ROGER F. CRESSBY Revision of the

f Genus Alebion

(Copepoda: Caligoida)

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 123
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111
SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 123

Roger F. Cressey Revision of the

Genus Alebion

(Copepoda:

Caligoida)

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON

1972
ABSTRACT

Cressey, Roger F. Revision of the Genus Alebion (Copepoda: Caligoida). Smithsonian Contributions to Zoology, number 123, 29 pages, 132 figures, 1972.—Eight

species of Alebion are recognized as valid: A. carchariae Krayer, A. maculatus Wilson, A. elegans Capart, A. glaber Wilson, A. gracilis Wilson, A. lobatus Cressey, and

A. pacificus, new species. Other species have been placed in synonymy. The new

species is described from the eastern Pacific Ocean where it is apparently endemic.

The spermatophore attached to the female is shown to be a very useful character in

recognizing the 8 species. The patterns of canaliculations on the surface of the postoral adhesion pad is also shown to be taxonomically important in both sexes. Keys

to both sexes of the species are given.

Library of Congress Cataloging in Publication Data

Cressey, Roger F, 1930-

Revision of the genus Alebion (Copepoda: Caligoida)

(Smithsonian contributions to zoology, no. 123)

Bibliography: p.

1. Alebion. I. Title. II. Series: Smithsonian Institution. Smithsonian contributions to zoology,

no. 123.

QL1.S54 no. 123 [QL444.C7] 591'.08s [595'.34] 72-540

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

in the Institution's annual report, Smithsonian Year.

For sale by the Superintendent of Documents, U.S. Government Printing Office

Washington. D.C. 20402 - Price 40 cents i paper cover)
Roger F. Cressey Revision of the

Genus Alebion

(Copepoda:

Caligoida)

Introduction

Collections of copepods parasitic on shallow water

sharks invariably contain members of the genus

Alebion. Collections made in recent years from several geographic areas have resulted in sufficient material available to revise the genus and to redescribe

the 8 species where needed. Thirteen species have

been described in the genus since the first was described by Kreyer in 1863. The revision to follow

recognizes 8 of these species as valid, including one

described as new.

Members of the genus are found in all major

oceans with the exception of the polar regions. Some

species are geographically restricted while others are

more ubiquitous. Both sexes seem to wander freely

over the body surface of the host with females generally more abundant in collections than males.

All material is preserved in 70 percent alcohol

and deposited in the collections of the Smithsonian

Institution.

All figures were drawn with the aid of a Wild

drawing tube.

The morphology of the spermatophore, a male

structure, is discussed with the descriptions of the

females since it is with this sex that the fully formed

spermatophore is found.

Roger F. Cressey, Department of Invertebrate Zoology,

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

Unless otherwise stated all measurements are from

a single specimen.

Acknowledgments

I thank Dr. Torben Wolff of the Copenhagen Museum for the loan of the type specimen of Alebion

carchariae Kroyer.

Mrs. Molly Griffin assisted in the preparation of

the figures.

Alebion Krpyer, 1863

Alebion Kreyer, 1863:165

Caligera van Beneden, 1892:258

DIAGNOSIS—Euryphoridae. Frontal plate separate,

without lunules. Thoracic segments bearings legs 1-3

fused. Thoracic segment bearing legs 4 free and bearing bilobed dorsal plate in female. Abdomen 2-segmented. First antenna 2-segmented. Second antenna

clawlike. Postoral adhesion pads present. Second

maxilla with two terminal flagella. Sternal furca absent. Maxilliped clawlike. Legs 1-3 biramous; rami

of leg 1 2-segmented, rami of legs 2 and 3 3-segmented. Leg 4 reduced, uniramous, 1-segmented. Leg

2 of male with two modified spines on exopod. Egg

strings uniseriate.

Type-species, by monotypy, Alebion carchariae

Kroyer. Gender masculine.

1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to the Known Species of Alebion

FEMALES

1. Genital segment with posterior processes 2

Genital segment without posterior processes A. glaber Wilson

2. Genital segment without prominent lateral bulges (Figures 58, 100, and 111) 3

Genital segment with prominent lateral bulges (Figures 1, 29, 44, and 88) 5

3. Posterior processes of genital segment (leg 5) not extending beyond caudal rami

A. pacificus, new species

Posterior processes extending well beyond caudal rami 4

4. Attached spermatophores divergent along entire length (Figure 60); surface of postoral

adhesion pad without long canals (Figure 64) A. elegans Capart

Attached spermatophores divergent in anterior half only (Figure 103); surface of postoral

adhesion pad with long canals (Figure 104) A. lobatus Cressey

5. Attached spermatophores widely divergent (Figure 45); maxilliped claw with accessory

process A. crassus Wilson

Attached spermatophores not divergent (Figures 3, 30, and 89); maxilliped claw simple 6

6. Attached spermatophores with central sinus (Figure 3) or anterior end wider than posterior

(Figure 4) ; central area of postoral adhesion pad with canals broken to form "islands"

(Figure 9) A. carchariae Kroyer

Attached spermatophores without central sinus and anterior end not widest (Figures 30 and

89) ; surface of postoral adhesion pad with long, straight canals (Figures 34 and 92) . 7

7. Sclerotized ring lateral to anterior end of attached spermatophores (Figure 89); first abdominal segment with prominent lateral alae A. gracilis Wilson

No sclerotized ring as above; first abdominal segment with only slight bulge at posterior

corners A. maculatus Wilson

MALES

1. Modified spine on second segment of exopod of leg 2 extending beyond, to, or nearly to

tip of modified spine of third segment (distalmost) 2

Modified spine of second segment extending only to middle or less of modified spine of third

segment 4

2. Modified spine of second segment at least twice as thick as modified spine of third segment 3

Modified spine of second segment of about same thickness as that of third segment

A. maculatus Wilson

3. Outermost of the two modified spines thumblike, tapered but not constricted along its

length; innermost spine of uniform thickness throughout its length and reaching to tip of

outer spine A. crassus Wilson

Outermost spine swollen in basal half, distal half narrowed; innermost spine shorter than

outer, distal fourth narrowed A. glaber Wilson

4. Canaliculations on surface of postoral pad long and straight, not forming "islands" (Figure

92) A. gracilis Wilson

Canaliculations forming "islands" in part or on all of surface 5

5. Canaliculations forming islands over entire surface of postoral adhesion pad

A. elegans Capart

Canaliculations long and straight on part of adhesion pad surface 6

6. Canaliculations forming islands in middle of pad, long and straight near anterior and

posterior borders A. carchariae Kreyer

Canaliculations forming islands along posterior border, long and straight in anterior region

only 7

7. Proximal modified spine on exopod of leg 2 bearing 20-30 heavy spinules and reaching to

just beyond base of distalmost spine (Figure 110) A. lobatus Cressey

Proximal spine bearing 50-100 spinules and reaching nearly to middle of distalmost spine

(Figure 132) A. pacificus, new species
NUMBE R 12 3 3

Alebion car char iae Kr^yer, 1863

FIGURES 1—28

Alebion carchariae Kroyer, 1863:165.—Brady, 1883:135.—

Bassett-Smith, 1899:462.—Brian, 1908:3; 1912:11.—

Wilson, 1907:704; 1932:422.—Bere, 1936:593.—Capart,

1953:655.—Barnard, 1955:254.—Heegaard, 1955:49;

1962:175.—Vaissiere, 1959:535.—Yamaguti, 1963:99.—

Cressey, 1970:4.

Caligera difficilis van Beneden, 1892:258.—Wilson, 1907:

703.

Alebion difficile Basset-Smith, 1899:462.

Alebion fuscus Wilson, 1921:2, 1932:421.—Carvalho, 1940:

275; 1951:137.—Vaissiere, 1959:549.—Yamaguti, 1963:

100.

Alebion difficilis Yamaguti, 1963:100.

Alebion gracilis.—Lewis, 1966:136.—Cressey, 1967:5.

MATERIAL EXAMINED.—Holotype female on loan

from the Copenhagen Museum, 11 collections identified as A. carchariae in the Smithsonian collections

(USNM 12664, 56596, 56603, 64051, 74303, 78956,

79157, 79455, 110819, 120747, and 120748), 3 collections [USNM 6083, 6210, and 32724 (co-types)],

type specimens of A. fuscus (USNM 58559 and

53560), 5 previously unrecorded collections from the

western Atlantic and Gulf of Mexico, M/V Delaware

and M/V Oregon II cruises), 71 collections from the

west coast of Florida, 8 collections from the western

Indian Ocean.

FEMALE.—Body form as in Figure 1. Total

length 8.1 mm (7.65-8.70) and greatest width 4.05

mm (3.60—4.50 mm) based on an average of 6

specimens from the north Atlantic. Genital segment

2.25 X 2.78 mm somewhat wider than long; widest

in posterior third, lateral margin at widest part with

prominent spines, posterior outer corners produced

to form processes extending beyond caudal rami.

Abdomen (Figure 2) 2-segmented; first segment

with lateral processes extending nearly to junction

of caudal rami. Spermatophores attached ventrally

on genital segment near attachment of abdomen.

Spermatophore of 2 forms. Most common form as in

Figure 3, each with an inner median sinus so that

when they are in position a central cavity is formed.

Second form as in Figure 4, no resultant central

cavity when they are in position. Caudal ramus (Figure 5) bears 4 plumose terminal setae and 2 smaller

subterminal setae; each ramus longer than wide,

520/* x 260/t.

First antenna (Figure 6) 2-segmented, basal segment with several short plumose setae, terminal segment with 13 setae (2 fused at base), plumose setae

indicated in figure. Second antenna (Figure 7) in

form of claw; terminal claw with seta near middle,

tip lightly pigmented. Tip of mouth tube with mandible blades within as in Figure 8. Adhesion pads

lateral to mouth tube as in Figure 9, surface of pad

canaliculate, forming islands at center. First maxilla

represented by a group of 3 small setae lateral to

mouth tube. Second maxilla (Figure 10) with 2

terminal flagella, shorter one about half the length

of longer; second segment with patch of setules (Figure 11); longest flagellum with rows of spinules

along its length, shorter flagella with serrate edge as

in Figure 12. Maxilliped (Figure 13) in form of

short stout claw, patches of scales on basal segment,

seta on inner edge near base of terminal claw.

Legs 1—3 biramous. Leg 1 (Figure 14) exopod 2-

segmented, basal segment with short spine on outer

distal corner, second segment (Figure 15) with 3

terminal spines (median spine modified as a flattened paddlelike process) and 4 plumose setae along

inner edge, distalmost seta much reduced. Leg 2

(Figure 16) exopod 3-segmented; basal segment with

membranous outer sheath attached at outer margin

and lying over most of ramus, toothed spine on outer

distal corner with row of spinules near base; last 2

segments each bear an inwardly directed modified

spine, palmate process near base of each, long plumose

setae on all segments as indicated in the figure;

endopod 3-segmented, each segment with long plumose setae as in the figure. Leg 3 (Figure 17)

exopod similar to leg 2 except last segment bears

2 modified spines (Figure 18) instead of 1: endopod

with second segment longer than other 2 combined.

Leg 4 (Figure 19) uniramous and reduced, bearing 4

short setae (1 lateral, 3 terminal). Leg 5 (Figure 20)

produced from outer distal corners of genital segment,

spines along outer margin gradually increase in size

posteriorly; 3 plumose setae present at tip in addition

to stout spines (terminal spines usually 3 in number).

Leg 6 absent.

MALE.—Body form as in Figure 21. Total length

6.0 mm and greatest width 2.85 mm based on a

single specimen from the north Atlantic. Genital segment somewhat longer than wide (1.5 X 1.3 mm) ;

edge of genital segment (Figure 23) with many

spinules near lateral margins and a group of stouter
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

spines anterior to leg 5. Abdomen 2-segmented, each

segment measures 0.55 mm and 0.69 mm respectively. Caudal ramus (Figure 24) armed as in female

and about twice as long as wide (0.51 X 0.25 mm).

Cephalic appendages as in female except as noted

below. Second antenna (Figure 25) in form of a

stout claw; basal segment with large adhesion pad on

surface, second segment with 2 smaller adhesion pads

and stout inner spine, terminal claw with stout (but

weak) spine near base, median seta and median

accessory clawlike process. Second maxilla as in female except serrations on edge of shorter flagellum

pointed rather than blunt as in female. Maxilliped

(Figure 26) in form of stout claw; median segment

with prominent patch of scalelike spinules as in figure, terminal segment with inner seta.

Legs 1—4 as in female except as noted below. Exopod spine at inner distal corner of last segment

(Figure 27) bifid and bearing accessory spinule at

tip. Leg 2 exopod (Figure 28) bearing modified

spine on outer surface of each of last 2 segments;

each spine with papillae on surface and clublike in

appearance, these spines correspond to the 2 modified spines on female second leg exopod. Leg 5

(see Figure 23) represented by three plumose setae

and another plumose seta near base along midmargin

of genital segment. Leg 6 (see Figure 23) represented by 3 short plumose setae at outer distal corner of genital segment.

REMARKS.—This species, described by Kroyer in

1863, and recorded many times since by various

authors is found in all major oceans and on a number

of shark species. Examination of the type material

was essential in correcting the identifications of several Smithsonian collections by various authors (Wilson, Lewis, and Cressey).

In an earlier paper (Cressey, 1970) I reported

on the occurrence of 2 forms taken by the spermatophore of this species and A. lobatus. I will not repeat

the discussion here but I should point out that the

two forms occurred in the sample studied in the ratio

of approximately 5:1. I could offer no explanation

for the two forms at that time and I am still not able

to do so.

Alebion pacificus, new species, described below

from the eastern Pacific Ocean seems to be closely

related to A. carchariae based on morphological characters.

Alebion maculatus Wilson, 1932

FIGURES 29-43

Alebion carchariae.—Bassett-Smith} 1898:366.

Alebion maculatus Wilson, 1932:423

Alebion alatus Gnanamuthu, 1951:1237.—Vaissiere, 1959:

548._Yamaguti, 1963:99. — Cressey, 1967:5. —Pillai,

1967:1618.

Alebion spinosus Rangnekar, 1959:53.—Pillai, 1967:1617.

Alebion megacephalus Kirtesinghe, 1956:15; 1964: 82.—

Yamaguti, 1963:100.—Pillai, 1967:1618.

MATERIAL EXAMINED.—Four females and 2 males

collected by the author from the body surface of

Carcharinus limbatus from Nosy Be, Malagasy Republic.

FEMALE.—Body form as in Figure 29. Total

length 4.88 mm and greatest width 2.63 mm based

on a single specimen. Genital segment wider than

long (2.06 X 1.67 mm), each lateral margin with

prominent spinose bulge; posterior outer corners

produced, extending well beyond caudal rami. Abdomen 2-segmented, first segment with short alate processes laterally. Spermatophores (Figure 30) slender,

not divergent. Caudal ramus (Figure 31) armed as

in A. carchariae; each ramus longer than wide,

170 n X 92 u.

Cephalic appendages as in A. carchariae except

as noted. Second antenna (Figure 32) similar to that

of other species in genus but with adhesion pads

as shown in figure. Second maxilla similar to A.

carchariae; short flagellum (Figure 33) with serrate

edges as shown in figure. Adhesion pads lateral to

mouth tube as in Figure 34. Maxilliped (Figure 35)

in form of short, stout claw; basal segment with 2

scaly areas on inner surface and small raised adhesion

pad near outer distal corner, short seta near inner

edge of terminal segment.

Legs 1—3 biramous and as in A. carchariae. Terminal segment of leg 1 exopod armed as in Figure

36. Leg 4 as in A. carchariae. Leg 5 (Figure 37)

with 2 plumose setae near tip.

MALE.—Body form as in Figure 38. Total length

1.69 mm and greatest width .90 mm, based on a

single specimen. Genital segment (Figure 39) nearly

twice as long as wide (.97 X .58 mm). Abdomen 2-

segmented; segments measure .29 and .27 mm respectively. Caudal rami (Figure 40) about twice as

long as wide (230 n X 110 |i).

Cephalic appendages as in female except as noted.

Second antenna (Figure 41) with adhesion pads on
NUMBE R 12 3

basal segment as shown in figure; accessory process

on claw in addition to median seta. Maxilliped (Figure 42) basal segment with conical inner process

bearing few serrations as shown in figure, terminal

claw well-developed and bearing median seta.

Legs 1-4 as in female except for modified spines

on exopod of leg 2 (Figure 43). Leg 5 represented

by group of 3 setae near lateral midarea of genital

segment. Leg 6 represented by 3 plumose setae near

posterior outer corner of genital segment.

REMARKS.—This species was first figured by Basset-Smith in 1898 as another record of A. carcariae. Wilson in 1932 noted differences in BassettSmith's description and the original description of

A. carchariae and suggested that the material of Bassett-Smith from Aden actually represented another

species for which he proposed the name maculatus.

This rather obscure reference by Wilson has been

largely unnoticed by later authors and Gnanamuthu

and Rangnekar both described this species as new

under the names alatus and spinosus respectively. I

identified specimens from Malagasy Republic as

alatus in 1967. Reexamination of this material and

the literature have convinced me that these are all

one species and should now be referred to as A.

maculatus.

The specimen described by Kirtesinghe as A.

megacephalus is actually an immature female of A.

maculatus.

This species appears to be restricted to the Indian

Ocean.

Alebion crassus Wilson, 1932

FIGURES 44-57

Alebion crassus Wilson, 1932:423.—Vaissiere, 1959:549.—

Yamaguti, 1963:99.—Shiino, 1965:426.

Platyporinus alata Rao, 1950:306.

Alebion echinatus Capart, 1953:655.—Shiino, 1955:177.—

Vaissiere, 1959:544. — Yamaguti, 1963:100. — Lewis,

1966:130.—Pillai, 1967:1618.

MATERIAL EXAMINED.—Holotype female (USNM

56614) and paratynes (1 female and 1 male, USNM

56615), 2 collections from Hawaii (USNM 110817

and 110818, identified originally as echinatus) and a

single collection from Florida (USNM 10174, previously unidentified).

FEMALE.—Body form as in Figure 44. Total

length 14.4 mm, greatest width 10.0 mm (measurements from holotype). Genital segment (Figure 45)

about as long as wide (6.35 X 6.00 mm) ; lateral

edges with prominent spinose bulge at posterior third,

posterior corners produced. Abdomen 2-segmented,

first segment with conspicuous lateral alae, second

segment slightly swollen laterally. Caudal ramus (Figure 46) longer than wide (820 \i X 520 \i) and somewhat inflated, terminal setae plumose but much

shorter in relation to length of ramus when compared with other species in the genus.

The excellent description and figures provided by

Shiino in 1955 (as A. echinatus) make a complete

redescription of this species unnecessary. The following

points supplement Shiino's description.

Second maxilla (Figure 47) second segment with

patch of broad scalelike setules; shorter terminal

flagellum much shorter in relation to longer flagellum

than in other species of genus and armed as in the

figure. Postoral adhesion pad lateral to mouth tube

(Figure 48) triangular and surface with canaliculations as in figure. Maxilliped (Figure 49) basal segment with 3 patches of scales as shown in figure

(scales in inner patch larger than those in outer

patches); claw with accessory process (Figure 50).

Last exopod segment of leg 1 as in Figure 51.

MALE.—Body form as in Figure 52. Total length

9.75 mm, greatest width 4.95 mm. Genital segment

about twice as long as wide (2.78 X 1.35 mm).

Abdomen 2-segmented; first segment 0.52 mm long,

second segment 0.75 mm long and wider than first.

Caudal rami (Figure 54) about twice as long as wide

(0.75 X 0.37 mm) and armed as in figure.

Second antenna (Figure 55) with prominent adhesion pads as in figure. Maxilliped (Figure 56)

basal segment with raised scale-covered area along

inner edge, claw with stout accessory process.

Second leg exopod (Figure 57) with modified

spine on each of last two segments; outermost spine

thumblike, inner surface covered with papillae; innermost spine much thinner and bearing small papillose

knob at tip.

REMARKS.—This species has been reported a number of times. Shiino redescribed the species in 1955

(as A. echinatus) from specimens of both sexes

found on Sphyrna zygaena in Japanese waters. He

commented on the similarity between Wilson's A.

crassus and Capart's A. echinatus but maintains the

integrity of the two species on the basis of Wilson's

illustration of the male showing a constriction in the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

fourth segment not found in the Japanese material.

After examining the type male, I have determined

that Wilson's figure misrepresents that character and

that the two species are actually synonymous. The

type male has been redrawn.

This species is apparently restricted to sharks of

the genus Sphyrna and is worldwide in distribution.

Alebion elegans Capart, 1953

FIGURES 58-73

Alebion elegans Capart, 1953:656.—Vaissiere, 1959:541.—

Yamaguti, 1963:100.—Cressey, 1967:5; 1970:5.

MATERIAL EXAMINED.—Four collections from Sarasota, Florida, and one collection from the Indian

Ocean.

FEMALE.—Body form as in Figure 58. Total length

9.9 mm, greatest width 5.7 mm based on a single

specimen. Genital segment (Figure 59) about as

long as wide (2.3 mm X 2.5 mm) ; spinose area on

lateral margin but not noticeably bulging. Abdomen

first segment with prominent lateral alae, second segment narrow anteriorly. Spermatophores (Figure 60)

oval, widest in anterior third and divergent when in

place on genital segment. Caudal rami (Figure 61)

longer than wide (410 ji X 190 \i).

First antenna as in A. carchariae. Second antenna

(Figure 62) without prominent adhesion areas, last

segment in form of claw and bearing a seta near

midinner margin. Second maxilla short flagellum with

serrations as shown in Figure 63. Adhesion pad

lateral to mouth tube (Figure 64) with canaliculations forming rounded islands on surface (see detail). Maxilliped (Figure 65) in form of stout claw,

without noticeable adhesion areas, a small patch of

scales near outer distal corner of basal segment (on

underside of maxilliped shown in Figure 65).

First leg exopod last segment as in Figure 66.

Legs 2, 3, and 4 as in A. carchariae. Leg 5 (Figure

67) with 3 short plumose setae at tip in addition to

stout spines; fifth leg extends well beyond caudal

rami.

MALE.—Body form as in Figure 68. Total length

5.55 mm, greatest width 3.10 mm, based on a single

specimen. Genital segment (Figure 69) longer than

wide (1.3 X 0.9 mm) with scattered spinules on

anterior lateral margins. Abdomen 2-segmented, segments measure 0.52 and 0.46 mm long respectively.

Caudal ramus (Figure 70) about twice as long as

wide (0.32 X 0.15 mm).

Appendages as in female except as noted below.

Second antenna with conspicuous adhesion pads;

claw with accessory process near midmargin, seta

near base of process. Maxilliped (Figure 72) with

bilobed, scale covered, raised area on inner margin

opposing claw, claw with seta near midmargin.

Leg 2 exopod with modified spines (Figure 73)

on last two segments, each spine with conspicuous

papillae. Leg 5 represented by small process bearing

3 plumose setae near midmargin of genital segment.

Leg 6 represented by 3 plumose setae at each distal

corner of genital segment.

REMARK.—This species is a common parasite on

sharks of the genus Sphyrna and so far has only

been reported from the Atlantic and Indian Oceans.

Alebion glaber Wilson, 1905

FIGURES 74-87

Alebion glabrum Wilson, 1905:129.—Rathbun, 1905:93.

Alebion glaber Wilson, 1907:708.—Fowler, 1912:481.—

Wilson, 1932:419. —Vaissiere, 1959:548.—Yamaguti,

1963:100.

MATERIAL EXAMINED.—Type specimens (USNM

8123, both sexes, no holotype designated) and twenty

additional collections in the Smithsonian Institution

(USNM).

FEMALE.—Body form as in Figure 74. Total

length 10.80 mm, greatest width 4.80 mm, based

on a single specimen. Genital segment about as long

as wide (2.63 X 2.55 mm), lateral margins without

ornamentation and without long processes originating

on outer posterior corners. Abdomen 2-segmented,

first segment wider than second. Spermatophores

(Figure 75) with anterior two-thirds slightly divergent, anteriormost portion more heavily sclerotized.

Caudal ramus (Figure 76) only slightly longer than

wide (300 y. X 240 \i) and armed as in other members of the genus.

First antenna as in A. carchariae. Second antenna

similar to other members of the genus, claw (Figure

77) stout and heavily sclerotized. Second maxilla as

in other species in the genus, shorter flagellum with

serrations as in Figure 78. Maxilliped (Figure 79)

a short, stout claw with patches of scales on basal

segment as indicated in figure. Adhesion pad lateral
NUMBE R 12 3

to mouth tube (Figure 80) with canaliculations on

surface as in figure.

Legs 1—4 as in A. carchariae except for slight

differences in exopod last segment (Figure 81). Leg

5 represented by small podomere bearing 3 short

setae attached at outer posterior comer of genital

segment. Leg 6 absent.

MALE.—Body form as in Figure 82. Total length

5.7 mm and greatest width 2.55 mm, based on a

single specimen. Genital segment somewhat longer

than wide (1.6 X 1.16 mm), edge of genital segment

unarmed except for legs 5 and 6. Abdomen (Figure

83) 2-segmented, segments measuring 0.43 and 0.64

mm in length respectively. Caudal ramus (Figure 84)

longer than wide (383 \i X 248 u) and armed as in

other members of genus.

Cephalic appendages as in female except as noted

below. Second antenna (Figure 85) with stout claw,

tip of claw slightly bifid, adhesion pads present as indicated in figure. Maxilliped (Figure 86) in form of

claw, basal segment with large patch of scalelike

spinules on inner margin, tip of claw with fine

striations.

Legs 1-4 as in female except for modified spines

on exopod of leg 2 (Figure 87); spine on basal

segment stout, bearing knoblike spinules and markedly narrowed in distal third; more proximal spine

smaller and with spinules on distal tip only. Leg 5

located at midmargin of genital segment, consisting

of small podomere bearing 3 short setae and basal

seta. Leg 6 represented by 3 short setae at outer

posterior corners of genital segment.

REMARKS.—This species differs noticeably from

other members of the genus by the absence of the

fifth leg process in the female. Its affinity to the

other species can be seen by a comparison of the

appendages, notably the modified spines on the second leg exopod of the male.

This species has been collected from the east coast

of the United States. It seems to be restricted primarily to smaller inshore species of sharks, however,

two Smithsonian collections are from rays.

Alebion gracilis Wilson, 1905

FIGURES 88-99

AUbion gracile Wilson, 1905:128.—Rathbun, 1905:93

Alebion gracilis Wilson, 1907:704; 1932:420.—Vaissiere,

1959:549.—Yamaguti, 1963:100.—Cressey, 1970:5.

MATERIAL EXAMINED.—Holotype female (USNM

8122), 9 collections identified as A. gracilis in the

Smithsonian collections, and a single collection made

by the author off Sarasota, Florida.

FEMALE.—Body form as in Figure 88. Total

length 9.3 mm (9.0-9.6 mm) and greatest width

5.4 mm (4.95—5.55 mm) based on an average of 5

specimens. Genital segment (Figure 89) somewhat

wider than long (3.68 X 3.23 mm), widest in posterior third, lateral margin with prominent spinules

as in A. carchariae; posterior outer corners produced and extending well beyond caudal rami. Abdomen 2-segmented, first segment with lateral expansions much shorter than in A. carchariae. Spermatophores slender, posterior half only slightly wider

than anterior, anterior portions lie between two

sclerotized rings on ventral surface of genital segment. Caudal ramus (Figure 90) nearly twice as

long as wide (370 n X 200 \i), armed as in other

members of the genus.

Cephalic appendages as in A. carchariae except

as noted below. Tip of shorter flagellum of second

maxilla with' serrations as indicated in Figure 91.

Adhesion pad lateral to mouth tube with canaliculations on surface as in Figure 92. Maxilliped (Figure

93) with short, stout claw and patch of fine scales

on surface of basal segment.

Legs as in A. carchariae. Last exopod segment of

leg 1 as in Figure 94.

MALE.—Body form as in Figure 95. Total length

5.7 mm (5.3-6.3 mm) and greatest width 3.4 mm

(3.3—3.5 mm) based on an average of 5 specimens.

Genital segment (Figure 96) longer than wide (1.5

X 1.1 mm), widest portion with a few spinules on

lateral margin. Abdomen 2-segmented, each segment

measures 0.44 and 0.55 mm respectively, second

segment wider than first. Caudal ramus (Figure 97)

longer than wide (230 \i X 180 n) and armed as

in other members of genus.

Cephalic appendages as in female except as noted

below. Second antenna (Figure 98) with stout claw

bearing well-developed accessory process, adhesion

pads as indicated in the figure.

Legs 1-4 as in the female except for the presence

of modified spines on the exopod of leg 2 (Figure 99).

Legs 5 and 6 as in A. carchariae.

REMARKS.—This copepod superficially resembles

A. carchariae and has been confused with that species in recent years (Lewis, 1966 and Cressey, 1967).
8 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Several collections in the Smithsonian identified by

Wilson as A. gracilis are in fact A. carchariae. As

a result of this revision and reassessment of the

collections and literature it appears that A. gracilis

is restricted to the east coast of North America and

is relatively uncommon. Females of A. gracilis and

A. carchariae can be easily separated by the nature

of the attached spermatophores, the canaliculations

on the postoral adhesive pad, and the lateral expansions on the first abdominal segment.

Alebion lobatus Cressey, 1970

FIGURES 100-110

Alebion lobatus Cressey, 1970:7.

MATERIAL EXAMINED.—Thirty collections (including

types) from off west coast of Florida as previously

reported (Cressey, 1970).

FEMALE.—A description of the female of this species will not be repeated here since it has been recently provided elsewhere (Cressey, 1970). Some

illustrations are provided, however, to enable the

reader to identify females of this species without

consulting the previous work. The only feature here

added to the original description of the female is the

nature of the canaliculations on the surface of the

postoral adhesion pad (see Figure 104).

MALE.—Body form as in Figure 105. Total length

6.2 mm (6.1-6.4 mm) and greatest width 3.2 mm

(3.1—3.4 mm) based on an average of five specimens. Genital segment (Figure 106) longer than

wide (1.50 X 1.16 mm); few scattered spinules on

anterior lateral edge, widest anteriorly. Abdomen 2-

segmented, segments measuring 0.54 and 0.70 mm

long respectively. Caudal ramus (Figure 107) nearly

twice as long as wide (940 \i X 580 u), armed as in

other members of genus.

Cephalic appendages as in female except as noted

below. Second antenna (Figure 108) with stout terminal claw bearing small accessory spine near middle

of inner curved margin; prominent adhesion pads as

indicated in figure. Maxilliped (Figure 109) with

stout claw; inner margin of basal segment with

raised scale-covered areas opposing claw.

Legs \—4 as in female except for modified spines

on exopod of leg 2 (Figure 110); proximal spine

stout and bearing several heavy spinules, distal

spine narrowed in basal fourth and continuing same

width to tip. Leg 5 represented by small podomere

bearing three setae near midlateral margin of genital

segment. Leg 6 represented by three setae at posterior outer corners of genital segment.

REMARKS.—This species so far has been reported

only from Carchdrinus milberti in the Gulf of Mexico.

Alebion pacificus, new species

FIGURES 111-132

Alebion gracilis.—Shiino, 1959:316.

MATERIAL EXAMINED.—Holotype female (USNM

139170), allotype (USNM 139171), and 4 paratypes females (USNM 139172) from the body surface of Carcharinus malpeloensis caught near Tres

Madres Is. Four additional collections from C.

azureus, C. galapagensis, and C. malpeloensis (2)

from the eastern Pacific off Mexico and Lower California. An additional collection was found in the

British Museum labeled A. gracilis from "sand shark"

off Baija Honda, Panama.

FEMALE.—Body form as in Figure 111. Total

length 9.6 mm ( 9.0-10.2 mm) and greatest width

4.5 mm (4.3^.8 mm) based on an average of 5

specimens. Cephalon only slightly longer than wide

4.5 X 4.6 mm) and comprising nearly one-half total

length. Thoracic segments bearing legs 1—3 fused.

Thoracic segment bearing leg 4 with dorsal bilobed

plate. Genital segment (Figure 112) about as long

as wide (2.5 X 2.5 mm), a row of spinules along

anterolateral edge; genital segment with slight lateral

bulge, superficially resembling form produced in A.

elegans and lobatus, posterior outer corners produced. Abdomen 2-segmented, segments incompletely divided; first segment with prominent lateral winglike expansions, second segment narrowed anteriorly. Spermatophores (Figure 113) each with anterior

outwardly directed lobe, widest anteriorly, the two

spermatophores joined along most of midline and

lying nearly parallel with each other, central sinus

between them similar to that seen in A. carchariae.

Caudal ramus (Figure 114) longer than wide (630/x

x

 410^) and armed as in figure.

First antenna (Figure 115) as in other members

of genus except for two terminal setae with blunt

tips. Second antenna (Figure 116) in form of claw

with small rugose areas as indicated in figure. Mandible and first maxilla as in A. carchariae. Second maxilla (Figure 117) having same general form as other
NUMBE R 12 3

members of genus with rows of setae near middle of

terminal segment (Figure 118) and edge of shorter

flagellum as in Figure 119. Postoral adhesion pad with

canaliculations as indicated in Figure 120. Maxilliped

(Figure 121) in form of stout claw, claw incompletely divided proximal to seta.

Legs 1—3 biramous. Leg 1 (Figure 122) exopod 2-

segmented, basal segment with small spine on outer

distal corner and row of hairs along inner edge, second segment (Figure 123) with three terminal

spines, middle one modified as in other members of

genus, and four inner setae, distalmost one short;

endopod 2-segmented, first segment unarmed, second segment with three setae. Leg 2 (Figure 124)

exopod 3-segmented, first segment with mantlelike

membrane (not shown in figure) covering most of

exopod; endopod 3-segmented; both rami armed as

in A. carchariae. Leg 3 (Figure 125) rami 3-segmented and armed as in A. carchariae. Leg 4 as in

A. carchariae. Leg 5 (Figure 126) produced as an

extension of posterior outer corners of genital segment, tip armed with two short spines and three

stout plumose setae. Leg 6 absent.

Egg strings uniseriate.

MALE.-—Body form as in Figure 127. Total length

6.45 mm and greatest width 3.23 mm based on a

single specimen. Cephalon slightly wider than long

(3.23 X 3.10 mm). Genital segment (Figure 128)

longer than wide (1.12 X 0.90 mm), widest in anterior third and bearing many small spinules laterally.

Abdomen 2-segmented; first segment 0.45 mm long,

second segment 0.64 mm long. Caudal ramus (Figure 129) about twice as long as wide (560 \i X 295 \i),

and armed as in the figure.

Cephalic appendages as in female except as noted

below. Second antenna (Figure 130) with stout claw

bearing prominent accessory claw, inner surface of

basal segment with prominent adhesion pad, outer

edge with rugose area. Maxilliped (Figure 131) basal

segment with raised area covered with blunt spinules

as indicated in the figure.

Legs 1—4 as in female except for modified spines

on exopod of leg 2 (Figure 132).

ETYMOLOGY.—The name pacificus refers to the

geographic area to which this species is restricted.

REMARKS.—Females of this species can be separated from all other species except A. elegans and

lobatus by the shape of the genital segment (lack of

prominent lateral bulge). It can be separated from

these two species by the length of the fifth leg process

which extends well beyond the caudal rami in A.

elegans and lobatus and does not in pacificus. When

the spermatophore is attached it can be separated

from all species by the nature of its form. The pattern

of the canaliculations on the surface of the postoral

adhesion pad further serves to separate both sexes of

this species from all others. Males can be separated

by the nature of the modified spines on the exopod of

leg 2.

On the basis of its morphology it seems most

closely related to A. carchariae.

This species is probably endemic to the eastern

Pacific.

Literature Cited

Barnard, K. H.

1955. South African Parasitic Copepoda. Annals of the

South African Museum, 41 (5): 223-312.

Bassett-Smith, P. W.

1898. Some New or Rare Parasitic Copepods Found on

Fish in the Indo-Tropic Region. Annals and

Magazine of Natural History, 2(7) :359-372.

1899. A Systematic Description of Parasitic Copepoda

found on Fishes, with an Enumeration of the

Known Species. Proceedings of the Zoological

Society of London, 2:438-507.

Beneden, P. J. van.

1892. Quelques Nouveaux Caligides de la Cote

d'Afrique, et de l'Archepel des Azores. Bulletins

de VAcademie Royale des Sciences, des Lettres et

des Beaux-arts de Belgique, 24(3) :241-262.

Bere, R.

1936. Parasitic Copepods from Gulf of Mexico Fish.

The American Midland Naturalist, 17(3):577-

625.

Brady, G. S.

1883. Report on the Copepoda Collected by H.M.S.

Challenger during the years 1873-76. Part 23 of

Volume 8 in Zoology in Report on the Scientific

Results of the Voyage of H.M.S. Challenger,

1873-76.

Brian, A.

1908. Note Preliminaire sur les Copepodes Parasites des

poissons Provenant des Campagnes Scientifiques

de S.A.S. le Prince Albert Ier de Monaco ou

Deposes dans les Collections du Musee Oceanographie. Bulletin de I'Institut Oceanographique,

110:1-19.

1912. Copepodes Parasites des Poissons et des fichinides

Provenant des Campagnes Scientifiques de S.A.S.

le Prince Albert Ier de Monaco. Re suit at s des

Campagnes Scientifique, 38:1-58.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Capart, A.

1953. Quelques Copepodes Parasites de Poissons Marins

de la Region de Dakar. Bulletin de I'Institut

francais d'Afrique noire. 15(2):647—671.

Carvalho, J. P.

1940. Notas Sobre alguns Caligoida com a Descricao de

Pandarus marcusi, sp. nov. Universidade de Sao

Paulo. Boletins de Faculdade de Filosofia, Ciencias 1 Letras XIX Zoologia, 4:271-302.

1951. Notas Sobre alguns Cope'podos Parasitos de Piexes

Maritimos da Costa do Estado de Sao Paulo.

Boletim do Instituto Paulista de Oceanografia,

2(2):135-144.

Cressey, R. F.

1967. Caligoid Copepods Parasitic on Sharks of the

Indian Ocean. Proceedings of the United States

National Museum, 121 (3572) : 1-21.

1970. Copepods Parasitic on Sharks from the West

Coast of Florida. Smithsonian Contributions to

Zoology, 38:1-30.

Fowler, H. W.

1912. The Crustacea of New Jersey. Annual Report.

New Jersey State Museum with a report of the

Crustacea of New Jersey, pages 9-650.

Gnanamuthu, C. P.

1951. New Copepod Parasites of Sharks. Annals and

Magazine of Natural History, 4( 12): 1236-1256.

Heegaard, P.

1955. Parasitic Copepods from Tropical West Africa.

Atlantide Report No. 3. Scientific Results of the

Danish Expedition to the Coasts of Tropical West

Africa, 1945-1946, pages 41-56.

1962. Parasitic Copepoda from Australian Waters.

Records of the Australian Museum, 25(9): 149-

234.

Kirtesinghe, P.

1956. Parasitic Copepods of Fish from Ceylon. IV.

Parasitology, 46(3): 14-21.

1964. A Review of the Parasitic Copepods of Fish recorded from Ceylon with Description of Additional Forms. Bulletin of the Fisheries Research

Station, Ceylon, 17(1) :45-132.

Kreyer, H.

1863. Bidrag til kundskab om snyltekrebsene. Naturhistorisk Tidsskrift, 3(2): 75-426.

Lewis, A. G.

1966. Copepod Crustaceans Parasitic on Elasmobranch

Fishes of the Hawaiian Islands. Proceedings of

the United States National Museum, 118(3524):

57-154.

Pillai, K.

1967. Copepods Parasitic on Indian Fishes—A Review.

Proceedings of Symposium on Crustacea, 5:1556—

1680.

Rangnekar, M. P.

1959. Parasitic Copepods from Fishes of the Western

Coast of India with Description of One New and

Redescription of Four Known Species. Journal of

the University of Bombay, 28(3):43-58.

Rao, T. S. S.

1950. On a New Caligid Parasite from the Indian

Hammerhead Shark. Proceedings of the Indian

Academy of Sciences, 31(5) section B: 302-307.

Rathbun, M. J.

1905. Fauna of New England. 5. List of the Crustacea.

Occasional Papers of the Boston Society of Natural History, VII, pages 1-117.

Shiino, S. M.

1955. Alebion echinatus Capart from Japanese Waters,

with Observations on the Newly Found Male

Form. Pacific Science, 9(2): 177-182.

1959. Ostpazifische Parasitierende Copepoden. Report

of the Faculty of Fisheries, Prefectural University

of Mie, 3(2): 267-333.

1965. Parasitic Copepods of the Eastern Pacific Fishes

6. Euryphoridae. Report of Faculty of Fisheries,

Prefectural University of Mie, 5(2) :421-433.

Vaissiere, R.

1959. Parasites de Poissons de mer ouest-africains recolte's par J. Cadenet. II. Copepodes. Bulletin

de I'Institut Francois Afrique Noire, 21, series A

(2):534-553.

Wilson, C. B.

1905. New Species of Parasitic Copepods from the

Massachusetts Coast. Proceedings of the Biological Society of Washington, 18(20): 127-132.

1907. North American Parasitic Copepods Belonging to

the Family Caligidae. Part 2. The Trebinae and

Euryphorinae. Proceedings of the United States

National Museum, 31 (1504): 669-720.

1921. New Species and a New Genus of Parasitic Copepods. Proceedings of the United States National

Museum, 59(2354): 1-17.

1932. The Copepods of the Woods Hole Region, Massachusetts. United States National Museum Bulletin, 158:1-635.

Yamaguti, S.

1963. Parasitic Copepods and Branchuria of Fishes.

Interscience Publications, pages 1-1104. New

York: John Wiley and Sons.
NUMBE R 12 3 11

FIGURES 1-6.—Alebion carchariae Kreyer, female: 1, dorsal; 2, genital segment, ventral;

3, spermatophores attached; 4, spermatophores attached; 5, caudal ramus, ventral; 6, first

antenna.
12

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 7-13.—Alebion carchariae Kroyer, female: 7, second antenna; 8, tip of mouth tube;

9, postoral adhesion pads and mouth tube; 10, second maxilla; 11, row of setules on second

maxilla; 12, flagella of second maxilla; 13, maxilliped.
NUMBE R 12 3 13

18

20

14

FIGURES 14-20.—Alebion car chariot Kreyer, female: 14, leg 1; 15, last exopod segment of

leg 1; 16, leg 2; 17, leg 3; 18, exopod of leg 3; 19, leg 4; 20, tip of leg 5.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

26 25

FIGURES 21-28.—Alebion carchariae Kroyer, male: 21, dorsal; 22, genital segment and abdomen; 23, ventral edge of genital segment; 24, caudal ramus, ventral; 25, second antenna;

26, maxilliped; 27, exopod spine of leg 1; 28, exopod of leg 2.
NUMBE R 12 3 15

34

FIGURES 29-35.—Alebion maculatus Wilson, female: 29, dorsal; 30, spermatophores attached;

31, caudal ramus, ventral; 32, second antenna; 33, short flagellurn of second maxilla; 34, postoral adhesion pad; 35, maxilliped.
16 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 36-43.—Alebion maculatus Wilson, female: 36, last exopod segment of leg 1: 37, tip

of leg 5. Male: 38, dorsal; 39, genital segment and abdomen; 40, caudal ramus. ventral; 41,

second antenna; 42, maxilliped; 43, exopod of leg 2.
NUMBE R 12 3 17

46

FIGURES 44—50.—Alebion crassus Wilson, female: 44, dorsal; 45, genital segment ventral;

46, caudal ramus, ventral; 47, second maxilla, area near flagella; 48, postoral adhesion pad;

49, maxilliped; 50, tip of claw of maxilliped.
18 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

51

57

FIGURES 51-57.—Alebion crassus Wilson, female: 51, last exopod segment of leg 1. Male: 52,

dorsal; 53, genital segment and abdomen, ventral; 54, caudal ramus; 55, second antenna;

56, maxilliped; 57, exopod of leg 2.
NUMBE R 12 3 19

FIGURES 58-65.—Alebion elegans Capart, female: 58, dorsal; 59, genital segment and abdomen,

ventral; 60, spermatophores attached; 61, caudal ramus, ventral; 62, second antenna; 63, tip

of short flagellum of second maxilla; 64, postoral adhesion pad; 65, maxilliped.
20 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

67

FIGURES 66-73.—Alebion elegans Capart, female: 66, last exopod segment of leg 1; 67, tip of

leg 5. Male: 68, dorsal; 69, genital segment and abdomen, ventral; 70, caudal ramus, ventral;

71, second antenna; 72, maxilliped; 73, exopod of leg 2.
NUMBE R 12 3 21

FIGURES 74—80.—Alebion glaber Wilson, female: 74, dorsal; 75, spermatophores attached;

76, caudal ramus, ventral; 77, claw of second antenna; 78, tip of short flagellum of second

maxilla; 79, maxilliped; 80, postoral adhesion pad.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURES 81-87.—Alebion glaber Wilson, female: 81, last exopod segment of leg 1. Male:

82, dorsal; 83, genital segment and abdomen; 84, caudal ramus, ventral; 85, second antenna;

86, maxilliped; 87, exopod of leg 2.
NUMBE R 12 3 23

92

FIGURES 88-94.—Alebion gracilis Wilson, female: 88, dorsal; 89, genital segment and abdomen,

ventral; 90, caudal ramus. ventral; 91, tip of short flagellum of second maxilla; 92, postoral

adhesion pad; 93, maxilliped; 94, last exopod segment of leg 1.
24 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

95

FIGURES 95-99.—Alebion gracilis Wilson, male: 95, dorsal; 96, genital segment and abdomen,

ventral; 97, caudal ramus; 98, second antenna; 99, exopod of leg 2.
NUMBE R 12 3 25

103

102

104

FIGURES 100-104.—Alebion lobatus Cressey, female: 100, dorsal; 101, genital segment and

abdomen, ventral; 102, spermatophores attached; 103, posterior portion of genital segment and

abdomen, ventral; 104, postoral adhesion pad.
26 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

105

110

108

106

FIGURES 105-110.—Alebion lobatus Cressey, male: 105, dorsal; 106, genital segment and abdomen, ventral; 107, caudal ramus; 108, second antenna; 109, maxilliped; 110, exopod of leg 2.
NUMBE R 12 3

27

115

FIGURES 111—116.—Alebion pacificus, new species, female: 111, dorsal; 112, genital segment

and abdomen, ventral; 113, spermatophores; 114, caudal ramus. ventral; 115, last segment

of first antenna; 116, second antenna.

113
28 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

125

FIGURES 117-125.—Alebion pacificus, new species, female: 117, second maxilla; 118, row of

setules on second maxilla; 119, tip of short flagellum of second maxilla; 120, postoral process;

121, maxilliped; 122, leg 1; 123, last exopod segment of leg 1; 124, leg 2; 125, leg 3.
NUMBE R 12 3 29

126

FIGURES 126-132.—Alebion pacificus, new species, female: 126, tip of leg 5. Male: 127, dorsal;

128, genital segment and abdomen, ventral; 129, caudal ramus, ventral; 130, second antenna;

131, maxilliped; 132, exopod of leg 2.

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