Seven Lubbockia Species

(Copepoda: Cyclopoida) from the

Plankton of the Northeast Pacific,

with a Review of the Genus

GAYLE A. HERON

and

DAVID M. DAMKAER

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 267
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SMITHSONIA N CONTRIBUTION S T O ZOOLOG Y • NUMBE R 26 7

Seven Lubbockia Species

(Gopepoda: Cyclopoida) from the

Plankton of the Northeast Pacific,

with a Review of the Genus

Gayle A. Heron

and David M. Damkaer

SMITHSONIAN INSTITUTION PRESS

City of Washington

1978
ABSTRAC T

Heron, Gayle A., and David M. Damkaer. Seven Lubbockia Species (Copepoda:

Cyclopoida) from the Plankton of the Northeast Pacific, with a Review of the

Genus. Smithsonian Contributions to Zoology, number 267, 36 pages, 22 figures,

2 tables, 1978.—Among the marine plankton cyclopoids, the oncaeids represent a

great number and wide variety of relatively little studied species. The family

Oncaeidae is defined in this report, and a key is given for its five genera. Of these,

the genus Lubbockia is defined in detail, and a key (Figure 2) is given for its 11

species. Relationships within the genus, as well as to other oncaeids, are discussed.

The bulk of the report is based on seven Lubbockia species collected from the

plankton of the northeast Pacific Ocean; most of these species are also widely

distributed elsewhere. Descriptions, illustrations, and notes on distributions are

provided for these seven species, four of which are new. Descriptive notes and

illustrations are also given for two other Lubbockia species known from tropical

and subtropical areas of the Pacific.

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

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

cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data.

Heron, Gayle A.

Seven Lubbockia species (Copepoda: Cyclopoida) from the plankton of the northeast Pacific,

with a review of the genus.

(Smithsonian contributions to zoology ; no. 267)

Bibliography: p.

1. Lubbockia—Classification. 2. Oncaeidae—Classification. 3. Crustacea—Classification. 4. Crustacea—North Pacific Ocean. I. Damkaer, David M., joint author. II. Title. III. Series:

Smithsonian Institution. Smithsonian contributions to zoology ; no. 267.

QL1.S54 no. 267 [QL444.C73] 591'.08s [595'.34J 77-16470
Contents

Page

Introduction 1

Material and Methods 1

Acknowledgments 2

Family ONCAEIDAE Giesbrecht, 1892 3

Key to the Genera of the Family Oncaeidae 3

Genus Lubbockia Claus, 1863 3

Lubbockia carinata, new species 5

Lubbockia forcipula, new species 11

Lubbockia wilsonae Heron and Damkaer, 1969 11

Lubbockia aculeata Giesbrecht, 1891 15

Lubbockia squillimana Claus, 1863 18

Lubbockia minuta Wolfenden, 1905 19

Lubbockia glacialis Sars, 1900 23

Lubbockia species 23

Lubbockia flemingi, new species 25

Lubbockia petersoni, new species 29

Appendix: Tables 33

Literature Cited 35

Seven Lubbockia Species

(Copepoda: Gyclopoida) from the

Plankton of the Northeast Pacific,

with a Review of the Genus

Gayle A. Heron

and David M. Damkaer

Introduction

Among the Poecilostoma (Copepoda: Cyclopoida), most attention has been given to the numerous families associated with benthic invertebrates

(see, for example, Gooding, 1963; Humes and Stock,

1973). The marine plankton poecilostomes, except

perhaps for the Corycaeidae, have by comparison

been little studied. The "planktonic" poecilostomes

(Corycaeidae, Sapphirinidae, and Oncaeidae) are

possibly not truly planktonic, but may be associated, at least at times, with larger plankton organisms (Sars, 1918). Sapphirina species, especially, are

known to associate as parasites and predators on

salps (A. C. Heron, 1973). Oncaea species have been

seen to graze on appendicularian houses (Alldredge,

1976). There is no doubt that Lubbockia species

would be mechanically able to hold other organisms

with their well developed maxillipeds, but their

actual food sources are not definitely known. The

broad specific differences in maxilliped construction suggest that food sources vary among Lubbockia species.

At present, 11 named species can be assigned to

Gayle A. Heron and David M. Damkaer, Pacific Marine

Environmental Laboratory, Environmental Research Laboratories, National Oceanic and Atmospheric Administration,

Seattle, Washington 98105.

the genus Lubbockia and are included in our key.

Notes on a North Atlantic species, L. extenuata,

were provided by G. Boxshall (personal communication), but the published report (Boxshall, 1977) was

not available before the present paper went to

press. Neither L. extenuata nor L. brevis, originally

described from off Ireland, were found in the

northeast Pacific, and they are not described in this

report. A description of both sexes of L. brevis, and

also of L. glacialis, is included in a report of Arctic

Oncaeidae to be published soon (Heron and

English, in prep.). Lubbockia squillimdna and L.

aculeata, from tropical and subtropical areas, also

were not found in our northeast Pacific samples,

but descriptive notes are given here for these species

as well as for the seven species (including four

which are new) that are known for the northeast

Pacific. Only the male of L. forcipula remains unknown.

Most Lubbockia species have a wide distribution,

and of those species found in the northeast Pacific,

we have also studied specimens from other areas: the

Antarctic (L. carinata, L. flemingi, L. forcipula, and

L. wilsonae; Tables 1, 2), the Atlantic (L. minuta;

Tables 1, 2), and the Arctic oceans (L. glacialis;

Heron and English, in prep.). Lubbockia petersoni

has been found only in the northeast Pacific.

MATERIAL AND METHODS.—Sources of the sped-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mens examined during this study are listed in

Table 1 and the species are itemized in Table 2.

Those specimens that were studied in detail are

enumerated under "Material Studied" in each

species presentation.

Type specimens or reference specimens of all

species described in this report have been deposited

in the National Museum of Natural History, Smithsonian Institution, Washington, D. C, where they

bear the catalog numbers of the former United

States National Museum (USNM).

The samples from NOAA Surveyor were collected

using a 60 cm diameter closing ringnet of 211 /tin

mesh, hauled vertically through discrete layers. The

Brown Bear and Thompson samples from Mr.

Willis K. Peterson were collected with a specially

designed plankton net, mesh aperture 110 jan,

which accompanied a deep water-bottle cast. Two

Clarke-Bumpus nets were sewed together with the

open cod end of the shorter one reduced to about

5 cm diameter and extended into the mouth of the

other; a small collecting bucket was attached to the

end of the outer net. A steel ring was sewed into

the mouth of the common opening, and fastened

by a short bridle to the hydrographic wire. Another

short bridle extended from the bucket, and this

bridle was fastened to the wire beneath the net.

Thus the sampler was held parallel to the hydrographic wire and it sampled only as it was retrieved. The net was attached beneath the deepest

water-bottle on a hydrographic cast (1800 m or

more). The maximum depth of the net sample was

estimated from the deepest thermometric depth.

The constricted cod end of the inner net and the

relatively great length of the entire device limited

the escape of plankton when an ascending haul was

stopped to remove water bottles.

The other Thompson samples were collected using a multiple net trawl described by Frost and

McCrone (1974).

Figures were drawn with the aid of a Wild M20

drawing tube. In the figure legends, the capital

letter following the explanation of each figure component refers to the particular scale to which the

component was drawn. These scales are illustrated

in Figure 1. Some Lubbockia species have a large

number of spinules, sensilla, and gland vents

(Fahrenbach, 1962), the latter associated with internal tubules. Only such external features that are

conspicious have been illustrated. Swimming legs

A

B

C

D

E

F

G

FIGURE 1.—Scales used in drawing figure components (each

scale = 0.1 mm).

are illustrated in anterior view.

Terminology used in describing the armature of

swimming legs is as follows: setae are represented

by arabic numerals, spines by roman numerals;

Si = inner border of segments, Se = outer border,

St = terminal border. In describing the armature of

antenna 1, arabic numerals used alone represent the

number of setae. For use of "uropod" and "pediger,"

see Bowman (1971, 1976).

ACKNOWLEDGMENTS.—We are indebted to and

thank the following persons for loans of specimens

or samples: Dr. Thomas E. Bowman, National

Museum of Natural History (NMNH), Smithsonian

Institution, Washington, D. C, the depository of

specimens bearing catalog numbers of the former

United States National Museum (USNM); Dr.

Frank D. Ferrari, Smithsonian Oceanographic Sorting Center (SOSC), Washington, D. C; and Dr.

Bruce W. Frost, Mr. Willis K. Peterson, and Mr.

David S. Thoreson, Department of Oceanography,

University of Washington (UW), Seattle, Washington.

We are grateful to Dr. Takasi Tokioka, Seto

Marine Biological Laboratory, for providing the

rare Marukawa (1927) publication, and also to Dr.

Shigeko Ooishi, Faculty of Fisheries, Mie University, for translating the relevant text.

We also appreciate the information kindly given

by Drs. T. Bjornberg, Universidade de Sao Paulo,

Brasil, and G. Boxshall, British Museum (Natural

History), concerning Lubbockia species studied by

them.

This study was supported in part by the Alaska

Outer Continental Shelf Environmental Assessment

Program, sponsored by the National Oceanic and
NUMBER 267

Atmospheric Administration (NOAA) and the U.S.

Department of Interior, Bureau of Land Management (BLM). This publication is Contribution

Number 950, Department of Oceanography, University of Washington WB-10, Seattle, Washington.

Family ONCAEIDAE Giesbrecht, 1892

Prosome and urosome divisions well defined, the

latter generally slender. Prosome, and urosome of

female, 5-segmented; urosome of male 6-segmented.

Prosome elongate to elongate-oval, or cyclopiform.

Antenna 1 short, with reduced number of segments.

Antenna 2 3-segmented, subprehensile to prehensile.

Labrum medially incised. Mandible complex, with

3-5 subterminal elements. Maxilla 1 small, bilobed.

Maxilliped a well developed claw in both sexes. Legs

1-4, exopods and endopods essentially 3-segmented.

Leg 5 a single free segment or represented by 1-3

setae.

Sars revised the family Oncaeidae and questioned

the inclusion of the genus Pachos (Claus, 1863). On

the basis of a female and a male specimen in the

genus Pachos that occurred in sample TA 45 (Table

1), it is apparent that the antenna 1 and the oral

appendages of this genus differ from any of the

other genera Sars included in the family Oncaeidae:

Conaea, Lubbockia, Oncaea, and Pseudolubbockia.

As a result of having examined specimens from

each of these genera and from the genus Epicalymma, recently described as an addition to the

Oncaeidae (G. A. Heron, 1977), we affirm Sars'

supposition and exclude Pachos from the Oncaeidae.

Humes and Stock (1973) stated that Oncaeidae

are rather closely related to the sabelliphilid and

lichomolgid poecilostomes, but that several important features establish their distinctions. It is likely

that Humes and Stock considered only the genus

Oncaea when they included among these principal

differing features a nonprehensile antenna 2 and

the form of the mandible. In our opinion, antenna

2 of Oncaeidae is at least subprehensile, grading

through Oncaea species toward a prehensile type,

found also in Epicalymma and Conaea species,

while in Pseudolubbockia and Lubbockia species,

especially in females, it is definitely prehensile. The

form of the mandible in Lubbockia species also

approaches the lichomolgoid type. We believe,

therefore, that Oncaeidae are even more closely

related to the Lichomolgoidea than has previously

been suggested.

Key to the Genera of the Family Oncaeidae

1. Legs 1-4 with inner coxal seta 2

Legs 1—4 without inner coxal seta 3

2. Mandible with narrow serrate tip; leg 5 a free segment armed with 4 elements

Pseudolubbockia Sars, 1909

Mandible terminating in a lash or lashed element; leg 5 absent (1 species) or a free segment

armed with 2 elements Lubbockia Claus, 1863

3. Uropod with conspicuous expansion on dorsal surface, surrounding insertion of dorsal seta .... 4

Uropod without expansion on dorsal surface Oncaea Philippi, 1843

4. Legs 1-3 third exopod segment with II, III, III outer spines ....Epicalymma G. A. Heron, 1977

Legs 1-3 third exopod segment with III, II, II outer spines Conaea Giesbrecht, 1891

Genus Lubbockia Claus, 1863

Prosome elongate to elongate-oval; urosome

slender. Antenna 1 7-segmented, with 1 or more

incomplete sutures on female, the third segment

with 2 spines; at least the 3 terminal segments always fused on male. Antenna 2 with elongate third

segment; female armature formula: 0, 1, 3 (subapical) f 3 (terminal) f 3 or 4 terminal claws.

Mandible blade with scalelike denticles on median

edge and terminating in a lash; 1 inner and 2

dorsal elements. Maxilla 1 a bilobed appendage,

lobes with 2, 3 or 3, 3 elements. Maxilla 2 with a

short setule and 3 (1 bifurcate) or 4 ornamented

elements. Oral appendages of male degenerate in

some species, maxilliped usually dimorphic.

Variable armature characteristics, as noted under

species descriptions, are: (1) inner edge of leg 1

basis, (2) outer edge of third exopod segment in

legs 1 and 2, and (3) inner edge of third endopod

segment in legs 3 and 4. Such characteristics appear

to reflect natural groups in the Oncaea-Epicalym-
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

[ A . Legs 1 and 2, third exopod segment, outer edge: II |

Leg 3, third endopod segment.

Inner edge: 2, I

Leg 4, third endopod segment.

Inner edge: 1, I

Leg 3, third endopod segment,

Inner edge: 2

Leg 4, third endopod segment,

Inner edge: 1

» Length sum of

anal segment

and uropod

shorter than

that of

genital

segment;

maxilliped

second

segment with

large keel

s " Leg 1 basis

without

Inner spine

« Length sum of

anal segment

and uropod

longer than

that of

genital

segment;

maxilliped

second

segment

without keel

* Leg 1 basis

with Inner

spine

Leg 6

with 1

element

« Maxilliped

second

segment

with teeth

i Leg 5 half

length of

genital

segment

« Maxilliped

second

segment

without

teeth

•f Leg S almost

as long as

genital

segment

s Maxilliped

firs t

segment

with tooth

* Anal segment

not concave

L. oarinata

new species

NE Pacific

« 1.40-1.53

* 1.41 mm

Antarctic

" 1.30-1.33 i

I. foraipula

new species

NE Pacific

s 1.03 im

Antarctic

« 1.02-1.06 i

L. wileonae

Heron i Damkaer,

1969

NE Pacific

» 2.64-2.98 ran

<f 1.96-2.08 ran

Antarctic

* 2.60 Ma

L. aculeata

Giesbrecht,

1891

Tropics

1 1.90-2.36

i 2.35 mi

» Maxilliped

first

segment

without

tooth

* Anal segment

lateral

margins

concave

L. aquillimana

Claus, 1863

Tropics

8 1.22-1.67 mn

* 1.75-2.09 mn

L. extenuata

Boxshail. 1977

NE Atlantic

• 1.72,1.74 im

i 1.34 mm

B. Legs 1 and 2, third exopod segment, outer edge: III

Leg 3, third endopod segment.

Inner edge: 2

Leg 4, third endopod segment,

Inner edge: 1

« Oviduct just

anterior to

midline of

genital

segment

" Uropod shorter

than any

urosome

segment

L. minuta

Wolfenden,

1905

NE Atlantic

» 1.30 mm

Off Florida

» 1.24 m

NE Pacific

« 1.29-1.50 i

* 2.00-2.21

» Oviduct just

anterior to

first 1/3

line of

genital

segment

' Uropod about

equal to

length of

each of 2

postgenital

segments

L. glaaialie

Sars, 1900

Arctic

' 1.60-2.45 m

" 1.72-1.92 ran

NE Pacific

» 1.58-1.71 ma

Leg 3, third endopod segment,

Inner edge: 3

Leg 4, third endopod segment.

Inner edge: 2

Leg 3, third endopod segment,

Inner edge: 2, I

Leg 4, third endopod segment,

Inner edge: 1, I

» Seta of leg 6

short. First

postgenital

segment

longer than

anal segment

* Uropod shorter

than genital

segment

L. flemingi

new species

NE Pacific

« 0.83-0.91

* 0.84-0.85

Antarctic

* 0.82 mi

s Seta of leg 6

about as long

as outer spine

on leg 5.

First

postgenital

segment about

same length as

anal segment

* Uropod length

equal to that

of genital

segment

L.

Fa

NE

s

brevie

-ran, 1908

Atlantic

0.85 mi

Arctic

s

a

0.88-1.00 m

0.84-0.94 mm

L. petersoni

new species

NE Pacific

» 0.95-1.06

* 0.85-1.03

FIGURE 2.—Diagrammatic key to the species of the genus Lubbockia.
NUMBER 267

ma-Conaea complex but probably no such groups

exist within Lubbockia species. The characteristics

enumerated above are among those used to prepare

a diagrammatic key to the species of Lubbockia

(Figure 2). The generic pattern of armature is as

follows, with a numeral representing a constant

value and an asterisk indicating a variable count.

All species, except Lubbockia flemingi, with hyaline papilla partially covering vent on outer edge

of leg 1 coxa. Lubbockia, as well as Oncaea, species

usually have small or conspicuous conical projections between 2 terminal spines on the endopods of

legs 2 and 3, and sometimes leg 4. These terminal

projections appear to support a vent, which is

Coxa Basis Endopod Exopod

Leg

1

2

3

4

Si

1

1

1

1

Si

•

0

0

0

Se

1

1

1

1

1

Si

1

1

1

1

2

Si

1

2

2

2

Si

4

3

•

•

3

St

I

II

II

II

Se

I

1

I

I

1

Se

I

I

I

I

Si

1

1

1

1

2

Se

I

I

I

I

Si

4

5

5

5

3

St

I

I

I

I

Se

•

•

II

II

present even when the projection is lacking. Often

a vent is also on the inner distal corner of the basis

of all swimming legs.

Leg 5 a free segment bearing 2 terminal elements,

or leg 5 absent and represented by 1 lateral seta.

Lubbockia is an interesting and complex genus,

far more variable than many other polytypic poecilostome genera. Specific differences include slight

variations in body shape, but major differences in

maxilliped and leg 5 structure, and armature and

relative length of endopods and exopods of legs 1-4.

The individual Lubbockia species show various

relationships to other oncaeid genera, depending on

the morphological features compared. The relationship of Lubbockia to Pseudolubbockia is seen to be

close when the comparison is based on L. carinata,

L. jorcipula, and L. petersoni.

With considerable foresight, Claus (1863) named

Lubbockia "for the deserving English investigator"

John Lubbock, afterward Sir John Lubbock, Lord

Avebury (1834-1913). It is difficult to select a

single area of endeavor to characterize Lubbock,

for he became preeminent in many fields and was

the author of a great number of scientific and

popular books. His earliest published papers dealt

with Copepoda (1853-1863), including specimens

collected by Darwin on H.M.S. Beagle. Although

much of Lubbock's efforts were in taxonomy, he

felt strongly that these investigations were not

ends in themselves, but only instruments for the

understanding of animal behavior. Lubbock was

the first to experiment on color vision in insects,

and he did extensive research on social-insect behavior. By profession a banker, Lubbock instigated

major alterations in English banking practices. He

was a Member of Parliament, where he introduced

far sighted legislation having a present day cast.

Among these proposals were: Open Spaces Act,

Ancient Monuments Acts, and Wild Birds Protection Act. Lubbock was the first to correctly attribute Stonehenge to the Bronze Age; his archeology texts, in which he coined the terms "paleolithic" and "neolithic," were standard for a generation. Lubbock's father, Sir John Lubbock, was

a physical astronomer, dealing especially with tidal

theory; both men were vice-presidents of the Royal

Society. Young Lubbock was a friend, neighbor,

and finally pallbearer of Charles Darwin, who said

that he relied on the opinions of three men only—

Hooker, Huxley, and Lubbock. Lubbock's name

is also perpetuated in some genera of insects, and

in some copepod species. There are several biographies of Lubbock, including a recent one by Pumphrey (1959).

Lubbockia carinata, new species

FIGURES 3-6

MATERIAL STUDIED (Tables 1, 2).—NE PACIFIC: 4 females,

1.40-1.53 mm, NP 4 et al. (including: holotype, 1.44 mm, NP

4, USNM 168487; 1 paratype, 1.41 mm, NP 12, USNM

168489); 1 male, allotype, 1.41 mm, NP 7, USNM 168488;

1 juvenile, stage IV, 1.16 mm, NP 32. ANTARCTIC OCEAN:

2 males, 1.30-1.33 mm, AA 41, 42.

FEMALE.—Average length 1.47 mm. Prosome
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 3.—Lubbockia carinata, new species, female: a, lateral (A); b, urosome, dorsal (C); c,

uropod, dorsal {E); d, cephalosome, ventral (maxilliped keel darkened) (£).
NUMBER 267

FIGURE 4.—Lubbockia carinata, new species, female: a, labrum, ventral (F); b, mandible, right

(G); c, maxilla 1, left (G); d, maxilla 2, right (G); e, maxilliped, right (E); /, leg 1 (E).
8

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figure 3a) about 1.6 times length of urosome, with

rounded posterior corners.

Pediger 5 (Figure 36) with faint transverse ridge

dorsoposteriorly. Genital segment with areas of external genital apparatus on anteriolateral third of

segment, each area with 3 setules; genital segment

and each postgenital segment with delicate spinules

on posteroventral margin. Uropod approximately

half the length of genital segment, with short spinules on ventral surface; dorsal, innermost, and

outer long terminal setae nearly as long as inner

long terminal seta; rounded laminal flap (Figure 3c)

extending over base of dorsal seta.

Rostrum (Figure 3d) with thickened posteroventral margin, truncate as shown in ventral view.

Antenna 1 (Figure 3d) 7-segmented, with rudimentary articulation between segments 3-4 and

5-6. Armature formula: 4, 13 + 1 spinule, 1+ 2

spines, 4, 4 + 1 esthete, 2+ 1 esthete, 7+ 1 esthete.

Antenna 2 (Figure 3d) 3-segmented, with incomplete suture at distal third of terminal segment.

Second segment with 1 seta. Third segment with

inner setules and 3 subapical setae; armed terminally with 3 setae and 4 claws, all short.

Labrum (Figure 4a) with free margin divided

into 2 deltoid posteroventral lobes; a number of

conspicuous gland vents on outer surface and rows

of setules inserted on under surface.

Mandible (Figure 46) with 2 outer elements ornamented with rows of setules; mandible blade with

row of graduated, scalelike denticles on the median

edge; apical lash slender and long; concave edge

of base with a hirsute element and rows of long

setules supported by 2 riblike processes.

Maxilla 1 (Figure Ac) flat, bilobed; 3 barbed elements on outer lobe and 1 barbed and 1 setose

element on inner lobe.

Maxilla 2 (Figure 4d) 2-segmented. First segment

with expanded base and outer row of setules. Second segment outer margin with a hyaline setule

and setose, stout seta; terminally bearing 2 setulose

elements; a constriction and sclerotization suggesting segmentation near the anterior base but posterior base fused; curved setulose element on inner

margin.

Maxilliped (Figure 4e) apparently 4-segmented;

complicated sclerotized pattern in area of segment

3. Second segment with a remarkable anterior keellike process, curving posteriorly beside an inner

pocket which serves as a sheath for the terminal

claw; segment 2 with short outer spinules.

Leg 1 (Figure 4/) without inner spine on basis,

third exopod segment with 2 short outer spines.

Leg 2 (Figure 5a) third exopod segment with 2

short outer spines. Leg 3 (Figure 56) inner edge of

third endopod segment with 2 setae and 1 spine.

Leg 4 (Figure 5c) inner edge of third endopod segment with 1 seta and 1 spine. Legs 2-4 endopods

terminate with a small vented conical projection

between spines.

Leg 5 (Figure 36) with short free segment; 2

terminal elements, the longer tapering in a way

which suggests there is a fine serrate, hyaline flange;

length less than twice that of the outer. Seta on

body near leg.

Leg 6 probably represented by 3 short, hyaline

setules in area of external genital apparatus.

MALE.—Average length 1.35 mm. Prosome (Figure 6a) about 1.7 times the length of urosome.

Urosome (Figure 66) with the 4 postgenital segments bearing spinules on posteroventral margins.

Uropodal setae with relative lengths as in female.

Rostral area and mouthparts, except antenna 1

and maxilliped, as in female.

Antenna 1 with segments corresponding to terminal 3 of female, fused in male; armature similar

to corresponding segments of female.

Maxilliped (Figure 6c) with segment 2 and claw

elongate compared to female; 2 short segments

proximal to claw.

Swimming legs and leg 5 resemble those of

female.

Leg 6 (Figure 66) probably represented by posteroventral pointed flap on genital segment.

STAGE IV.—Length 1.16 mm, with 4 segments

and uropod in urosome; leg 5 small and segmented

(Figure 6d). Maxilliped with keel-like process well

developed.

ETYMOLOGY.—The specific name carinata, from

the Latin carinatus (keeled) refers to the keel-like

process on the maxilliped of the female.

REMARKS.—Lubbockia carinata superficially resembles L. forcipula, but differs from it by the relatively shorter uropods, distinctively keeled female

maxilliped and by the lack of an inner spine on

leg 1 basis. Lubbockia carinata, L. glacialis, and

L. minutus are the only Lubbockia species without

a spine on leg 1 basis.

Lubbockia carinata also resembles L. forcipula
NUMBER 267

FIGURE 5.—Lubbockia carinata, new species, female: a, leg 2 (E); b, leg 3 (E); c, leg 4 (E).

in having a seta on body near leg 5. Lubbockia

petersoni has such a seta but lacks leg 5. These are

the only known Lubbockia species with this seta,

which is probably a remnant of leg 5 basis. A similarly placed seta is characteristic of the other genera

of Oncaeidae: Pseudolubbockia, Oncaea, Epicalymma, and Conaea.

Lubbockia carinata and L. forcipula have armature of female antenna 1 resembling that of Pseudolubbockia dilatata Sars, 1909, except for having 2

additional setae and 1 spinule on segment 2. These

3 Oncaeidae species also differ from all other Lubbockia females by having an additional seta on segments 3 and 5, and 7 setae and 1 esthete on the

terminal segment instead of the usual 5 setae and 2

esthetes.

We have restudied Pseudolubbockia dilatata for

comparison with Lubbockia species and have modi-
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 6.—Lubbockia carinata, new species, male: a, lateral (A); b, urosome, ventral (C); c,

maxilliped, left (£). Stage IV: d, ventral (maxilliped keel darkened; antenna 1, except basal

segment, and swimming leg rami omitted) (A).

fied our diagnosis concerning details of P. dilatata

antenna 1. Antenna 1, 7-segmented, with an incomplete suture on inner surface between 5-6.

Armature formula: 4, 11, 1 + 2 spines, 4, 4 + 1

esthete, 2 + 1 esthete, 7+ 1 esthete. The esthetes

are relatively short, narrow, and each is associated

with a short seta, on proximal third of segment 5

and distal margins of segments 6 and 7. The short

seta on the posterior margin of segment 5 is an

error in figure 2, Heron and Damkaer, 1969.

The maxillipeds of several species of caligoid

copepods have been described with a prominent

second segment keel or similar process associated

with the tip of the terminal claw, as in L. carinata:

Dysgamus atlanticus Steenstrup and Lutken, 1861

(pi. 4: fig. 8), Dysgamus pacificus Wilson, 1950 (pi.

6: fig. 55), and Lewis (1968, fig. 8d) for Dissonus

heronensis Kabata, 1966. Lewis (pers. comm.) has

stated that the legends for his figure %d,e were inadvertently transposed and that figure Sd is the

maxilliped of female D. heronensis.

The long, resilient uropodal setae of L. carinata

and L. forcipula resemble the uropodal setae in

Conaea and Epicalymma species, but differ from

those of Oncaea, Pseudolubbockia, and the other

Lubbockia species.

DISTRIBUTION.—Northeast Pacific and Antarctic

Ocean (Tables 1, 2).
NUMBER 267

11

FIGURE 7.—Lubbockia forcipula, new species, female: a, dorsal (C); b, lateral (C); c, cephalosome,

ventral (E); d, antenna 2, right (E); e, labrum, ventral (G); /, mandible, left (G); g, maxilla 1,

right (G); h, maxilla 2, right (G); i, maxilliped, right (G).
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lubbockia forcipula, new species

FIGURES 7-8

MATERIAL STUDIED (Tables 1, 2).—NE PACIFIC: 1 female,

holotype, 1.03 mm, NP 11, USNM 168492. ANTARCTIC

OCEAN: 2 females, 1.08, 1.12 mm, A A 42.

FEMALE.—Length about 1.08 mm. Prosome (Figure 7a,b) 1.2 times the length of urosome.

Pediger 5 with transverse sclerotized ridge dorsoposteriorly. Genital segment with areas of external

genital apparatus on anteriolateral third of segment; each area with a setule. Genital segment and

each postgenital segment with delicate minute spinules on posteroventral margin. Uropod about z/3

length of genital segment, posteriorly dilated over

base of dorsal seta; dorsal, innermost, and outer

long terminal setae nearly as long as inner long

terminal seta.

FIGURE 8.—Lubbockia forcipula, new species, female: a, leg 1 (E); b, leg 2 (E); c, leg 3 (E);

d, leg 4 (E).
NUMBER 267

13

Rostrum with thickened posteroventral margin

slightly depressed and indented as shown in ventral

view of cephalosome (Figure 7c).

Antenna 1 (Figure 7c) and antenna 2 (Figure

lc,d) with armature similar in number to that of

L. carinata.

Labrum (Figure le) with free margin divided

into 2 posteroventral lobes, each with a subapical

cluster of setules; complex pattern of layers of tissues with hyaline setules inserted on under surfaces.

Mandible (Figure 7/), maxilla 1 (Figure 7g), and

maxilla 2 (Figure Ik) with armature similar in

number to that of L. carinata.

Maxilliped (Figure 7t) 4-segmented, cheliform.

Second segment beaklike, projecting beyond and

somewhat enclosing the tip of short terminal claw,

the complex forming a short, strong pincer.

Legs 1-4 (Figure Sa-d) with armature similar in

number to that of L. carinata except for the addition of an inner spine on leg 1 basis.

Leg 5 (Figure la) similar in general form to that

of L. carinata, including seta on body near leg.

Leg 6 probably represented by short, hyaline

setule in area of external genital apparatus.

MALE.—Unknown.

ETYMOLOGY.—The specific name, from the Latin

forcipula (nippers, tongs), refers to the appearance

of the maxilliped in the female.

REMARKS.—The truncate shape of the L. forcipula rostrum resembles L. carinata, but the depressed and indented posterior margin is similar

to that of Pseudolubbockia dilatata.

Of the known species of Lubbockia, only L. forcipula, L. carinata, and L. petersoni have an inner

spine on legs 3 and 4 third endopod segment; L.

carinata and L. forcipula differ from L. petersoni

by having 2 (rather than 3) outer spines on legs

1 and 2 third exopod segment. Lubbockia forcipula

and L. carinata also differ from other Lubbockia

species by having 13 setae (rather than 7) on

antenna 1 second segment.

The unusual maxilliped of L. forcipula, having

a short, hooded terminal claw, separates it from L.

carinata, which it otherwise resembles. The maxilliped of L. forcipula resembles that of Phyllotherus

cornutus (Milne-Edwards, 1840), a caligoid copepod

parasitic on sharks (see Cressey, 1967, fig. 130).

DISTRIBUTION.—Northeast Pacific and Antarctic

Ocean (Tables 1, 2).

Lubbockia wilsonae Heron and Damkaer, 1969

FIGURE 9

Lubbockia aculeata.—Vervoon, 1951:151; 1957:148. [Not L.

aculeata Giesbrecht, 1891.]

Lubbockia wilsonae Heron and Damkaer, 1969:9-15, figs. 11-

19.

The original description of Lubbockia wilsonae

was based on 2 females (2.70, 2.75 mm) from the

Gulf of Alaska. Additional specimens of both sexes

from the northeast Pacific have made it possible to

describe the male and amend the original description of the female.

MATERIAL STUDIED (Tables 1, 2)—NE PACIFIC; Prince

William Sound, Alaska: 3 females, 2.80-2.98 mm, NP 1. Near

the Canadian weatherstrip sta P: 6 females, 2.64-2.88 mm,

NP 38, 39; 3 males, 1.96-2.08 mm, NP 38, USNM 168503,

168504; 8 juveniles, stage V, NP 38, 39; 2 juveniles, stage IV,

both 1.48 mm, NP 39; 2 juveniles, stage III, 0.97, 1.06 mm,

NP 7, 4 (frequency of sex and stage categories by depth for

samples NP 38, 39 shown in accompanying tabulation).

ANTARCTIC OCEAN: 3 males, 2.42-2.44 mm, AA 42.

6 Aug 7 Aug

m

165-110

220-165

275-220

330-275

385-330

440-385

$

1

1

1

$

1

2

V

1

3

1

$

2

1

V

1

2

IV

1

1

FEMALE.—Rostrum (Figure 9a) rounded posteriorly. Antenna 1 in all Lubbockia species appears

to be fundamentally 7-segmented; segment 3 often

with incomplete or no suture separating 4. Female

Lubbockia specimens have been described as having

from 4 or 5 to 7 segments in antenna 1; some sutures are incomplete on inner surface, while articulation is usually more distinct on outer surface.

Lubbockia wilsonae female antenna 1 is 7-segmented (rather than 6) with segments 3-4 partially

or completely fused. Armature should be corrected

from original description to 4, 6 + 1 esthete + 1

spinule, 2 spines, 4, 3 + 1 esthete, 2+ 1 esthete,

5 + 2 esthetes (1 seta, about equal to combined

length of segments 5-7, in place of 2 short setae

shown on figure 14 of Heron and Damkaer, 1969).

Maxilla 1 (Figure 96) bilobed; inner lobe with 2

elements and short hyaline setules; 3 elements on

larger lobe.

MALE.—Length about 2.01 mm (NE Pacific) to
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 9.—Lubbockia wihonae Heron and Damkaer, female: a, rostrum, ventral (E); b, maxilla

1, right (F). Male: c, lateral (A); d, dorsal (A); e, first 3 urosomal segments, ventral (D); /,

rostrum, ventral (E); g, antenna 1 distal esthete bases x, y, and z (G); h, antenna 2, left (E);

i, maxilliped, right (E). Stage V, female: ;, urosome, ventral (A). Stage V, male: k, urosome,

ventral (A). Stage III: /, ventral (antenna 1, except basal segment, and swimming leg rami

omitted) (A).
NUMBER 267 15

2.43 mm (Antarctic). Body (Figure 9c,d) slender,

with prosome to urosome ratio and relative lengths

of urosomal segments similar to female. Prosome

with conspicuous sensilla, gland vents, and minute

spinules. Urosomal segments covered with minute

spinules, somewhat larger on most ventral surfaces;

posteroventral margins with spinule patterns similar

to those of female. Genital segment (Figure 9e) with

conspicuous spinules on ventral surface. Relative

lengths of uropodal setae as in female.

Rostrum (Figure 9/) more tapered posteroventrally than in female.

Antenna 1 distinctly segmented only between

segments 4 and 5 and outer surface of 1 and 2. Armature similar in number to that of female except for

additions of esthete on segment 2 and 2 esthetes on

segment 4. All esthetes, except 1 each on segments

corresponding to segments 2 and 7 (labeled "x" in

Figure 9g) longer and of a different structure than

those of female. Terminal esthete (labeled "y" in

Figure 9g) with broad base and wide setalike rib

supporting entire length; nearly reaches urosome

segment 3. The 5 other esthetes extend from a short

base (labeled "z" in Figure 9g) and also have a supporting rib.

Antenna 2 (Figure 9h) third segment sexually dimorphic; 3 terminal claws of female modified to 3

short setae, and 1 seta modified to long lash; 2 clusters of long setules on inner surface.

Labrum with hood extending to cover vertex between the 2 posteroventral lobes.

Mandible, maxilla 1, and maxilla 2 degenerate,

variable between specimens and right and left.

Maxilliped (Figure 9i) with outer surfaces of segments 1 and 2 rugose; second segment with inner

row of setules; cluster of long, delicate setules on

inner distal surface; terminal claw undulant.

Swimming legs and leg 5 as in female.

Leg 6 probably represented by posteroventral flap

on genital segment, bearing 2 elements.

STAGE V.—Female (Figure 9;") 1.98 mm, male (Figure 9k) 1.75 mm, with 5 segments and uropod in

urosome; leg 5 well developed. Male with a faint

sclerotized ventral line on genital segment in area

of adult anterior margin of posteroventral flap.

STAGE IV.—Length 1.48 mm, with 4 segments and

uropods in urosome; leg 5 small with 2 spines; inner

spine reaching beyond posterior edge of next segment.

STAGE III.—Two specimens, 0.97, 1.06 mm, with

5 segments in the prosome and 3 segments and

uropods in urosome (Figure 91); leg 5 represented by

1 spine. Antenna 2 with 6 elements similar to that

in adult female. Maxilliped with relative size similar to that in adult female, 2 small triangular

spines on second segment and small hyaline spinules

on terminal claw.

REMARKS.—The 3 L. wilsonae males from the

Antarctic differ from the northeast Pacific specimens by their larger size and greater number of

spinules on the urosome. All other morphological

characters appear to be similar.

Among the Lubbockia species with 2 outer spines

on legs 1 and 2 third exopod segment, only L. wilsonae has 2 elements representing leg 6.

DISTRIBUTION.—Northeast Pacific and Antarctic

Ocean (Tables 1, 2; Heron and Damkaer, 1969).

Lubbockia aculeata Giesbrecht, 1891

FIGURES 10-11

Lubbockia aculeata Giesbrecht, 1891:477; 1892:606-611, pi. 48:

figs. 3,9,11, 13,16,20.

Giesbrecht described Lubbockia aculeata from

specimens collected in the tropical eastern Pacific

Ocean (female, 2.30 mm; male, 2.35 mm).

Our specimens agree with the characters which

Giesbrecht outlined for the species. Additional

species in the genus now require more detailed

comparisons, so we include figures of several features not emphasized by Giesbrecht.

MATERIAL STUDIED (Tables 1, 2).—ATLANTIC OCEAN, off

Bahamas: 2 females, 1.90, 213 mm, TA 44. Near Florida: 2

females, 210, 236 mm, TA 45. PACIFIC OCEAN, near

Philippine Islands: 1 female, 2.05 mm, TP 43.

FEMALE.—Average length about 2.09 mm. Body

and all appendages with unusually thick exoskeleton. Rostrum with complex pattern of sclerotization and pointed posteroventral margin as shown

in ventral view of cephalosome (Figure 10a).

Antenna 1 as in L. wilsonae.

Antenna 2 (Figure 106) and mandible {Figure

10c) with armature similar in number to that of

L. carinata.

Maxilliped similar to L. squillimana and L. wilsonae, with large dentiform processes on second

segment, but L. aculeata with 1 large dentiform

process on first segment (Figure lla). First segment
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 10.—Lubbockia aculeata Giesbrecht, female: a, cephalosome and somite of leg 1, ventral

(C); b, antenna 2, left (E); c, mandible, left (G).

may have small spinules in addition to dentiform

process (Figure 10a).

Leg 1 (Figure 116) with inner spine on basis,

third exopod segment with 2 short outer spines.

Leg 2 (Figure lie) third exopod segment with 2

short outer spines. Leg 3 (Figure lid) inner edge

of third endopod segment with 2 setae. Leg 4 (Figure 1 le) inner edge of third endopod segment with

1 seta. Legs 2-4 endopods terminate with a vent

between spines.

REMARKS.—Fryer (1957:14) stated that the swimming legs of certain freshwater cyclopoids are always directed forward while feeding, but that the

legs play no part in the actual feeding mechanism.

When legs are directed either posteriorly or anteriorly, the usual cases in preserved specimens, the

direction of the inner spine on the basis is not

clearly apparent (see spine darkened on Figure 10a).

However, legs of some specimens were preserved in

intermediate positions, and the direction of this

spine was obvious (see spine darkened on Figure

lla); it appears that this spine would be useful in

holding food. Three of the 11 named Lubbockia

species (4 of 12 including stages IV and V of "Lubbockia species") lack this spine: L. carinata, L.

glacialis, and L. minuta.
NUMBER 267

17

FIGURE 11.—Lubbockia aculeata Giesbrecht, female: a, maxilliped basal segment and leg 1,

ventral (C); b, leg 1 (D); c, leg 2 (D); d, leg 3 (D); e, leg 4 (D).
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

DISTRIBUTION.—Lubbockia aculeata has a definite

tropical distribution, although a few specimens have

been collected just outside tropical waters. Records

and summaries are from the Pacific Ocean (Tables

1, 2; Giesbrecht, 1891; Farran, 1936; Bjornberg,

1973); Indian Ocean and Indo-Pacific (Sewell,

1947); Atlantic Ocean (Tables 1, 2; Cleve, 1904;

Farran, 1926; Owre and Foyo, 1967; Vives, 1972);

and the Mediterranean Sea (Razouls, 1974).

Lubbockia squillimana Claus, 1863

Figure 12

Lubbockia squillimana Claus, 1863:164, 165, pi. 25: figs. 1-5.—

Giesbrecht, 1892:606-611, pi. 4: fig. 6, pi. 48: figs. 1, 2, 4-8

17-19,21.

Lubbockia minuta.— Marukawa, 1927:1237, fig. 2384. [Not L.

minuta Wolfenden, 1905.]

Lubbockia marukawai Mori, 1937:122, 123, pi. 67: figs. 10-13.

Specimens that we studied from different areas

varied as little as did ones from within the same

sample, and all specimens agreed with Giesbrecht's

(1892) excellent description. We have supplemented

this description of L. squillimana, because of the

additional Lubbockia species that have been added

since Giesbrecht's study.

MATERIAL STUDIED (Tables 1, 2)—MEDITERRANEAN

SEA, Gulf of Naples: 20 females, 1.29-1.67 mm, M 47. USNM

FIGURE 12.—Lubbockia squillimana Claus, female: a, rostrum, ventral (E); b, rostrum, lateral (E);

c, labrum, ventral (G); d, maxilla 2, right (G); e, leg 2 (E); /, leg 3 (E).
NUMBER 267

19

168505; 3 males, 1.98-2.09 mm, M 47, USNM 168505. Western

Mediterranean: 3 females, 1.33-1.48 mm, M 46. ATLANTIC

OCEAN, near Florida: 82 females, 1.22-144 mm, TA 45;

3 males, 1.75-1.86 mm, TA 45, USNM 168506. Off Bahamas:

1 damaged male, TA 44.

FEMALE.—Body and all appendages with unusually thick exoskeleton, with complex sclerotized

pattern in some areas (Figure I2a,b).

Antenna 1 as in L. wilsonae.

Labrum (Figure 12c) with free margin divided

into 2 posteroventral lobes, each margin with a row

of processes on under surface; short medial hood

over apex.

Maxilla 2 (Figure 12d) with armature similar in

number to that of L. carinata.

Legs 1-4 armature similar to that of L. aculeata;

legs 2 and 3 (Figure I2e,f) and leg 4 endopods terminate with a vent between spines.

DISTRIBUTION.—Lubbockia squillimana, like L.

aculeata, has a definite tropical distribution, although a few specimens have been collected just

outside tropical waters. Principal records and summaries are from the Pacific Ocean (Giesbrecht, 1891;

Farran, 1936; Mori, 1937, 1942; Wilson, 1950); Indian Ocean and Indo-Pacific (Scott, 1909; Krishnaswamy, 1953); Atlantic Ocean (Giesbrecht, 1892;

Marques, 1958; Owre and Foyo, 1967; Vives, 1972);

and the Mediterranean Sea (Razouls, 1974).

Lubbockia minuta Wolfenden, 1905

FIGURES 13-15

Lubbockia minuta Wolfenden, 1905:15, 16.

Lubbockia glacialis.—Heron and Damkaer, 1969:16-18, figs.

20, 21.—Boxshall, 1977:118, fig. 8.—Deevey and Brooks, 1977:

287. [Not L. glacialis Sars, 1900.]

Wolfenden collected 1 female (1.30 mm) off the

west coast of Ireland, which he described without

illustrations. The copepod collection of Richard

Norris Wolfenden, M.D. (1856-1923) was recently

made available to the authors by his heirs. While a

number of copepod type specimens have been found

in this collection, we have unfortunately not located

the Lubbockia minuta. Some L. squillimana were

found (see above), and also an empty slide labeled

"Lubbockia." The Wolfenden Collection will be

cataloged and deposited in NMNH.

MATERIAL STUDIED (Tables 1, 2).—NE PACIFIC: 32 females,

1.29-150 mm, NP 2 et al., USNM 168508-168512; 2 males,

2.00, 2.03 mm, NP 9, 14, USNM 168507; 5 juveniles, stage V,

1.03-1.10 mm, NP 9 et al.; 8 juveniles, stage IV, 0.89-0.97

mm, NP 9 et al. ATLANTIC OCEAN, near Florida: 1

female, 1.24 mm, TA 45.

FEMALE.—Average length about 1.41 mm. Prosome of equal or slightly greater length than

urosome (Figure 13a). Urosome (Figure 136,c) with

delicate hyaline spinules on posteroventral margin

of postgenital segments. Genital segment length

twice that of uropod; area of external genital apparatus located just anterior to midlength of segment, each area with a setule. Second postgenital

segment usually about y4 length of genital segment,

occasionally as long as genital segment

Rostrum rounded posteriorly as shown in ventral

view of cephalosome (Figure 13d).

Antenna 1 (Figure 13a,d) 7-segmented with variable rudimentary articulations between segments

3^1 and 5-6. Armature formula: 4, 7 + 1 spinule,

2 spines, 4, 3 + 1 esthete, 2 + 1 esthete, 5 + 2

esthetes; esthetes elongate.

Antenna 2 (Figure 13d) with 1 seta on second

segment. Third segment with inner setules and 3

subapical setae; armed terminally with 4 short claws

and 3 short setae, 1 nude, 1 setose, and 1 lashlike.

Labrum (Figure 13e) with free margin divided

into 2 rounded posteroventral lobes, each margin

with a row of short setules on under surface.

Mandible (Figure 13/) with stout, setulose outer

element; mandible blade with row of size-graduated

scalelike denticles and lashlike element adjacent to

blade; concave inner edge of mandible base with a

setulose element and row of long setules.

Maxilla 1 (Figure 13g) flat, bilobed; 3 elements

on each lobe.

Maxilla 2 (Figure 13A) with expanded first segment. Second segment outer margin with a hyaline

setule and stout setulose element; terminates in

element adjacent to setulose seta; curved setulose

element on inner margin.

Maxilliped (Figure 13i) 4-segmented. Second segment with inner cluster of short, delicate hyaline

setules. Terminal segment a sharply pointed claw.

Leg 1 (Figure 14a) without inner spine on basis,

third exopod segment with 3 short outer spines.

Leg 2 (Figure 146) third exopod segment with 3

short outer spines. Leg 3 (Figure 14c) inner edge

of third endopod segment with 2 setae. Leg 4 (Figure 15a) inner edge of third endopod segment with

1 seta. Legs 2-4 endopods terminate with a small

vented projection between spines.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 13.—Lubbockia minuta Wolfenden, female: a, lateral (A); b, urosome, dorsal (C); c,

postgenital segments and uropods, ventral (C); d, cephalosome, ventral (E); e, labrum, ventral

(G); /, mandible, right (G); g, maxilla 1, left (G); h, maxilla 2, left (G); i, maxilliped, right (E).
NUMBER 267

21

FIGURE 14.—Lubbockia minuta Wolfenden, female: a, leg 1 (E); b, leg 2 (E); c, leg 3 (E).

Leg 5 (Figure 156) with slender free segment not

extending to area of external genital apparatus; 2

terminal spines with narrow, delicate hyaline flange.

Leg 6 probably represented by small, hyaline

setule in area of external genital apparatus.

MALE.—Urosome about 1.3 times length of prosome (Figure 15c). Solophenyl blue 2RL stain

(English and Heron, 1976) revealed minute hyaline

setules on ventral posterior margins of 3 postgenital

segments. Third postgenital segment shows sexual

dimorphism in being i/g longer than preceding segment rather than only slightly longer as in female.

Rostrum as in female.

Antenna 1 with a single distinct articulation between segments 2, 3 of female; a partial line separates segments 1, 2; armature similar to corresponding segments of female, except for addition of 1

esthete on segment 2 and 2 esthetes on segment 4.

These additional esthetes and esthete on segment 6

elongate, supported by setalike rib; a terminal

esthete reaches fifth urosomal segment.

Antenna 2 third segment with the 4 terminal

claws of the female modified into 4 setae.

Mandible, maxilla 1, and maxilla 2 degenerate.

Maxilliped 3-segmented, without short segment

proximal to claw of female.

Swimming legs as in female.

Leg 5 with relative length of spines differing

slightly from female.

Leg 6 probably represented by posteroventral flap
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 15.—Lubbockia minuta Wolfenden, female: a, leg 4 (E); b, last prosomal segment and

first 2 urosomal segments, lateral (E). Male: c, ventral (A). Stage V: d, dorsal (B). Stage IV: e,

dorsal (B).
NUMBER 267 23

on genital segment, bearing a small pointed posterior process.

STAGE V.—Average length of 5 specimens 1.06 mm

(1.03-1.10 mm). These specimens resemble L. glacialis, but are slightly smaller and more slender, and

have shorter uropods in relation to the preceding

(anal) segment and urosome (Figure \5d).

STAGE IV.—Average length of 8 specimens

0.95 mm (0.89-0.97 mm). The same criteria used to

separate stage V were used to distinguish this stage

(Figure \5e).

REMARKS.—A single damaged male of Lubbockia

minuta was reported earlier as L. glacialis from the

Gulf of Alaska (Heron and Damkaer, 1969); the size

and swimming legs of that male resembled those of

female L. glacialis (male unknown at that time).

Since the females of L. minuta have now been recognized with males in northeast Pacific samples, it is

believed that these males are sexually dimorphic

males of L. minuta. Lubbockia minuta closely resembles L. glacialis, including the sexually dimorphic characters. A forthcoming study of Arctic

Ocean Oncaeidae includes the description of the

male L. glacialis, 1.72-1.92 mm x = 1.82 mm).

Lubbockia minuta has a shorter leg 5 and more

distal female genital operculum than L. glacialis.

The inner spine of L. glacialis male leg 5 reaches

the middle of the second postgenital segment.

The female specimen of L. minuta found near

Florida is slightly smaller (1.24 mm) than the smallest northeast Pacific female (1.29 mm), but appears

to be similar except for more distinct segmentation

between segments 3, 4 on antenna 1.

DISTRIBUTION.—NE Atlantic (Wolfenden, 1905)

and tropical Atlantic Ocean (Tables 1, 2); NE

Pacific Ocean (Tables 1, 2).

Lubbockia glacialis Sars, 1900

FIGURE 16

Lubbockia glacialis Sars, 1900:114-118, pi. 33: figs. 1-15.

MATERIAL STUDIED (Tables 1, 2).—NE PACIFIC: 6 females,

1.58-1.71 mm (x = 1.63 mm), NP 6 et al., USNM 168513-

168515; 1 juvenile, stage V, 1.20 mm, NP 4; 1 juvenile, stage

IV, 1.03 mm, NP 27.

REMARKS.—The specimens from the northeast

Pacific enumerated above were so much smaller

than those which Sars described (2.45 mm) from the

Norwegian North Polar Expedition that it took a

considerable amount of study to conclude that

these are the same species. Seventy-five L. glacialis

females from the Arctic Ocean (which are not listed

in Tables 1, 2 but are to be discussed in a forthcoming report, Heron and English, in prep.),

measured 1.60-1.92 mm (x = 1.72 mm).

The relative lengths of the 2 postgenital segments of L. glacialis females are slightly variable.

The second postgenital segment of the northeast

Pacific specimens is slightly longer than the first

postgenital segment, while these 2 segments are

about the same length in Arctic specimens. The

prosome appeared larger in relation to the urosome

on Arctic specimens, but this could be an effect of

their diet. The intestinal structures of all the Arctic

females and juveniles appear to be bulging with a

material which is conspicuously darker than the

exoskeleton, even in unstained specimens. The expanded prosome, variable in extent and occasionally

asymmetrical, may be an effect of the gut content.

The Pacific specimens resemble the Arctic specimens in all other characters.

Lubbockia glacialis females resemble L. minuta

in many characters, but differ in having the external

genital apparatus on the anterior third of the genital segment, and in having a longer uropod in relation to the anal segment (Figure \6a,b).

STAGE V.—One specimen, 1.20 mm, NP 4, differed

from the very similar L. minuta by the slightly

larger size and longer uropodal length relative to

the preceding segment (Figure 16c).

STAGE IV.—One specimen, 1.03 mm, NP 27, was

separated from the similar specimens of L. minuta

on the same criteria as those used for stage V

(Figure 16d).

DISTRIBUTION.—Arctic Ocean (Sars, 1900; Grice,

1962; Johnson, 1963; Minoda, 1967; Heron and

English, in prep.) and the North Pacific Ocean

(Tables 1, 2; Minoda, 1971). Hardy and Gunther

(1935) reported L. glacialis from the Antarctic, but

with some doubt.

Lubbockia species

FIGURE 17

STAGE V.—Seven specimens, 1.37-1.44 mm, (x =

1.40 mm) from the northeast Pacific (Tables 1, 2).

Body proportions (Figure 17a) and appendages are

similar to those in L. minuta and L. glacialis. Areas
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 16.—Lubbockia glacialis Sars, female: a, urosome, dorsal (C); b, last prosomal segment

and first 2 urosomal segments, lateral (£). Stage V: c, dorsal (B). Stage IV: d, dorsal (B).

of dense refractile points on fourth and fifth urosomal segments separate this species from the others.

These specimens were larger than the juveniles of

L. minuta and L. glacialis from the northeast Pacific, but were approximately the same size as L.

glacialis juveniles from the Arctic Ocean (Heron

and English, in prep.).

STAGE IV.—A specimen (1.14 mm) (Table 2) was
NUMBER 267

25

FIGURE 17.—Lubbockia species, stage V: a, dorsal (B). Stage

IV: b, dorsal (B).

compared with similar size specimens of Arctic L.

glacialis, and each urosomal segment of this undescribed species appeared more robust and wider in

relation to the length (Figure 176).

Lubbockia flemingi, new species

FIGURES 18-20

MATERIAL STUDIED {Tables 1, 2).—NE PACIFIC: 6 females,

0.83-0.91 mm, NP 7 et al., {including: holotype, 0.83 mm,

NP 7, USNM 168493; 2 paratypes, 0.84 mm, NP 13, 17,

USNM 168495, 168496); 2 males, 0.84, 0.85 mm, NP 16, 26

{including: allotype, 0.84 mm, NP 16, USNM 168494).

ANTARCTIC OCEAN: 1 male, 0.82 mm, AA 42.

FEMALE.—Average length about 0.85 mm. Prosome (Figure 18a,b) about 1.7 times length of

urosome. Pediger 5 with slight transverse dorsoposterior sclerotized ridge. Genital segment with areas

of external genital apparatus on anterior third of

dorsolateral surface, each area with a setule. Genital

segment and each postgenital segment with fine

spinules on posteroventral margins. Uropod about

3 times as long as wide; outermost proximal seta

about midsegment.

Rostrum with thickened posteroventral margin,

deltoid in ventral view as shown in figure of orientation of oral appendages (Figure 18c).

Antenna 1 with armature and segmentation similar in number to that of L. minuta.

Antenna 2 (Figure 18c) with 1 seta on second

segment. Third segment with inner setules and 3

subapical setae; armed terminally with 4 short claws

and 3 setae.

Labrum (Figure 18d) with free margin divided

into 2 rounded posteroventral lobes, each margin

with row of hyaline setules and small dentiform

processes on under surface; complex pattern of

layers of tissue over- and underlying sclerotized

areas.

Mandible (Figure 18e) with 2 outer elements,

both with rows of setules; mandible blade with row

of size-graduated scalelike denticles on median edge,

and apical lash; concave edge of mandible base with

a setulose element and rows of long setules.

Maxilla 1 (Figure 18/) flat, bilobed; 3 elements

on each lobe.

Maxilla 2 (Figure 19a) first segment expanded.

Second segment with a hyaline setule and stout

setulose element; terminal bifurcate, setulose element; curved setulose element on inner margin.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 18.—Lubbockia flemingi, new species, female: a, lateral (C): b, dorsal (C); c, cephalosorae, ventral (E); d, labrum, ventral {G); e, mandible, right (G); /, maxilla 1, right (G).

Maxilliped (Figure 196) 4-segmented. Second segment with a row and cluster of setules on inner surface (Figure 18c). Terminal segment a long claw.

Leg 1 (Figure 19c) with inner spine on basis, third

exopod segment with 3 short outer spines. Leg 2

(Figure 19<2) third exopod segment with 3 short

outer spines. Leg 3 (Figure 19e) inner edge of third

endopod segment with 3 setae. Leg 4 (Figure 19/)
NUMBER 267

27

FIGURE 19.—Lubbockia flemingi, new species, female: a, maxilla 2, right (G); b, maxilliped, left

(E); c, leg 1 (E); d, leg 2 (E); <r, leg 3 (E); /, leg 4 (E).

inner edge of third endopod segment with 2 setae.

Legs 2-4 endopods with a small vented projection

between terminal spines. Leg 1 coxa lacks an outer

papilla, but a vent is present.

Leg 5 (Figure 186) with free segment about i/s

length of genital segment; 2 terminal spines with

serrate, hyaline flange; outer spine slightly more

than 14 length of inner, which reaches beyond pos-
28

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

terior margin of genital segment.

Leg 6 probably represented by hyaline setule in

area of external genital apparatus.

MALE.—Prosome (Figure 20a) about 1.7 times as

long as urosome. Urosome (Figure 206) with the 4

postgenital segments and uropods bearing spinules

on ventral margin. Uropodal setae with relative

lengths as in female.

FIGURE 20.—Lubbockia flemingi, new species, male: a, lateral

(C); b, urosome, ventral (E).

Rostrum and mouthparts, except antenna 1, as in

female.

Antenna 1 with segments corresponding to terminal 3 of female fused in male; armature similar

to corresponding segments of female, except for addition of 1 esthete on segment 2 and 2 esthetes on

segment 4; additional esthetes and 1 esthete on each

of segments 6 and 7 somewhat longer than on

female.

Swimming legs and leg 5 as in female with additions of clusters of short spinules on legs 1-4, most

numerous on anterior surfaces of legs 1-3 and outer

coxa and basis of leg 4.

ETYMOLOGY.—With the name Lubbockia

flemingi, we wish to express our sincere appreciation to Professor Richard H. Fleming, Department

of Oceanography, University of Washington, for our

memorable introduction to oceanography.

REMARKS.—Lubbockia flemingi resembles L.

brevis Farran, 1908, described from a deep sample

off the coast of Ireland. Additional descriptions of

both female and male L. brevis from the Arctic

Ocean will be published in a report on Arctic

Oncaeidae (Heron and English, in prep.). In L.

brevis the length of the spiniform setule of female

leg 6 is about i/3 the length of the genital segment.

Lubbockia brevis uropod is 4 to 5 times as long as

wide on the female and about the same length as

the genital segment of the male. Lubbockia flemingi

and L. brevis differ from all other known Lubbockia species by having 3 inner setae on leg 3 third

endopod segment and 2 inner setae on leg 4 third

endopod segment. In addition, they are the only

Lubbockia species without a sexually dimorphic

maxilliped.

The vents associated with the terminal projections on the swimming leg endopods occasionally

have an exuding material, most apparent on stained

specimens.

Dr. T. Bjornberg informed us that the single

female reported as L. brevis from the Marchile

I Expedition (Bjornberg, 1973:364) measured 1.25

mm. She kindly sent us illustrations of the urosome

which showed that the uropod and the seta representing leg 6 resemble those of L. flemingi more

closely than those of L. brevis.

DISTRIBUTION.—Northeast Pacific and Antarctic

Oceans (Tables 1, 2); Southeast Pacific Ocean

(Bjornberg, 1973).
NUMBER 267 29

Lubbockia petersoni, new species

FIGURES 21-22

MATERIAL STUDIED {Tables 1, 2).—NE PACIFIC: 26 females,

0.95-1.06 mm, NP 10 et al. (including: holotype, 0.99 mm,

NP 24, USNM 168497; 9 paratypes, 0.95-1.05 mm, NP 10, 24,

33, USNM 168499-168501); 12 males, 0.85-1.03 mm, NP 10

et al. (including: allotype, 0.99 mm, NP 24, USNM 168498;

4 paratypes, 0.91-0.95 mm, NP 10, 24, 33, USNM 168499-

168501.

FEMALE.—Average length about 1.00 mm. Prosome about 1.8 times the length of the slender urosome, posterior corners rounded (Figure 2la,b).

Urosome first segment with slight transverse,

sclerotized dorsoposterior ridge. Genital segment

with areas of external genital apparatus posterior

to midpoint on dorsal surface of segment, each area

with 2 setules. Genital segment and each postgenital

segment with fine spinules on posterior margins.

Uropod with hyaline flap extending over base of

dorsal seta.

Rostrum with sclerotized pattern accentuating a

thickened rhomboidal ventral surface as shown in

ventral view of cephalosome (Figure 21c). Protruding sclerotized process posterior to rostrum (Figure

2Id), with vent near apex of process.

Antenna 1 with armature similar in number to

that of L. minuta; variable rudimentary articulation

between all segments, except distinct articulation

between segments 4, 5.

Antenna 2 with armature similar to that of L.

minuta, third segment with 4 terminal claws, 1 long

seta, and 2 short setae (Figure 21c).

Labrum (Figure 21e) with free margin divided

into 2 rounded posteroventral lobes, separated by a

triangular vertex, from which arises a short hyaline

hood.

Mandible (Figure 21/) with 2 adjacent, setulose

outer elements; mandible blade with row of scalelike denticles on median edge; concave edge of base

with a setulose element, spinules, and row of long

setules.

Maxilla 1 (Figure 21g) and maxilla 2 (Figure

21/?) with armature similar in number to that of

L. minuta; ornamentation differs as shown in figures.

Maxilliped (Figure 21i) first segment with posterior protuberant process (Figure 21a,c). Second

segment and terminal claw relatively reduced in

length; incomplete suture separating terminal claw.

Leg 1 (Figure 21;) with inner spine on basis, third

exopod segment with 3 short outer spines, proximal

spine reduced. Leg 2 (Figure 22a) third exopod segment with 3 short outer spines, proximal spine

reduced. Leg 3 (Figure 226) inner edge of third

endopod segment with 2 setae and 1 spine. Leg 4

(Figure 22c) inner edge of third endopod segment

with 1 seta and 1 spine. Legs 2, 3 endopods terminate with a small vented projection; leg 4 endopod

terminates with a vent between 2 most distal spines.

Leg 5 represented only by 1 seta on midlateral

surface of first urosomal segment (Figure 21a,&).

Leg 6 probably represented by 2 hyaline setules

in area of external genital apparatus.

MALE.—Average length about 0.93 mm. Prosome

about 1.8 times length of slender urosome (Figure

2U,e).

Urosome with the 4 postgenital segments bearing

fine spinules on posterior margins.

Rostral area and mouthparts, except antenna 1

and maxilliped, as in female; sclerotized process

posterior to rostrum.

Antenna 1 with segmentation similar to that of

female; armature similar to female except for addition of long esthete on segment 2 and 2 esthetes on

segment 4; all esthetes elongate.

Maxilliped (Figure 22/) with small segment between second segment and slender, elongate claw.

Swimming legs and uropod segment 1 (Figure

22g) as in female, except narrower flange on swimming leg spines.

Leg 6 probably represented by posteroventral

pointed flap on genital segment.

ETYMOLOGY.—It is a pleasure to name this species

for Willis K. Peterson, Department of Oceanography, University of Washington, who designed and

built the net for collecting deep plankton samples

during hydrographic casts off the Washington

coastal shelf. Mr. Peterson has generously made

these samples available to us.

REMARKS.—Lubbockia petersoni differs from all

other species of Lubbockia by the absence of leg 5.

The complex pattern of sclerotization in the rostral

area of L. petersoni resembles that in L. squillimana

and L. aculeata. The stocky antenna 2 resembles

that in L. flemingi except for having a longer lashlike seta. Maxilla 2 resembles that in L. flemingi.

Among Lubbockia species, only L. petersoni, L.

carinata, and L. forcipula have an anterolateral seta
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 21.—Lubbockia petersoni, new species, female: a, lateral (A); b, dorsal (A); c, cephalosome, ventral (E); d, rostrum and ventral process, lateral (antenna 1 and antenna 2 omitted)

(E); e, labrum, ventral (G); /, mandible, right (G); g, maxilla 1, left (G); h, maxilla 2, left (G);

i, maxilliped, right (E); ;, leg 1 (E).
NUMBER 267

31

FIGURE 22.—Lubbockia petersoni, new species, female: a, leg 2 (E); b, leg 3 (E); c, leg 4 (E).

Male: d, lateral (A); e, dorsal (A); /, maxilliped, right (E); g, first 2 urosomal segments, ventral (E).
32 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

on pediger 5, which is common to all other Oncaei- Sample NP 24 (Tables I, 2) also contained a

dae species. number of the pelagic tunicate Doliolum species.

The sclerotized process posterior to the rostrum Two L. petersoni specimens were inside of 1 dolioresembles a structure illustrated for the cyclopoid lid and 1 in another; this might not reflect a natural

copepods Nasomolgus cristatus Sewell, 1949 (fig. association.

355) (Lichomolgidae) ' and Hippomolgus latipes DISTRIBUTION.—Northeast Pacific Ocean (Tables

Humes and Ho, 1967 (fig. 44) (Clausidiidae). 1, 2).
Appendix

Tables

Table 1.—Sources of specimens (AA

TA = tropical west Atlantic Ocean, TP

Antarctic Ocean, M = Mediterranean Sea, NP = northeast Pacific Ocean,

tropical Pacific Ocean; for institutional abbreviations see "Acknowledgments")

Sample Cruise Station Date Location Depth (m) Collection

1

2

3

4

5

6

7

8

9

10

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP 11

NP 12

NP 13

NP 14

NP 15

NP 16

NP 17

NP 18

NP 19

NP 20

NP 21

NP 22

NP 23

NP 24

NP 25

NP 26

NP 27

NP 28

NP 29

NP 30

NP 31

NP 32

NP 33

NP 34

NP 35

NP 36

NP 37

NP 38

NP 39

NP 40

AA 41

AA 42

TP 43

TA 44

TA 45

M 46

M 47

NOAA Surveyor 75D

Brown Bear 332

344

349

352

357

368

380

Oshawa 6

Thompson TT002

TT030

TT082

Eltanin 27

Albatross

Bache

Gosnold 74

Silver Belle

1,2

8

AF6

4

5

11

19

21

25

26

31

35

25

16

16

20

21

19

19

22

20

23

25

28

30

31

32

33

36

37

38

40

13

12

12

32

65

P

P

P

1811

1817

5185

10209

2348

44

4 Oct 75

5Oct 75

26 Aug 63

19 May 64

20 May 64

23 May 64

27 May 64

28 May 64

30 May 64

31 May 64

2Jun 64

4Jun 64

24 Aug 64

22 Jan 65

22 Jan 65

25 Jan 65

26 Jan 65

20 Apr 65

20 Apr 65

22 Apr 65

10 Aug 65

12 Aug 65"

13 Aug 65

14 Aug 65

15 Aug 65

16 Aug 65

16 Aug 65

17 Aug 65

18 Aug 65

18 Aug 65

19 Aug 64

20 Aug 65

10 Nov 65

22 Oct 64

22 Oct 64

10 Aug 67

23 Jul 68

6 Aug 73

7 Aug 73

8 Aug 73

7 Jan 67

9 Jan 67

30 Mar 08

22 Mar 14

21 Aug 65

22 Aug 04

1 Feb 67

60°40.7'N, 147°41.1'W

60°4O.7'N, 147°43.1'W

45°20.0'N, 130°00.0'W

47°33.5'N, 126°48.5'W

47°33.8'N, 128°22.7'W

47°44.8'N) 134°55.8'W

49°56.0'N, 144°54.0'W

47°49.2'N, 144°57.7"W

45°17.0'N, 141°48.0'W

45°15.7'N) 132°44.0'W

45°15.7'N, 132°44.0'W

45°29.3'N, 126°58.0'W

45°19.5'N, 132°26.0'W

45°13.0'N, 136°04.0'W

45°13.0'N, 136°04.0'W

45°22.0'N, 128°10.0'W

45°27.0'N, 126°33.0'W

45°17.5'N, 138°34.0'W

45°17.5'N, 138°34.0'W

47°59.5'N, 139°58.0'W

45°22.0TSf, 128°36.0'W

45°20.0'N, 132°28.O'W

45°20.6'N, 134°56.5'W

45°17.5TSr, 139°09.6'W

46°20.8'N, 139°50.5'W

47°05.8'N, 139°51.0'W

47°58.5'N, 139°44.4'W

47°58.5'N, 139°44.4'W

47°47.6TST, 135°34.7'W

47°43.1'N, 134°17.0'W

47°33.3'N, 132°32.0'W

47°55.3'N, 129°09.5'W

45°32.0'N, 139°48.0'W

45°20.0'N, 139°06.0'W

45°20.0'N, 139°06.0'W

44°37.0'N, 128c00.0'W

47°66.0'N, 128°50.0'W

49°56.0'N, 144°54.0'W

49°56.0'N, 144°54.0'W

49°56.0'N, 144°54.0'W

62°59.0'S, 177°4O.O'E

65°59.0'S, 176°23.0'E

10°05.8'N, 124°18.5'E

off Bahamas

29°44.9'N, 76°5O.8'W

35°45.0'N, 3°41.0'W

Gulf of Naples

99-50

212-70

453-0

1800-0

2128-0

2536-0

4014-0

4306-0

4500-0

2000-0

3183-0

2750-0

1800-0

3500-0

4150-0

2000-0

2750-0

4000-0

4150-0

4000-0

2700-0

2000-0

2000-0

4250-0

4200-0

2000-0

4000-0

4000-0

3700-0

2O0O-O

3000-0

1967-0

4650-0

2250-0

4200-0

147-0

75-0

440-110

440-220

3000-0

2000-1000

2000-1000

1167

100-0

0-200-0

183

100-0

Heron and Damkaer

U.W. (Peterson)

UW (Frost)

UW (Thoreson)

SOSC

USNM 73654

USNM 79578

SOSC

R. N. Wolfenden

USNM 168505
34 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Table 2.—Distribution of specimens of Lubbockia species by samples (for identification of samples see Table 1)

Sample

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

NP

AA

AA

TP

TA

TA

M

M

1

2

3

4

5

6

7

8

9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

30

31

32

33

34

35

36

37

38

39

40

41

42

43

44

45

46

47

carinata forcipula

9 <? IV 9

1

1

1

1

1

1

1

1

1 2

wilsonae

9 d1 V IV

3

1

1

1

3 3 5

3 3 2

1

3

III

1

1

1

1

aculeata

9

1

2

2

squillimana

9 d1 V IV III

1 1

82 311 7 2

3

20 3 3

9

1

2

1

3

1

2

3

1

2

2

1

1

1

1

2

2

1

1

4

1

minuta

d

1

 V

1 1

1

1

1

1

1

IV

2

1

1

1

2

1

glacialis L

9 V IV

1

1

1

1

1

1 1

1

. species

V IV

2

1 1

1

1

2

flemingi

9

1

1

1

1

1

1

d

1

 V

l

1

l

1

1

9

1

1

1

1

2

8

1

1

1

2

1

2

1

3

petersom

d

1

1

1

1

3

1

1

1

2

1

V

1

1

1

1

1

1

1

3

IV

1

1

1

1

1
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1976. Discarded Appendicularian Houses as Sources of

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1971. The Case of the Nonubiquitous Telson and the

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