, 1904:34-36.-Fowler, 1904:49, 53 [table].-Tattersall, 1906:23.-Stewart. 1913:256.-Stephensen, 1924:93-94.-

Chevreux and Fage, 1925:407-408, fig. 405.-Spandl, 1927:

164,-Pirlot, 1929:124-126; 1930:19-20; 1939:37.-Schellenberg, 1927:637, fig. 42.-Barnard, 1930:414-415; 1932:276;

1937:184.-Ruffo, 1938:148.-Shoemaker, 1945b:238.-Bulycheva, 1955:1048 [in table].-Hure, 1955:48.-Hurley, 1956:

15; 1960b:280; 1969:33.-Tregouboff and Rose, 1957:460,

pi. 132.-Kane, 1962:302.- Vinogradov, 1962:25.-Siegfried,

1963:8.-Hure, 1965:48.—Pillai, 1966:211-212, fig. 5.-Vives,

1966:96, table 19; 1968:460, table l.-Dick, 1970:57.-Hure,

Scotto di Carlo, and Basile, 1971:passim.—Yoo, 1971:57.

Hyperia sibaginis ? Stebb.—Vosseler, 1901:60-64, pi. 7: fig.

6-20 [misidentification].

Hyperia sibaginis var. longipes Vosseler, 1901:63 [conditional

name].

Hyperia longipes Vosseler, 1901:63 [conditional name].

Hyperia longipes Chevreux.—Reid, 1955:18 [lapsus].

DERIVATION OF NAME.—Not given; presumably

refers to length of P5-6.

TYPE-LOCALITY.—Eastern Atlantic, 4 Hirondelle

stations west of Bay of Biscay, 44°42'-48°25'N,

9°16'-22°14'W.

DIAGNOSIS.—See below, under H. sibaginis.

DISTRIBUTION.—//, longipes has been recorded

from warm parts of all the world oceans, but little

is known of the details of its distribution. Charts

of its occurrence on 4 CalCOFI cruises show its

consistent presence offshore from San Francisco

south, closer to the coast in the southern part of

the area. This pattern fits in general the eastern

limits of Brinton's (1962) pattern for his "central"

euphausiid species, which occupy ocean gyrals of

midlatitudes 15°-40° in both hemispheres in the

Pacific, but with such limited data I can only suggest this possibility.

13. Hyperioides sibaginis (Stebbing), new

combination

FIGURE 26

Hyperia sibaginis Stebbing, 1888:1379-1382, pi. 165.—Bovallius, 1889:201-203.-Barnard, 1931:127-128.-Chiba, Tsurutu, and Maeda, 1955:194, 196, 200.

Hyperia sibaginis ? Stebbing.—Pirlot, 1930:18-19, fig. 6.

[not] Hyperia sibaginis ? Stebb.—Vosseler, 1901 [= H. longipes Chevreux].

DERIVATION OF NAME.—Not expressly stated, but

presumably for Sibago Island, Philippines.

TYPE-LOCALITY.—Challenger station 200, northeast of Sibago Island, east of Zamboango Peninsula, Philippines: 6°47'N, 122°28'E.

DIAGNOSIS.—Both species of Hyperioides have

been well described and illustrated, and in view of

the ease with which this genus may be identified,

diagnoses of its two species may be presented conveniently by the following tabular comparison.

Hyperioides longipes

Dorsal profile of head

nearly straight

Eye facets limited to dorsal

part of head

Al 3-merous

A2 slightly> 1/2 Al

Epistome distinctly

posterior to gland cone

Carpal process of P2

reaches midlength of s6

Up3 protopod about 2

times as long as wide;

only slightly longer

than exopod

Hyperioides sibaginis

Dorsal profile of head

rounded

Eye facets cover both dorsal

and lateral parts of head

Al 2-merous

A2 nearly = Al

Epistome nearly reaches

anterior margin of gland

cone

Carpal process of P2 reaches

about 125 length of s6

Up3 protopod about 2.8

times as long as wide,

about 1.5 times longer than

exopod

IV. Lestrigonus Milne Edwards

Lestrigonus Milne Edwards, 1830:392; 1840:81-82.

DIAGNOSIS.—Small species, with rather plump

pereon. Head globular; eyes occupying most of its
34

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 24.—Hyperioides longipes, off California (a-g, 9): a, lateral view; b, head, lateral; c,

Al, dorsal; d, A2, dorsal; e, incisor of right Md; /, Mxl, apex of palp; g, Mxl outer lobe; h,

Mx2 (setae omitted); i, Mxp, anterior; /, Mxp, lateral; k, PI; /, P2; m, P4; n, P3; o, P5; p,

P6; ^, P7; r, $ urosomite 1, lateral.
NUMBER 136 35

FIGURE 25.—Distribution of Hyperioides longipes on 4 CalCOFI cruises.
36 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 26.—Hyperioides sibaginis, Golfo Elena, Panama (a-q, 9) : a, lateral view; b, head,

lateral; c, Al; d, A2; e, left Md, incisor and lacinia mobilis; /, right Md, incisor; g, Mxl, outer

lobe; h, Mxl, palp; i, Mxp, anterior surface; /, PI; k, P2; I, P4; m, P5; n, P6; o, P7; p, pleonal

epimera; q, urosome; r, $ urosomite 1, lateral; 5, 5 right Md.
NUMBER 136 37

surface. Some of anterior pereonites fused (2?), 5-5

in 9» 2-4 in cf > always more in 9 . Coxae fused

with pereonites. Telson of moderate size. 9 Al

2-merous. 9 A 2 1-merous, usually very small;

gland cone conspicuous, with pointed or rounded

apex. Epistome prominent, strongly convex anteriorly. Md with dentate incisor; palp absent in 9 .

Mxl outer lobe with 3 large terminal spines and

usually a smaller subterminal spine. Mxp outer

lobes separate, tapering distally; inner lobe usually

well developed. PI subchelate or barely chelate.

P2 distinctly chelate; carpal process spoon-shaped,

with spines along margin of spoon. P5-7 usually

longer than P3-4; P5 and P7 subequal, slightly

shorter than P6.

DERIVATION OF NAME.—"Seemingly Irom Uecldfpv

/WS, an ancient savage tribe of Italy and Sicily"

(Stebbing, 1888). Gender, masculine.

TYPE-SPECIES.—By monotypy, Lestrigonus fabrei

H. Milne Edwards, 1830.

REMARKS.—Next to Hyperia, Lestrigonus is the

oldest available generic name for this group of species. Since Hyperia is restricted herein to a different group of species, Lestrigonus becomes the valid

name of the genus, and L. fabrei is automatically

the type-species. This is not an ideal situation,

since the true identity of L. fabrei is uncertain.

The species called Hyperia fabrei by most authors

is an entirely different species, H. vosseleri Stebbing, which I am transferring herein to the new

genus Hyperietta. The true Lestrigonus fabrei has

not been recorded since the original accounts by

Milne Edwards (1830, 1838, 1840), which were limited to the male. Milne Edwards unfortunately

gives little information concerning the characters

that I have found most useful for distinguishing

genera and species of Hyperia s.l. His illustrations

(1840) include a lateral view of the male and drawings of the Md and Mxp; copies of these drawings

are given in Figure 27. I am unable to add any

details because the type-specimen appears to have

been lost. Dr. J. Forest, Museum National d'Histoire Naturelle, Paris, has kindly informed me that

a vial containing a label reading "Lestrigonus fabrei. Mer des Indes. Raynaud coll." contains a

second label in the handwriting of Bouvier stating

that the vial was returned empty by Bovallius. For

the time being L. fabrei remains a nomem dubium.

FIGURE 27.—Lestrigonus fabrei: a, $ lateral; b, Md; c, Mxp. Redrawn from Milne Edwards,

1840, pi. 30.
38 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Milne Edwards' species can be excluded without

difficulty from the genera of Hyperiidae other than

Lestrigonus considered herein. Following each

genus in the list below, I give the characters of

L. fabrei that exclude it from that genus.

Hyperia and Hypericlla: Pereonites 1-2 fused; coxae fused

with pereonites.

Hyperioides: P5-7 subequal.

Themis tella: Pereonites 1-2 fused; P5-7 subequal; Mxp with

well-developed inner lobe.

Hyperietta: Mxp with long narrow outer lobes and welldeveloped inner lobe; no long spines on s4-5 of PS-4.

Hyperionyx: Mxp with long narrow outer lobes; P5-7 subequal.

Only the rediscovery of Lestrigonus fabrei will enable us to know with certainty whether it agrees

with the details in my emended diagnosis of Lestrigonus. There is nothing seriously incompatible

and illustrations except possibly the body length

cited by him, "long d'environ cinq lignes." This

length is about 11 mm, much longer than the species herein assigned to Lestrigonus, which rarely

exceeds 4 mm. I suspect, but cannot prove, that

Milne Edwards' measurement was inaccurate.

Attempting to sort out the valid species of Lestrigonus and their synonyms from published

descriptions and illustrations has been very frustrating. In some instances previous authors assigned males and females or adults and juveniles

of one species to different species or even different

genera. Much of the confusion can be attributed

to the fact that these authors, following Bovallius,

used the number of fused pereonites as a primary

character for separating species, but failed to realize that in any species the fusion is greater in

females than in males and greater in juveniles than

with this diagnosis in Milne Edwards' description in adults. One can scarcely blame Pirlot (19S9)

Key to the Species of Lestrigonus1

FEMALES

1. Pereonites 1-3 fused 2

Pereonites 1-4 fused 3

Pereonites 1-5 fused 19. L. bengaletuis

2. Gland cone pointed, extending ventrally beyond buccal mass. Spine on distal margin of

s6 of P6-7 slender, simple 14. L. schixogeneios

Gland cone rounded, not reaching ventral border of buccal mass. Spine on distal margin of

s6 of P6-7 robust, with 1-2 teeth on anterior margin 15. L. crucipes

3. Head more than half as long as high, as long as fused pereonites 1-4. Gland cone reaching

or exceeding ventral border of buccal mass. Telson about half as long as Up3 protopod

17. L. macrophthalmus

Head about half or less than half as long as high. Gland cone not reaching ventral border

of buccal mass. Telson about 3/5 as long as protopod of Up3 4

4. Fused pereonites with partial sutures dorsally. S6 of Pl-2 with 1 spine on anterior margin

16. L. shoemakeri

Fused pereonites 1-4 without partial sutures. S6 of Pl-2 with 1-2 spines on anterior margin

18. L. latissimus

MALES

1. Pereonites 1-2 fused 2

Pereonites 1-4 fused 19. L. bengalensis

2. P5-7, distal margin of s6 produced into recurved dentate hook 15. L. crucipes

Distal margin of s6 not produced in P5, in P6-7 produced into smooth spine 3

3. Gland cone bluntly rounded, clearly not reaching ventral margin of buccal mass 4

Gland cone rather pointed, reaching or nearly reaching ventral margin of buccal mass

14. L. schixogeneios

4. Head more than half as long as high 17. L. macrophthalmus

Head less than half as long as high 5

5. From Pacific Ocean 16. L. shoemakeri

From Atlantic Ocean or Mediterranean 18. L. latissimus

1L. fabrei is omitted because its characters are largely unknown.
NUMBER 1S6 39

for retreating from the resultant problem by reducing most of the species herein assigned to Lestrigonus to synonyms of Hyperia bengalensis. Pirlot's action only added to the confusion, and

unfortunately a number of subsequent workers

have preferred to take refuge in his synonymy

rather than to attempt the formidable task of sorting out the valid synonyms without the aid of adequate collections.

A definitive taxonomic treatment of the species

of Lestrigonus is not possible with the material

now available to me, but I hope that the arrangement of species proposed herein will decrease somewhat the chaos that now confronts us.

14. Lestrigonus schizogeneios (Stebbing)

FIGURES 28-30

Hyperia schizogeneios Stebbing, 1888:1391-1394, pi. 168.-

Bovallius, 1889:221-224.-Chevreux, 1892:233-237. figs. 1-3;

1900:139-143, pi. 17: fig. lam; 1935:188-189.-Vosseler,

1901:66-67.-Lo Bianco, 1902:446 [identification by Vosseler];

1903:table facing p. 278 [identification by Vosseler].-Stewart, 1913:255-256.-Stephensen, 1924:86-90, Chart 13 -

Spandl, 1924b:265.-Chevreux & Fage, 1925:402-404, fig.

402-Pirlot, 1929:120-122; 1930:16-18.-Candeias, 1934:4-

Irie, 1948:36 [in table]; 1957a:351, fig. 12; 1957b:passim;

1958:107 [in table]; i959:passim.-Alvarado, 1955:219-

Hure, 1955:47-48, fig. 57a-b; 1961:33.-Bulycheva, 1955:1048

[in list].-Tregouboff & Rose, 1957:456, pi. 132.-Yang,

1960:15-19, figs. l-3.-Laval, 1965:6197-6198; 1968:passim;

1972:passim.-Vives, 1966:96, table 19; 1968:460, table 1 -

Dick, 1971:56-57.-Hure, Scotto di Carlo, & Basile, 1971:

passim.—Yoo, 1971:56, fig. 14.

Hyperia promontorii Stebbing, 1888:1385-1387, pi. 166B;

1910:475-476.-Bovallius, 1889:214-218, pi. 11: figs. 3-13 -

Vosseler, 1901:64.-Lo Bianco, 1901:446 [identification by

Vosseler]; 1903:table facing p. 278 [identification by Vosseler].-Barnard, 1930:411; 1937:183-184.-Dakin & Colefax,

1940:121, fig. 207.-Siegfried, 1963:8.

Hyperia bengalensis (Giles).—Pirlot, 1939:35-36.—Hurley,

1955:137-140, figs. 70-82; 1956:14-15; 1960b:279; 1969:19-

Reid, 1955:17, fig. 4.-Kane, 1962:299-300.-Vinogradov,

1962:24-25.-Brusca, 1967a:388; 1967b:452.

Hyperia zebui Stebbing, 1888:1394.

DERIVATION OF NAME.—From the Greek

=1 cleave] -\-}f£Vgtor[=a chin], referring to the

emarginate lower border of the head.

TYPE-LOCALITY.—Off St. Vincent, Cape Verde Is.,

le^^N, 25°14'W.

DIAGNOSIS.—Length of 9 2.2-3.5 mm, of tf 3.5-

4.5 mm. Head about twice as high as long, about

half as long as pereon. Pereonites 1-3 fused in $,

1-2 in o". Gland cone in 9 conspicuous, rather

sharply pointed, extending beyond buccal mass; in

cf blunter, not quite or barely reaching ventral

border of buccal mass. Md incisor with 8 teeth;

lacinia with 6-8 teeth. Mxl inner lobe with 3 long

and 2 short terminal spines, 1 subterminal spine.

Mx2 outer lobe with 1 subterminal and 2 terminal

spines; inner lobe with a shorter terminal spine.

Mxp outer lobes conical, about 3 times as long as

wide, with 3-4 spines along inner margin. S2 of PI

with moderately convex anterior margin; s4 with

2-3 (rarely 4) posterodistal spines; carpal process

with 5-6 spines; s6 with 2 (Atlantic specimens) or

3 (Pacific specimens) spines near anterior margin.

P2 with carpal process about half as long as s6; s6

with 1-2 spines on anterior margin. P3-4, posterior

margin of s5-6 with comb of spinules; s5 also with

2 spines on posterior margin. P5-7 with moderately broad s2 armed with a few spines on anterior

margin; anterior margins of s5-6 with comb of

spinules; s7 of P5 with a few anteroproximal spinules, s6 of P6-7 with spine on distal margin overlapping base of s7 medially. Telson triangular, in

9 about half, in <$ about 2/5 as long as protopod

of Up3.

VARIATION.—Specimens from the Atlantic and Pacific are very similar, but have slight differences. I

have not studied these differences in detail, but

I have examined the armature of the anterior margin of s6 of PI, which was the most obvious difference to me, in my specimens from both oceans.

Most of the adult Atlantic specimens had 2 spines

on this segment, but occasionally there were 3.

One female had 3 spines on the right and 2 on the

left PI. Pacific specimens (from CalCOFI collections) usually had 2-3 spines in the adult female

(stage 7 of Laval, 1968) and 3-4 in the adult male.

The number of spines is less in younger individuals. Stage 5 and younger females have only 1 spine;

in stage 6 there are 1-2 spines. Other groups of

spines vary in the same way; the older the specimen the more spines in a group.

The number of fused pereonites also varies with

maturity, as shown by Yang (1960) and Laval

(1968). Laval found that young specimens from

the marsupium, characterized by rudimentary

pleopods and uropods and named by Laval "protopleon larvae" (stage 1), have all 7 pereonites
40 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 28.—Lestrigonus schizogeneios, (a-c, <J) a, lateral view; b, ventral part of head, lateral; c,

Al. (d-l, $ ) d, ventral part of head, lateral; e, left Md; f-g, incisors of right and left Md; h,

Mxl; i, Mxl palp, flattened; ;, Mx2; k, Mxp, lateral; /, Mxp outer lobe, posterior surface.

(California specimens, except d, from Sargasso Sea.)

free. CalCOFI specimens removed from the marsupium also have no fusion of pereonites. Laval

observed that after the protopleon larvae leave the

marsupium and settle on the host medusa, Phialidium sp., they molt into juveniles with pereonites

1-5 fused (stage 2). Three more instars, stages 3-5,

follow in which pereonites 1-5 are fused. The

female then enters the stage of prepuberty, stage

6, in which pereonites 1-4 are fused and rudimentary oostegites are present. Stage 7 is the adult,

with pereonites 1-3 fused and fully developed

oostegites. In the male the flagella of Al-2 remain

short and unsegmented in stage 6 (pereonites 1-4

fused) and stage 7 (pereonites 1-3 fused). Stage 8

is the prepuberty stage, with pereonites 1—2 fused

and the flagella of Al-2 segmented but only about
NUMBER 1S6 41

FIGURE 29.—Lestrigonus schizogeneios (a-g, $): a, PI; b, P2; c, P4; d, P5; e, P6; /, P7; g, urosome;

/i, $ urosome. California specimens.

as long as the pereon. The male does not become

fully adult until stage 9, thus he requires 2 more

molts than the female to attain maturity.

I have examined many specimens of L. schizogeneios from the CalCOFI collections and have

found that the sequence of development agrees

closely with that described by Laval.

It is generally agreed that Hyperia schizogeneios

and H. promontorii are the female and male respectively of the same species. Both specific names

have been used, but although H. promontorii has

page precedence, the first revisor, Stephensen

(1924), selected the name schizogeneios. Pirlot's

(1939) indiscriminate lumping of 12 nominate species under H. bengalensis has made it difficult to

know the real identity of specimens reported as H.

bengalensis by subsequent authors. I have, for example, examined those reported by Shoemaker

from Bermuda (1945) and Cuba (1948) and found

them to be Lestrigonus latissimus, hence Shoemaker's references are not included in the synonymy

given above for L. schizogeneios.

DISTRIBUTION.—Reported from warm waters

around the world. In the CalCOFI area it is the

second most abundant hyperiid, being exceeded

only by Parathemisto pacifica. It extends farther

south than the subarctic P. pacifica, but is replaced

in the southernmost part of the CalCOFI area by

the closely related L. shoemakeri. When its overall

distribution in the Pacific becomes known it may

prove to be an inhabitant of the transition zone

(Brinton, 1962). Its abundance in the samples of

the three 1949 CalCOFI cruises was least in March,

greater in July, and highest in November, the average number per 1000 m3

 for the positive stations

increasing from 17 to 64 to 102.
42 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE SO.—Distribution of Lestrigonus schizogeneios on 4 CalCOFI cruises.
NUMBER 1S6 43

15. Lestrigonus crucipes (Bovallius)

FIGURE 31

Hyperia crucipes Bovallius, 1889:225-228, pi. 11: fig. 14-25-

Walker, 1904:236.-Stephensen, 1924:90.-Barnard, 1937:

184.-Pirlot, 1939:36.-Laval. 1968:64-65, fig. 12.-Dick,

1970:55.

DERIVATION OF NAME.—From the Latin "crucis"

[=across] -)- "pes" [=foot], referring to spine overriding dactyl on P5-7.

TYPE-LOCALITY.—"The tropical region of the Atlantic: off Barbadoes."

DIAGNOSIS.—Length about 4 mm. Body rather

plump and compact. Pereonites 1-2 fused in o"»

1-3 in 9 • Pleonal epimera with rounded posterior

corners. A1-2 of $ well developed for a Lestrigonus; gland cone obtuse, projecting obliquely anteriad. Md with broad molar; incisors with 8-9

teeth. Mxl inner lobes with 5 apical spines. Mx2

with 2 and 1 spines on outer and inner lobes respectively. Pl-2 with 3-4 spines on anterior margin of s6; carpal process of P2 reaching beyond

midlength of s6; s7 of Pl-2 with spinulose posterior

margin. P3-4 longer than P5-7; s4-5 with 2-3

spines on posterior margin; s6 posterior margin

armed with close-set spinules. P5-7 with broad s2

and rather robust s3-6; s6 produced distally into

recurved hook with 1-2 spiniform teeth on convex

margin; hook overlaps s7 medially. Up 1-3 with

slender peduncles and rami. Telson oval, slightly

wider than long, about 0.4 the length of Up3

peduncle.

RELATIONSHIPS.—The specimens that I have examined, all from the Arabian Sea, differ from Bovallius' description so much that I was at first convinced that I was dealing with an undescribed

species. But no specimens agreeing more nearly

with Bovallius' account have come to hand, and

the frustrations of trying to reconcile the species

descriptions of Bovallius with the characters that

can be seen on his species have led me to share the

opinion of Bovallius' work expressed by Stephensen

(1924), who wrote in evident exasperation, ". . .

but in consideration of several positive misstatements in the work of Bovallius, there is hardly any

reason to pay much attention to his remarks . . ."

I am therefore assuming considerable error in Bovallius' account of Hyperia crucipes. The principal

discrepancies between Bovallius' description and

the specimens that I have examined are the

following:

Bovallius' description

pereonites 1-3 fused in $

P5-7 subequal in length

to P3-4

anterior margin s6 of Pl-2

without spines

P5-6 with forked

projection at base of s7,

1 branch of fork crossing

s7 on each side (Fig.

Sir)

P7 without projection at

base of s7

telson half length of UpS

protopod

HOE specimens

pereonites 1-2 fused in $

P5-7 distinctly shorter than

P3-4

same with 3-4 spines

P5-7 with toothed recurved

hook projecting on medial

side of s7 from distal

margin of s6 (Fig. 31q)

P7 as in P5-6

telson<half length of UpS

protopod

My disbelief in Bovallius' accuracy was reinforced by examination of a female amphipod in

the collections of the Stockholm Museum from the

Caribbean Sea east of Dominica (15°22'N,

62°41'W). This specimen, labeled Hyperia crucipes

var, macropis by Bovallius (an unpublished varietal name), agrees closely with the HOE specimens.

I have also examined Stephensen's (1924) specimens from Thor stations 266, 377, and 399; they

also closely resemble the HOE specimens.

DISTRIBUTION.—Warm regions of the Atlantic

(Bovallius, Pirlot, Laval, Stephensen) and Indian

(Walker, Barnard) Oceans. The few specimens

that I have examined were collected at three stations in the Arabian Sea during the International

Indian Ocean Expeditions.

16. Lestrigonus shoemakeri, new species

FIGURES 32-34

DERIVATION OF NAME.—After the late Clarence

R. Shoemaker, in recognition of his many contributions to amphipod taxonomy.

TYPES.-Holotype ? t USNM 139096, allotype d1.

USNM 139097, CalCOFI Cruise 9, station 1106,

27°48'N, 119°14'W, west of Punta Eugenia, Baja

California, 12 November 1949. 2 cf and 2 $ paratypes deposited at Scripps Institution of Oceanography.

DiAGNOsis.-Length of $ 2.3-2.7 mm, of <? 3.5-

4.0 mm. Head slightly more than twice as high as

long, about y3 as long as pereon. Pereonites 1-4

fused in $, but sutures present dorsally, that between pereonites 1-2 shorter than others. Pereonites

1-2 fused in <f; partial sutures usually absent in
44 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 31.—Lestrigonus crucipes, 9 : a, lateral view; b, head, anterior view; c, pleonal epimera;

d, telson and uropods, dorsal; e, protopod of Up3, showing distoventral serrations; /, Al; g,

A2; h, left Md; i, Mxl, inner lobe; ;, Mx2; k, Mxp; /, PI; m, P2; n, P3; o, P5; p, P7; q, P7, distal

end, medial; r, P6, distal end (copied from Bovallius, 1889, pi. 11: fig. 23).
NUMBER 136 45

FIGURE 32.—Lestrigonus shoemakeri: a, 9. lateral; b, 9 head and pereonites 1-5, lateral; c,

ventral part of $ head, lateral; d, $ head and pereonites 1-3, lateral; e, ventral part of

$ head, lateral; /, 9 Al; g, $ Al; h, incisor of 9 right Md; i, palp of $ Md; ;, 9 Mxl; k-l,

9 Mxl, apices of palp and outer lobe; m, 9 Mx2; n, Mxp; o, 9 PI; p, 9 P2. (b, d, f, g, i, j ,

m, n, o, p from off Baja California; a, c, e, h, k, I from off Pacific coast of Nicaragua.)
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 33.—Lestrigonus shot makeri (a-m, 9) a, PI; b, P2; c, carpal process of P2 from above; d,

meral process of P2 from above; e, P3; /, P4; g, P5; h, P6; i, P7; /-/, dactyl and distal part of

propus of P5-7, lateral; m, urosome. n. $ urosome. (e-i, m,n from off Baja California; a-d, j~l

from off Pacific coast of Nicaragua.)
NUMBER 136 47

9

FIGURE 54.—Distribution of Lestrigonus shoemakeri on 4 CalCOFI cruises.
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

adult, present in immature specimens with more

than 2 pereonites fused. Gland cone bluntly

pointed, overlapping epistome, but not reaching

ventral border of buccal mass. Md incisor with 8-9

teeth; lacinia with 7-8 teeth. Mxl inner lobe with

3 long terminal spines and 1 shorter subterminal

spine. Mx2 outer lobe with 1 subterminal and 2

terminal spines; inner lobe with shorter terminal

spine. Mxp outer lobes conical, with about 5 spines

along inner margin. S2 of PI with strong bulge

on anterior margin; s4 with 2-4 posterodistal

spines; carpal process with 5-6 spines; s6 with single

spine near anterior margin. P2 with carpal process

more than half as long as s6; s6 with 1-2 spines on

anterior margin. P3-4, posterior margins of s5-6

with comb of spinules; s5 with 2 spines, s4 with 1

spine on posterior margin. P5-7 with moderately

broad s2 armed with a few spines on anterior margin; s5-6 with comb of spinules on anterior margin;

s7 of P5 with a few anteroproximal spinules; s6 of

P6-7 with spine on distal margin overlapping s7

medially. Telson of 9 nearly %, of J about half

as long as Up3 protopod.

VARIATION.—As in Lestrigonus schizogeneios more

pereonites are fused in juveniles than in adults.

The youngest free-living individuals have pereonites 1-5 fused. With increasing maturity the

number of pereonites fused decreases until the

adult condition is reached: pereonites 1-4 fused

in the female, 1-2 fused in the male.

DISTRIBUTION.—Thus far I have identified L. shoemakeri only in samples from the warmest stations

of the 4 CalCOFI cruises (Figure 34) and in a few

samples in the eastern equatorial Pacific: (1) a few

miles north of Clipperton Island; (2) west of Nicaragua, 12°35'N, 93°40rW; (3) west of Costa Rica,

10°03'N, 88°52/W; (4) west of Panama, 7°36'N,

85°09^V; (5) near the Galapagos Islands, 00°07'S,

86055rWT. These few records suggest that when better known the distribution of L. shoemakeri may

be similar to that of Euphausia eximia and the

other euphausiids that Brinton (1962) classifies as

eastern Pacific equatorial species.

17. Lestrigonus macrophthalmus (Vosseler)

FIGURE 35

Hyperia macrophthalma Vosseler, 1901:70-72, pi. 6: fig. 16-

25.-Yang, 1960:19-28, figs. 4-5.-Dick, 1970:56.

[?] Hyperia hydrocephala Vosseler.—Dakin & Colefax, 1940:

121. fig. 206a-c.

DERIVATION OF NAME.—Not given, presumably

from the Greek )*/»(()(& [=long] -f Offdtdvcs

[=eye]. Although Vosseler was technically incorrect grammatically in giving the noun thus formed

a feminine ending and thereby treating it as an

adjective (the correct adjective would have been

macrophthalmata), I will continue the practice of

other authors in considering the name an adjective.

TYPE-LOCALITY.—Six stations of the PlanktonExpedition in the tropical Atlantic.

DIAGNOSIS.—Length of 9 2-3.5 mm. of o* 3-4

mm. Head more globular than in other species,

more than half as long as high, about as long as

pereonites 1-4 combined. Pereonites 1-4 fused in

9, 1-2 fused in cf. Gland cone bluntly rounded

below, in 9 reaching or extending slightly beyond

ventral border of buccal mass, in o*

 n °t reaching

this border. Md incisor and lacinia with 7 teeth

in immature 9 dissected. Mxl outer lobe with

somewhat tapering apex extending well beyond

distomedial spine. Mxp outer lobes slender, with

2 terminal setae and 2-3 setae on medial margin;

inner lobe with 2 terminal spines. S2 of PI 'with

strong convexity on anterior margin; s4 with 2

posterodistal spines; s6 with 1 (9) or 2 (cf) spines

on anterior margin. P2 with carpal process about

half as long as s6; s6 with 1 (9) or 2 (c?) spines

on anterior margin. S7 of P5 about Vi~V4 length

of s6, 14 shorter than s7 of P6-7, armed with a few

anteroproximal spinules; s6 of P6-7 with spine on

distal margin overlapping base of s7 medially. Telson triangular, in 9 about half, in o* less than

half as long as protopod of Up3.

VARIATION.—As in other species of Lestrigonus

the number of fused pereonites is greater in immature specimens, but never exceeds 5.

RELATIONSHIPS.—This species agrees with L. latissimus in having pereonites 1-4 fused in the female,

but is distinguished by its smaller size and more

globular head. In lateral view the gland cone overlaps the epistome less in macrophthalmus than in

latissimus. Up protopod is more slender and longer

in relation to the telson than in L. latissimus. S6

of Pl-2 bears a single spine, whereas in L. latissimus s6 usually has 2 spines, but has 1 in immature

specimens and in some adults. S7 of P5 is relatively shorter in latissimus than in macrophthalmus.

H. hydrocephala has been considered identical

with H. macrophthalma, but I consider it to be an
NUMBER 1S6 49

FIGURE 35.—Lestrigonus macrophthalmus, 9: a, lateral view; b-c, head, lateral; d, indsor,

right Md; e-f, outer lobe and palp of Mxl; g, Mxp; h-j, PI; k, P2; /, P3; m, P4; no, P5; p-q, P6;

r-s, P7; /, urosome. (a, i, k-m, o, q, s, t from East China Sea [Albatross station 4909]; b,j from

off South Carolina [T. N. Gi7/ Cruise 3, station 59]; d-g, n, p, r from Sargasso Sea [Weather

station "E"]; c, h from Gulf of Guinea [Ombango Cruise 14, station 329].)
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

immature L. latissimus (see discussion under the

latter species). Dakin and Colefax's (1940) H. hydrocephala has the globular head of L. macrophthalmus and may be a young specimen in which

pereonites 1-5 are fused.

DISTRIBUTION.—Tropical parts of the Atlantic, Pacific, and Indian Oceans. Not found in the CalCOFI area.

18. Lestrigonus latissimus (Bovallius)

FIGURE 36

Hyperia latissima Bovallius, 1889:229-232, pi. 11: figs. 26-

36.-Vosseler, 1901:67.-Chevreux & Fage, 1925:404-405-

Stephensen, 1928:590.-Candeias, 1934:4, fig. 3.-Vives,

1966:96, table 19; 1968:460. table 1.

Hyperia hydrocephala Vosseler, 1901:74-76, pi. 6: figs. 26-28,

pi. 7: figs. l-5.-Steuer, 1911:677-679, pi. 2.—Stephensen,

1924:91-92, Chart 14.-Gamulin, 1948:21.

Hyperia sp. ? hydrocephala Vosseler.-Pesta, 1920:30-31, fig.

5a-c.

Hyperia bengalensis (Giles) .-Shoemaker, 1945b:2S8; 1948:

12-13.

DERIVATION OF NAME.—Not given; presumably

from the Latin "latus" [=broad], referring to

the plump body.

TYPE-LOCALITY.—"The Southern temperate region of the Atlantic" (Bovallius, 1889).

DIAGNOSIS.—Length of ? 2-3 mm, of tf 3-4 mm.

Head about 2.2 times as high as long, shorter than

pereonites 1^1 combined. Pereonites 1-4 fused in

$, 1-2 fused in <$. Gland cone bluntly round below, not reaching ventral border of buccal mass.

Md incisor and lacinia with 10 teeth. Mxl outer

lobe with truncate apex. Mxp outer lobes slender,

with 3 apical setae and 2-4 setae on medial margin.

PI with pronounced bulge on anterior margin of

s2; s4 with 4 posterodistal spines; s6 with 1 or 2

spines on anterior margin. P2 carpal process about

half as long as s6; s6 with 2 spines on anterior

margin. S7 of P5 about 14-% length of s6, i/s

shorter than s7 of P6-7, armed with a few anterodistal spinules; s6 of P5-7 with spine on distal

margin overlapping base of s7 medially. Telson

triangular, in 9 about %, in cf about i/2 as long

as protopod of Up3.

RELATIONSHIPS.—L. latissimus is very similar to

L. shoemakeri, but the two species appear to be

allopatric, the former inhabiting the Atlantic, the

latter the Pacific. The partial sutures on fused

pereonites 1-4 of L. shoemakeri are absent in L.

latissimus. L. macrophthalmus has a distinctly

more globular head and is a smaller species.

I believe Vosseler's H. hydrocephala is an immature L. latissimus in which pereonites 1-5 are

fused and not, as Stephensen (1924) maintained,

conspecific with L. macrophthalmus. I have examined the specimens identified by Stephensen

(1924) as H. hydrocephala from Thor station 224.

The sample contains mature females with pereonites 1-4 fused and immature females with pereonites 1-5 fused. S6 of PI has 1 spine on the

anterior margin in immature specimens; mature

specimens usually have 2 spines, but some have

only 1. The Thor specimens agree with L. latissimus rather than with L. macrophthalmus in body

size, gland cone-buccal mass relationship, and

length of the telson in relation to Up3 protopod.

Contrary to Stephenson's opinion, therefore, I consider macrophthalma to be distinct from hydrocephala, and believe the latter to be an immature

L. latissimus.

DISTRIBUTION.—Reliably known only from the

eastern Atlantic and the Mediterranean.

19. Lestrigonus bengalensis Giles

FIGURES 37-38

Lestrigonus bengalensis Giles, 1887:224-227, plates 6-7.

Hyperia bengalensis (Giles) -Bovallius, 1889:199-201, fig.

[copied from Giles].-Walker, 1904:235; 1909:51.-Nayar,

1959:4fr-47, plate 16: figs. 1-5.

Hyperia dysschistus Stebbing, 1888:1388-1391, plate 167.—

Bovallius, 1889:204-206, plate 11: figs. l-2.-Spandl, 1924a:

23; 1924b:265.

Hyperia thoracica Bovallius, 1889:233-236, plate 11: figs.

37-41.-Vosseler, 1901:73-74, plate 6: figs. l-4.-Stephensen,

1924:91.-Lewis and Fish, 1969:9.

Hyperia gilesi Bovallius, 1889:236-239.

Hyperia atlantica Vosseler, 1901:67-70, plate 6: figs. 5-15.—

Yang, 1960:28-33, figs. 6-7.-Dick, 1970:55.

Hyperia latissima Bovallius.-Barnard, 1930:410-411 [misidentification].

Hyperia hydrocephala Vosseler.—Dakin and Colefax, 1940:

121, fig. 206 [misidentification].

DERIVATION OF NAME.—Not given, presumably

geographical (Bay of Bengal).

TYPE-LOCALITY.—"About 100 miles from land in

the Bay of Bengal, the depth of the water in the

locality being 850 fathoms."
NUMBER 1SS 51

FIGURE 36.—Lestrigonus latissimus (a-l, $): a, head and pereon, lateral; b, ventral part of head,

lateral; c, incisor of left Md; d-e, Mxl, outer lobe and palp; /, Mxp; g, PI; h, P2; i, P5; /, P6; k,

P7; /, urosome; m, ventral part of $ head, lateral, (a from Thor station 224; b-f, i-l from

Ombango Cruise 12, station 300; g-h from Ombango Cruise 14, station 329; m from Ombango

Cruise 13, station 308.)
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DiAGNosis.-Length of $ 2-2.5 mm, of tf 2.8-3.4

mm. Head of 9 about twice, of cf about 1.6 times

as high as long. Pereonites 1-5 fused in 9> 1-4

fused in cf. Gland cone of 9 produced anteroventrally, overlapping and extending anterior to

epistome; apex bluntly rounded; posterior margin

subparallel to body axis. Gland cone of cf produced ventrally as rounded lobe, extending only

slightly anterior to epistome. Md incisor with 8

teeth; lacinia with 7 teeth. Mxl inner lobe with 3

long terminal spines and 1 shorter subterminal

spine. Mx2 outer lobe with 1 subterminal and 2

slender terminal spines; inner lobe with 1 shorter

terminal spine. S2 of PI with strong bulge on

anterior margin; s4 with 2 posterodistal spines; s5

with 1-2 spines on posterior margin and 3 on carpal process; s6 with single spine near posterior

margin. S2 of P2 quite broad for a Lestrigonus;

carpal process slightly less than half as long as s6,

with 7 marginal spines. P3-4, posterior margins

of s5-6 with comb of spinules; s4-5 with posterodistal spine. P5-7 with moderately broad s2 armed

with 2-4 spines on anterior margin; s5-6 with comb

of spinules on anterior margin; s7 of P5 % as long

as s7 of P6-7, armed with a few anteroproximal

spinules; s7 of P6-7 sometimes with 1-2 anteroproximal spinules; s6 of P6-7 with spine on distal

margin overlapping s7 medially. Telson of 9

slightly more than i/2, in cf about % as long as

Up3 protopod.

VARIATION.—It is entirely possible that more than

1 species of Lestrigonus agrees with the above diagnosis. There is a noticeable variation in some characters in different populations included here under

L. bengalensis, and illustrations are provided to

show some of the variation in the armature of the

distal segments of P5-7. Until sufficient collections

for an adequate study of variations are available,

I am including all Lestrigonus with 5 pereonites

fused in the 9 and 4 in the cf under L. bengalensis, realizing that this may be a gross oversimplification. There is some indication from available

collections that L. bengalensis inhabits coastal

waters, which would lead to populations becoming

isolated and tending to differentiate from one another.

Stebbing's (1888) drawing of an immature male

H. dysschistus in lateral view (plate 167) shows a

complete suture between pereonites 1 and 2 and

fusion of pereonites 2-5. This suture is not mentioned in Stebbing's description, which states that

pereonites 1-5 are fused. I have examined the

holotype of H. dysschistus and can confirm the complete fusion of pereonites 1-5. Nevertheless, Bovallius (1889) considered a free pereonite 1 to be

the outstanding diagnostic character of H. dysschistus and stated that he had examined specimens

which exhibited this peculiar segmentation. Spandl

(1924a, 1924b) also claimed to have identified similar specimens of H. dysschistus.

Among the hundreds of specimens of Lestrigonus

that I have examined, I have never found one with

pereonite 1 free and succeeding pereonites fused,

and I am convinced that such a condition does not

exist in this genus. I suggest that Stebbing, Bovallius, and Spandl all mistook muscle bands or some

other structure beneath the integument for a suture. Such a mistake can be expected in the study

of small hyperiids, where sutures and other markings on the rather delicate and semitransparent

cuticle are often less apparent than subcuticular

structures. Such internal structures have frequently

been mistaken for sutures between the coxae and

their pereonites. In his lateral view of Hyperia

dysschistus Stebbing shows free coxae, as he also

does for Hyperia schizogeneios on plate 168. Bovallius (1889) shows distinct coxae on Hyperia

fabrei, H. promontorii, H. crucipes, H. latissima,

and H. thoracica, none of which have them. Other

carcinologists who have portrayed species of Lestrigonus with what appear to be sutures between the

coxae and their pereonites are Chevreux, Hurley,

Irie, and Stephensen.

Giles (1887) described Lestrigonus bengalensis

as having 7 free pereonites, but remarked on "the

junction between the pleura and the coxal plates

being hard to make out, as are also the junctions

of the terga of the first 5 thoracic segments." In

his illustrations the adult male has 7 free pereonites, but an immature male is shown with pereonites 1-5 fused and a female with pereonites

1-5 fused. Bovallius (1889) placed H. bengalensis among the species with 7 free pereonites,

but it seems evident that Giles portrayed correctly

the condition of the female and young male; L.

bengalensis has pereonites 1-5 fused in the 9 an ^

1-4 in the male.

The specimens that Barnard (1930) identified as

Hyperia latissima are placed here under L. bengalensis because of Barnard's statement "The present
NUMBER 1S6 53

FIGURE 37.—Lestrigonus bengalensis (a-q, 9): a, lateral view; b-c, head, lateral; d-h, ventral part

of head, lateral; i, Al; /, A2; k, incisor and lacinia of left Md; l-m, outer lobe and palp of Mxl;

n, Mx2; o, Mxp, lateral; p, outer lobe of Mxp, posterior; q, urosome; r, $ head, lateral; s,

$ Al; t-u, $ telson and Up3. (a, i, s, t from off southern Baja California; b, g from Gulf

of Camby, India; c, h, j-q from off Pacific coast of Central America; d, from off Savannah,

Georgia; e from Gulf of Guinea; /, r from Bora Bora, Society Islands; u from Moorea, Society

Islands.)
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Y /'/

FIGURE 38.—Lestrigonus bengalensis (a-s, $): a-c, PI; d, P2; e, P2, carpal process from above; /,

P3; g, P4; h, P5; i, P6; /, P7; *-*, dactyls of P5-7; t-v, dactyls of $ P5-7, from Moorea, Sodety

Islands, (a, d, f-j from off Baja California; b, e from off Pacific Coast of Central America; c

from Gulf of Cambay; km from off Savannah, Georgia; n-p from Gulf of Cambay, India; q-s

from off Pacific coast of Nicaragua.)
NUMBER 136 55

specimens have segments 1-4 in o", 1-5 in 9 coalesced."

DISTRIBUTION.—Worldwide in tropical waters,

tending to occur in coastal waters.

V. Hyperietta, new genus

DIAGNOSIS.—Small species, with body rather compressed laterally. Head rather short anteroposteriorly; eyes occupying most of its surface. Pereonites 1-2 fused in both sexes. Coxae fused with

pereonites. Telson inserted distinctly anterior to

insertion of Up3 peduncle; 9 telson at least half

as long as Up3 protopod. Epistome small, inconspicuous. 9 Al 2-merous. 9 A2 1-merous, rudimentary; gland cones rather inconspicuous, in

anterior view converging medially. Md with

smooth incisor and dentate lacinia; molar narrow;

palp absent in 9 • Mxl outer lobe with 3 terminal

spines. Mxp outer lobes fused medially; inner lobe

rudimentary. PI simple, weakly subchelate, or

barely chelate. P2 chelate, with spoon-shaped carpal

process bearing marginal spines. 9 P3-4 with 1

conspicuous spine on posterior margin of s4, 2

spines on posterior margin of s5; <f P3-4 with

shorter spines in same positions. Distal margin of

s6 of P6-7 and sometimes of P5 produced into

spinose lobe medial to base of s7; s7 unarmed. Up

1-3 slender; margins of rami smooth or with extremely fine serrations.

DERIVATION OF NAME.—Diminutive of Hyperia,

gender feminine.

TYPE-SPECIES.—By present designation, Hyperia

hizoni Stebbing.

20. Hyperietta luzoni (Stebbing)

FIGURES 39-40

Hyperia luzoni Stebbing, 1888:1382-1384, pi. 166A.—Bovallius,

1889:212-214.-Lo Bianco, 1902:424, 425, 446 [identification

by Vosseler]; 1903: table facing p. 278 [identification by

Vosseler]; 1904:42.-Stewart, 1913:255.-Stephensen, 1924:

84-86, fig. 34. Chart 12.-Pirlot, 1929:122-123; 1939:35.-

Barnard, 1930:410.-Hurley, 196Ob:279; 1969:pl. 19 [map].-

Kane. 1962:301.

[not] Hyperia luzoni Stebbing.—Vosseler, 1901:64-66, pi. 5:

figs. 16-28.-Stebbing, 1904:33-34.

DERIVATION OF NAME.—Not given, presumably

for the type-locality.

TYPE-LOCALITY.—South China Sea, west of Luzon

Island, Philippines, 16°35'N, 117°47'E.

DIAGNOSIS.—Length of 9 2-3 mm, of Q* 3-4 mm.

Head slightly more than twice as high as long,

about half as long as pereon. Gland cone just

reaching or barely exceeding ventral margin of

head, posteroventrally overlapping anterior part of

buccal mass. Mxp outer lobes subpyriform, bearing

a few setae on distal margin and on shelf. PI simple;

s2 sinuate, proximal part strongly bulging anteriorly, distal part much narrower; s5 twice as

long as wide, with 1 spine at midlength and 2 at

distal end of posterior margin; s6 with 1 spine at

midlength of anterior margin. P2 very slender,

chelate; s2 about 6 times as long as wide; carpal

process % as long as s6; s6 with 2 spines on anterior

margin. P3-4 very slender. P5-7 with moderately

broad s2; other segments slender; s4-5 without long

spines. Telson round-triangular, slightly longer

than wide; in 9 about %, in 6* % as long as

protopod of Up3.

Key to the Species of Hyperietta

1. S5 of P5-7 with strong spine on anterodistal corner S

S5 of P5-7 without strong spine on anterodistal corner 2

2. PI, anterior margin of s2 evenly convex. P5, s2 about yA as wide as long 21. H. vosseleri

PI, anterior margin of s2 with deeply concave distal part. P5, s2 about 2/3 as wide as

long 20. H. luzoni

3. Anterior margin of head in lateral view flattened below insertion of Al. Pereon about 4

times as long as head 24. H. parviceps

Anterior margin of head in lateral view evenly rounded. Pereon not more than 2-3 times

as long as head *

4. S5 of PI with spine at midlength of posterior margin. S6 of Pl-2 with 1 spine on anterior

margin. Mxp with only a few spinules along inner margins of outer lobes 23. H. stephenseni

S5 of PI without a spine at midlength of posterior margin. S6 of Pl-2 with 2 spines on

anterior margins. Outer lobes of Mxp with dense covering of fine setules at apex and few

blunt spines on anterior surface 22. H. stebbingi
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 39.—Hyperietta luzoni, 9 off southern California; a, lateral view; b, head, lateral; c, head,

anterior; d, Al; e, left Md, incisor and lacinia; /, left Md molar; g, Mxl palp, posterior; h,

Mx2; i, Mxp, anterior /, PI; k, P2: /, P3; m, P4; n, P5; o, P6; p, P7; q, urosome.
NUMBER 136 57

FIGURE 40.—Distribution of Hyperietta luioni on 4 CalCOFI cruises.
58 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

RELATIONSHIPS.—The above synonymy includes

all references to H. luzoni known to me, but only

Stebbing's original account and that of Stephensen

(1924) can be attributed to this species without

question. Two new species described below are

very similar to H. luzoni, and all three may be

found in the same plankton sample. The absence

of descriptions or illustrations of H. luzoni in the

works of most authors who have recorded it makes

it impossible to determine from their publications

whether they in fact had H. luzoni or a similar but

different species. Vosseler's (1901) H. luzoni is the

new species described below as Hyperietta stebbingi.

The species referred to as Hyperia luzoni by Stebbing in 1904 is not H. luzoni because of the presence of spines on s5 of P5-7, but might have been

either of two new species proposed herein, Hyperietta stebbingi or H. stephenseni.

Stebbing's (1888) description and figures are of

an immature male. Stephensen (1924) described

and illustrated the adult male, but the female has

not yet been described or figured.

DISTRIBUTION.—Reported by Stebbing (1888) from

the Philippines (west of Luzon and Zebu Harbor)

and the eastern South Pacific (38°6'S, 88°2'W).

Stephensen (1924) listed it from a number of Thor

Mediterranean and a few Atlantic stations, but it

is possible that some of these records may be referable to one of the new species of Hyperietta described below.

In the CalCOFI collections it was present off the

coasts of southern California and Baja California

and farther offshore as far north as the latitude of

San Francisco. Its pattern of distribution is consistent with that of Euphausia brevis and the other

Pacific central euphausiid species (Brinton, 1962)

which occupy oceanic gyrals of mid-latitudes 15°-

40° in both hemispheres.

I have not found H. luzoni in any Atlantic

plankton samples.

21. Hyperietta vosseleri (Stebbing)

FIGURES 41-42

Hyperia fabrei H. Milne Edwards.-Bovallius, 1889:206-211,

pi. 10: figs. 40-55.-Vosseler, 1901:58-60, pi. 5: figs. 5-15-

Stephensen, 1924:83-84.-Spandl, 1924:22-23; 1927:153.—

Barnard, 1930:410.-Pirlot, 1939:34.-Hurley, 1955:137.-Reid,

1955:15.-Yang, 1960:33-35, fig. 8.-Kane, 1962:301.-Dick,

1970:55.

Hyperia vosseleri Stebbing, 19O4:33-34.-Stewait, 1913:255.-

Chevreaux, 1935:189.

DERIVATION OF NAME.—Not given, but obviously

after J. Vosseler, who described and illustrated this

species in detail.

TYPE-LOCALITY.— Tropical Atlantic.

DIAGNOSIS.—Length of 9 2-3 mm, of tf 3-4 mm.

Head slightly more than twice as high as long, in

profile narrowing ventrally more than in other

species of Hyperietta. $ Al reaching well beyond

ventral margin of head. Gland cone extending

beyond margin of head; posterior margin slightly

concave, separated by short gap from epistome.

Left Md with lacinia nearly as wide as incisor,

divided into 11-12 sharp teeth. Mxp outer lobes

slightly more than 1.5 times as long as wide; apex

narrow, bearing dense tuft of setae; distal margin

rugose. PI simple; s2 about twice as long as wide,

with evenly convex anterior margin; s6 with single

spine on inner surface near anterior margin. P2

chelate; carpal process nearly reaching midlength

of sfi. P5-7 with very broad s2, especially that of

P5, which has very convex anterior margin. Telson

about % as long as Up3 protopod in 9> slightly

less than half as long in Q* •

RELATIONSHIPS.—Stebbing (1904) believed that

the species named Lestrigonus fabrei by Milne Edwards (1830) was much too large ("long d'environ

cinq lignes"=iabout 11 mm) to be identical with the

species called Hyperia fabrei by Bovallius (1889)

and subsequent authors, which measured only 3-6

mm. Stebbing therefore proposed the name Hyperia

vosseleri for the smaller species. Most authors have

not accepted Stebbing's proposal, but it seems certain that he was correct. Not only is the size difference well beyond the limits of reasonable variation,

but the maxilliped is entirely different in the two

species. That of H. vosseleri has short, broad outer

lobes and a rudimentary inner lobe. (Figure 41/^),

whereas Milne Edwards (1840, pi. 30: fig. 20) portrays the maxilliped of L. fabrei as having long

slender outer lobes and a well developed inner

lobe (Figure 27c). I consider the two species sufficiently distinct that I have placed them in different genera in this paper.

DISTRIBUTION.—Reported from the warm parts of

the Atlantic, Pacific, and Indian Oceans. In the

CalCOFI area its distribution was quite similar to

that of H. luzoni (compare Figures 40 and 42), and
NUMBER 1S6 59

FIGURE 41 .—Hyperietta vosseleri (a-q, 9, off Baja California): a, lateral view; b, head, lateral; c,

head, anterior; d-e, incisors of left and right Md; /, Mxp, anterior; g, PI; h, P2; i, PS; ;',

P4; *, P5; /, P6; m, P7; n-p, dactyls of P5-7, medial; q, urosome. (r-s, $, Gulf of Guinea) r,

Mxl inner lobe; s, Mxp outer lobe.
60 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICUKE 42.—Distribution of Hyperietta vosseleri on 4 CalCOFI cruises.
NUMBER 1S6 61

it may be also characterized as a Pacific central

species.

22. Hyperietta stebbingi, new species

FIGURES 43-45

Hyperia luzoni Stebbing.-Vosselei, 1901:64-66, plate 5: figs.

16-28 [misidentification].

DERIVATION OF NAME.—The species is named for

the distinguished English carcinologist, T. R. R.

Stebbing.

TYPES.-AH from CalCOFI Cruise 9, station 607,

12 November 1949, off central California, 35°58'N,

127°51'W, oblique tow from 66 m to surface. Holotype $ USNM 137510, allotype J, USNM 137511,

20$ and 7cf paratypes, USNM 137512; 2? and 2

o* paratypes deposited at Scripps Institution of

Oceanography.

DIAGNOSIS.—Length of 9 2-3 mm, of tf 3-4 mm.

Head about twice as high as long, evenly rounded

ventrally. 9 Al reaching ventral margin of head.

Gland cone barely reaching ventral margin of

head, broadly rounded, separated from epistome

by distinct gap in lateral view. Lacinia of left Md

with about 8 teeth. Outer lobe of Mxp % longer

than wide, bearing a series of spines on both surfaces near distal margin and scattered minute

setules on distal part of anterior margin. S2 of PI

slightly more than twice as long as wide, both

margins moderately convex; s5 with very small

carpal process bearing 3^1 spines, posterior margin

otherwise unarmed; s6 about twice as long as broad,

with 2-3 spines on anterior margin. S6 of P2 bearing 2 spines on anterior margin. P6 only slightly

longer than P5 and P7; s2 slightly wider in P5

than in P6-7, anterior margin convex and unarmed

in P5, with 1 spine in P6, 2 spines in P7; s4 of 9

with long spine at anterodistal corner, sometimes

reduced or absent; s4 of cf with 2 shorter spines

on anterior margin; s5 of 9 always with long

spine at anterodistal corner; s5 of o" with smaller

spines at same position. Telson slightly shorter than

width at base, in 9 about %, in o* slightly less

than half of Up3 protopod.

DISTRIBUTION.—Warm parts of the Atlantic, Pacific, and Indian Oceans. In the CalCOFI area its

distribution is similar to that of the much more

abundant Lestrigonus schizogeneios, and it seems

likely that H. stebbingi is also a transition zone

species.

23. Hyperietta stephenseni, new species

FIGURES 46-48

DERIVATION OF NAME.—The species is named for

the distinguished Danish carcinologist, Knut

Stephensen.

TYPES.-AH from CalCOFI Cruise 5, station 801,

13 July 1949, off southern California, 33° 19^,

120°45'W, oblique tow from 71 m to surface.

Holotype, 9, USNM 137507, allotype d", USNM

137508, 3 9 and 2 immature d1 paratypes USNM

137509; 29 paratypes deposited at Scripps Institution of Oceanography.

DIAGNOSIS.—Length of 9 about 2 mm, of cf 3-3.5

mm. Head about twice as high as long, evenly

rounded ventrally. 9 Al reaching ventral margin

of head. Gland cone barely reaching ventral margin

of head, broadly rounded, separated from epistome

by short gap in lateral view. Lacinia of left Md

with about 7 teeth. Mxp like that of H. luzoni;

outer lobe about % longer than wide, bearing a

few spinules on both surfaces near distal margin.

S2 of PI about 2.2 times as long as wide, anterior

margin convex in proximal %, concave in distal

y$; s5 bearing a single spine at midlength of posterior margin and 3 at posterodistal corner; s6

about 3 times as long as wide, bearing a single

spine on anterior margin in 9> *~2 spines in cfS6 of P2 with a single spine on anterior margin in

9, 1-2 spines in cf • P6 very slightly longer than

P5 and P7; s2 slightly wider in P5 than in P6-7,

anterior margin convex and unarmed in P5, nearly

straight and bearing 1 spine in 9» 2 spines in cf

in P6-7; s4-5 of 9 with long spines at anterodistal

corners; s4-5 ot cf with shorter anterodistal spine

and additional short spines on anterior margins.

Telson slightly shorter than width at base, slightly

more than half length of Up3 protopod.

DISTRIBUTION.—Warmer parts of the Atlantic,

Pacific, and Indian Oceans. In the CalCOFI area

its distribution is similar to that of H. stebbingi

and the much more abundant Lestrigonus schizogeneios. It seems likely that H. stephenseni, like

the latter two species, inhabits the transition zone.
62

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 4$.—Hyperietta stebbingi, 9 (a-m, from off southern California): a, lateral view; b, head,

lateral: r. Al; d, rieht Md: e, Mxl; f, Mx2; z, Mxp, posterior; h, Mxp, lateral; i, PI; ;, P2; *,

P3; /, P4; m, urosome; n (from Arabian Sea), head, lateral.
NUMBER 136 63

FIGURE 44.—Hyperietta stebbingi, from off southern California (a-c, $) : a, P5; b, P6; c, P7; (d-t,

3 ) d, urosome, e, P4; /, PS; g, P5; /», P6; i, P7.

24. Hyperietta parviceps, new species

FIGURES 49-50

DERIVATION OF NAME.—From the Latin "parvi-"

[=small] + "-ceps" [=head], referring to the small

head compared to other species of Hyperietta.

TYPEs.-Holotype ? , USNM 137505, allotype rf.

USNM 137506, both from CalCOFI Cruise 9, station 1011, west of Punta Eugenia, Baja California,

27°56'N, 122°59'W. 3 paratypes (2$, ltf) from

the same sample have been deposited in the Scripps

Institution of Oceanography.

DIAGNOSIS.—Length of 9 2-2.5 mm, of cf 3-3.5

mm. Head and pereon broader than in other species

of Hyperietta; head about 1.6 times as broad as

long, about twice as high as long, in profile rather

flat below insertion of Al. Pereon quite convex

dorsally, highest in pereonite 5. Gland cone bluntly

rounded, clearly visible in lateral view, separated

from epistome by distinct gap. Mxp outer lobe

about 1.4 times as long as wide, with many fine setae

on anterior surface and a few heavier setae on posterior surface; lateral margin with single long seta. PI

subchelate; s2 slightly more than twice as long as

wide; s5 with small carpal process bearing 3 spines

and 1 spine on posterior margin; s6 with long spine

on anterior margin. P2 with carpal process about

Y4 as long as s6, bearing about 6 spines on its

margins. P3-7 with rather robust distal segments.

P5-7 with strong spine on anterodistal corner of

s4-5 of $ and s5 of cT- Telson about as long as

wide; in 9 %> in cf half as long as Up3 protopod.

DISTRIBUTION.—Thus far H. parviceps has been

found only at two stations of CalCOFI Cruise 5

and five stations of CalCOFI Cruise 9, all well offshore in the southern part of the area (Figure 50).

VI. ThemisteUa Bovallius, 1887b

DIAGNOSIS.—Small species with rather broad

pereon. Head rather broad; eyes occupying most of

its surface. Pereonites 1-5 fused in both sexes.

Coxae fused with pereonites. Telson very short.

9 Al 2-merous, rather long. $ A2 1-merous, rudimentary; gland cone small. Md with serrate incisor;

palp absent in 9. Mxl outer lobe with 4 terminal

spines plus 2 lateral and 1 medial subterminal

spines. Mxp outer lobes narrow, tapering distally;

inner lobe almost completely absent, apparently

represented by slight unarmed bulge on basal segment. Pl-7 with rather broad segments. Pl-2

chelate; gauge-shaped carpal process bearing marginal spines. P5 much longer than P6-7. Dactyls

of P6-7 with flexure slightly distal to midlength.

DERIVATION OF GENERIC NAME.—Diminutive of

Themisto (a Greek nymph, daughter of Neptune

and Doris); gender, feminine.
64

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 45.—Distribution of Hyperietta stebbingi on 4 CalCOFI cruises.
NUMBER 1S6 65

FIGURE 46.—Hyperietta stephenseni, from off California {a-c, $): a, head, lateral; b, Al; c,

urosome; (d-o,9) d, Al; f, A2; /, left Md; g, Mxl; fc, Mx2; i, Mxp, posterior; ;, Mxp, lateral;

k-n, PI—4; o, urosome.
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Hyperietta stephenseni, from off California (a-/,

?.): a-c, P5-7; df, dactyls of P5-7, medial; g-k, $ P3-7.

TYPE-SPECIES.—By monotypy, Themistella steenstrupi Bovallius, 1887b.

REMARKS.—Themistella was originally a monotypic genus, but in 1889 Bovallius added to it

Lestrigonus fuscus Dana (1853). The differences

cited by Bovallius for considering the two species

distinct are not convincing, since the descriptions

and illustrations of both authors lack detail, and

neither Dana nor Bovallius can be depended upon

for a high degree of accuracy. Hence I am treating

the two nominal species as synonyms.

In assigning the amphipod described below to

Themistella I have assumed two major inaccuracies

in the accounts of Dana and Bovallius. The first

assumption concerns the number of fused anterior

pereonites: for L. fuscus "thorax seven-jointed, first

segment nearly concealed" (Dana, 1853); for T.

steenstrupi "The first two peraeonal segments are

coalesced, the following are free" (Bovallius, 1889).

Bovallius' pi. IS: fig. 47, a lateral view of the male,

shows faint sutures separating pereonites 2-5 from

one another and heavier sutures on the margins of

pereonites 6-7. I suggest that Bovallius misinterpreted muscle bands or other structures as the

faint sutures and that his specimens actually had

pereonites 1-5 fused.

My second assumption is that Dana and Bovallius did not notice the angular bend in the dactyls

of P6-7. It is difficult to believe that this feature,

so conspicuous to me, could have been overlooked,

but a character that attracts the attention of one

author may escape the notice of another.

If the above assumptions are accepted, there is

no difficulty in equating the species described below as T. fusca with those proposed by Dana and

Bovallius. The assumptions may appear overly bold

and are to some degree intuitive, but they are based

on considerable experience and much frustration

in attempting to reconcile the accounts of hyperiid

species by the above authors with the characters

of specimens thought to belong to their nominal

species.

25. Themistella fusca (Dana)

FIGURE 51

Lestrigonus fuscus Dana, 1853:983, pi. 67: figs. 8a-c.-Bate,

1862:291-292, pi. 48: fig. 8 [copied from Dana].

Hyperiella fusca (Dana) .-Bovallius, 1887b:20.
NUMBER 1S6 67

FIGURE 48.—Distribution of Hyperietta stephemeni on 4 CalCOFI cruises.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 49.—Hyperietta parviceps, from off Baja California (a-o, $) : a, lateral view; b, Al; c,

right Md; d, left Md; e, Mxl; /, Mx2; g, Mxp; h-n, Pl-7; o, urosome; p-t, $ P5-7: u, $ urosome.
NUMBER 136 69

FIGURE 50.—Distribution of Hyperietta parviceps on 2 CalCOFI cruises.

Themistella steenstrupi Bovallius, 1887b:23; 1889:313-316, pi.

13: figs. 47-60.

Themistella fusca (Dana) .-Bovallius, 1889:316-317, figs. 1-3

[copied from Dana].

Hyperia thoracica Bovallius.—Vosseler, 1901:73-74, pi. 6: figs.

l^l.-Stephensen, 1924:91.

DERIVATION OF NAME.—From the Latin "fuscus"

[=dusky, dark], referring to the color of this

species.

TYPE-LOCALITY.—Tropical Atlantic, 1°S, 17°-

18°W.

DIAGNOSIS.—Since the genus is monotypic the

generic diagnosis serves as a specific diagnosis also.

RELATIONSHIPS.—The short telson of Dana's L.

fusca is characteristic of Hyperioides as well as of

Themistella, and the possibility that Dana's L.

fuscus was actually Hyperioides sibaginis must be

considered. In Dana's fig. 8a, P7 is nearly as long

as P6, whereas in H. sibaginis P7 is much shorter

than P6. The uropods of L. fuscus in Dana's fig.

8c are slender as in Themistella, and the rami are

much shorter than the protopods. In H. sibaginis

the uropods are broader and the rami are much

longer in proportion to the protopod lengths,

especially in Up 1-2.

The short telson and the structure of the uropods

in Vosseler's (1901) pi. 6: fig. 4 are characteristic

of Themistella and quite different from Bovallius'

illustrations of these structures in Hyperia thoracica, hence I have listed Vosseler's reference to if.

thoracica in the synonymy of Themistella. Stephensen (1924) also identified specimens of Themistella

as H. thoracica. I have examined his specimen from

the Thor Expeditions and found it to be a typical

Themistella. The specimen in the Copenhagen

Museum called H. Reinhardi by Bovallius (on the

label) and H. thoracica by Stephensen (1924) is

also a Themistella.

DISTRIBUTION.—I have examined specimens from

the tropical Atlantic (off Barbados; Gulf of

Guinea), the eastern Pacific (off the southern end

of Baja California, Mexico; off Guatemala; off

Nicaragua), and the Indian Ocean (Arabian Sea)

and have found them all very similar. Previous

records are from the tropical mid-Atlantic (Dana:

1°S, 17-18°W; Bovallius: 3°N, 25°W), North and

South Atlantic (Vosseler), and near Madeira
70

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 51.—Themistella fusca (a-tn, $ from off Cape San Lucas, Baja California): a, lateral

view; b, right Md; c, Mxl; d, Mx2; e, Mxp; /-/, Pl-7; m, urosome. (n-s,. $ from off Pacific

coast of Central America) n, Md. palp; o, Mxl palp; p, Mxp, lateral; q, Mxp, outer lobe,

posterior; r, P5 dactyl, lateral; s, urosome; t, 9 from Gulf of Guinea, head, lateral.
NUMBER 1S6 71

(Stephensen). The very limited records suggest a

pantropical distribution for T. fusca.

VII. Hyperionyx, new genus

DIAGNOSIS.—Small species. Head globular, with

eyes occupying most of its surface. Pereonites 1-3

fused in both sexes. Coxae fused with pereonites.

Telson very short. $ Al 2-merous. 9. A2 1-merous, slender, moderately long; gland cone small,

triangular. Md reduced [?]; palp absent in <j>.

Mxl outer lobe with 1 terminal spine; palp with

very few marginal setae. Mx2 with few setae; outer

lobe with 2 terminal spines; inner lobe very short.

Mxp with very few setae; inner lobe slender; outer

lobes obovate. PI subchelate. P2 chelate, with

spinose spoon-shaped carpal process. P3-7 with

strong, curved, unarmed dactyls; P3-4 and P6 subequal, P5 much shorter, P7 somewhat longer.

DERIVATION OF NAME.—Hyperia + the Greek

"onyx" [= claw], referring to the strongly developed dactyls of P3-7; gender masculine.

TYPE-SPECIES.—Hyperia macrodactyla Stephensen,

1924.

REMARKS.—The distinctive characters of H. macrodactyla warrant the establishment of a new genus

for it. The species is well described and illustrated

by Stephensen (1924) and Yang (1960) except for

the omission of descriptions of any of the mouthparts. The small size of H. macrodactylus (2-3

mm) together with the reduced mouthparts makes

dissection of the latter difficult. After removing the

posterior mouthparts it was apparent that I would

not be able to remove the Md without excessively

damaging the only Smithsonian specimen (1 of

Yang's 2 $) and I did not attempt to do so. Even

when the specimen was cleared I could not see

the incisor or molar and I believe they are quite

reduced.

Hyperionyx is the only genus of Hyperiidae in

which pereonites 1-3 are fused in both sexes. It

resembles Themistella in having a short telson and

a small gland cone and shares with Hyperioides a

well developed female A2, but in other respects it

is quite distinct from the latter two genera.

26. Hyperionyx macrodactylus (Stephensen)

FIGURE 52

Hyperia macrodactyla Stephensen, 1924:90-91, fig. 35.—Hurley,

1960b:279.-Vives, 1968:460.-Dick, 1970:56.

Hyperia (Parahyperia) macrodactyla Stephensen.—Yang, 1960:

35-38, fig. 9.

DERIVATION OF NAME.—Not expressly stated; obviously refers to the long dactyls of P3-7.

TYPE-LOCALITY.—Mediterranean, from south of

the Balearic Islands to the Sea of Marmora.

DIAGNOSIS.—With the characters of the genus.

RELATIONSHIPS.—This species, the sole representative of the genus, can be recognized immediately

by the relatively short P5, illustrated clearly by

Stephensen and by Yang. Supplementing their figures, illustrations are given here of the mouthparts,

the head, showing the small pointed gland cone

FIGURE 52.—Hyperionyx macrodactylus, $, from off Miami,

Florida: a, head, lateral, b, Mxl; c, Mx2; d, Mxp, posterior;

e, Mxp, lateral, f-h, dactyls of P5-7.
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

widely separate from the labrum, and the dactyls

of P5-7. Yang shows a 2-merous A2, but I could

not make out a suture in A2, possibly because I

examined it only in situ.

DISTRIBUTION.—Mediterranean (Stephensen, 1924,

34 specimens from 4 stations; Vives, 1968); off Miami, Florida, in Florida Current (Yang, 1960, 2 $);

South Atlantic, off South Africa (Dick, 1970); Fiji

Islands (Hurley, 1960, 1 o"). Presumably these

records represent only a small part of its range.

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