-

SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM

PROCEEDINGS

OF THE

UNITED STATES NATIONAL MUSEUM

VOLUME 121

NUMBERS 3568-3580

SMITHSONIAN PRESS
WASHINGTON : 1967

Publications of the United States National Museum

The scientific publications of the United States National Museum include two
series, Proceedings of the United States National Museum and United States
National Museum Bulletins.

In these series are published original articles and monographs dealing with
the collections and work of the Museum and setting forth newly acquired facts
in the fields of anthropology, biology, geology, history, and technology. Copies
of each publication are distributed to libraries and scientific organizations and
to specialists and others interested in the various subjects.

The Proceedings, begun in 1878, are intended for the publication, in separate
form, of shorter papers. These are gathered in volumes, octavo in size, with
the publication date of each paper recorded in the table of contents of the volume.

In the Bulletin series, the first of which was issued in 1875, appear longer,
separate publications consisting of monographs (occasionally in several parts)
and volumes in which are collected works on related subjects. Bulletins are
either octavo or quarto in size, depending on the needs of the presentation.
Since 1902, papers relating to the botanical collections of the Museum have been
published in the Bulletin series under the heading Contributions from the United

States National Herbarium.
Frank A. TAYLOR

Director, United States National Museum
II

CONTENTS

BarnarpD, J. Laurens. New species and records of Pacific
Ampeliscidae (Crustacea: Amphipoda). 20 pages, 4 figures.
Published March 8, 1967. . ‘

New species: Byblis albatrossae, B. ampelisciformis, B. orientalis.

Cressey, Roger. Revision of the family Pandaridae (Cope-
poda: Caligoida). 133 pages, 356 figures. Published
February 16, 1967. en fee a vee

New genus: Pannosus.
New species: Pandarus floridanus, P. katoi, Dinemoura discrepans,
Nesippus tigris.

Cressey, Roger. Caligoid copepods parasitic on sharks of
the Indian Ocean. 21 pages, 54 figures, 2 plates. Published
March 15, 1967. saretbst repens nach.

New species: Nemesis aggregatus, Eudactylina pusilla, E. pollex,
Kroyeria gemursa.

Humes, Artuur G., and Ho, Ju-Sury. New cyclopoid
copepods associated with the alcyonarian coral Tubipora
musica (Linnaeus) in Madagascar. 24 pages, 69 figures.
Published March 8, 1967. . TAS RWNENOF te soe he

New species: Lichomolgus organicus, L. conjunctus, Hippomolgus
latipes, H. cognatus.

Krer, Porter. Bredin-Archbold-Smithsonian Biological Sur-
vey of Dominica: 1, The echinoids of Dominica. 10 pages,
3 figures, 2 plates. Published December 30, 1966 .

Kornicker, Louis S. Supplementary descriptions of two
myodocopid ostracods from the Red Sea. 18 pages, 6 figures.
Published February 16, 1967 .

Kornicxer, Louis S. The myodocopid ostracod families
Philomedidae and Pseudophilomedidae (new family). 35
pages, 12 figures, 1 plate. Published February 16, 1967 .

New family: Pseudophilomedidae.
New combination: Huphilomedes asper.

Lewis, Attan G. Copepod crustaceans parasitic on teleost
fishes of the Hawaiian Islands. 204 pages, 70 figures. Pub-
lished June 22, 1967. hh bte walls. Chucecsmrey eee

New species: Pseudotaeniacanthus puhi, Anchistrotos moa, Hat-
schekia breviramus, Lepeophtheirus? fallolunulus, Caligus kapuhili,
Brachiella regia.

III

Number

3576

3570

3572

3573

3577

3571

3580

3574

IV CONTENTS

McFappEen, Max W. Soldier fly larvae in America north of
Mexico. 72 pages, 156 figures. Published February 1,
POG TS Ire Se ot cao Ps, Sepang

Perripons, Marian H. Some bathyal polynoids from Central
and Northeastern Pacific (Polychaeta: Polynoidae). 15
pages, 5 figures. Published February 16, 1967 .

New genera: Bathyadmetella, Bathymoorea.
New species: Bathyadmetella commando.
New combination: Bathymoorea renotubulata.

Quate, LAURENCE W., and THompson, SaraH E. Revision of
click beetles of genus Melanotus in America north of Mexico
(Coleoptera: Elateridae). 83 pages, 12 figures, 1 plate.
Published April 18, 1967.

New species: Melanotus indistinctus, M. diesen. M. ‘iessie lite
M. lanei, M. beameri, M. lanceatus.

Reuper, Haraup A. Valid zoological names of the Portland

Catalogue. 51 pages, 2 figures. Published March 21, 1967 .

Stone, AuAN. Bredin-Archbold-Smithsonian Biological Sur-
vey of Dominica: 2, New species of Diptera from Dominica
(Anisopodidae and Bibionidae). 6 pages, 2 figures. Pub-
lished December 30, 1966 . EN

New species: Mycetobia limanda, Olbiegaseer “ddniitie O. evanst,
Plecia porca.

U.S. GOVERNMENT PRINTING OFFICE 262-119-h

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Proceedings of
the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3568

REVISION OF CLICK BEETLES OF GENUS MELANOTUS
IN AMERICA NORTH OF MEXICO
(COLEOPTERA: ELATERIDAE)

By Laurence W. QuaTe and Saran E. THomeson!

Click beetles and wireworms of the genus Melanotus are common
elements in the North American fauna. Some of the species are
important to agriculture because the larvae are destructive to seeds
and roots of corns, small grains, grasses, and some root crops; in
spite of their importance, the North American species of the genus
have never been revised. Early descriptions by LeConte (18538,
1866), Say (1823, etc.), Melsheimer (1846), Erichson (1842), and
others are of little value in identifying the species. General treat-
ments of local faunas (Blatchley, 1910; Dietrich, 1945; Brooks, 1960)
are improvements but generally insufficient for identification except
within a limited area. Descriptions of a few new species have ap-
peared recently (Knull, 1959, 1962) and have increased the need for
a thorough revision. M. C. Lane is the taxonomist most qualified to
do this, having identified most North American collections of Me-
lanotus in the past several decades, but the pressure of other duties
has prevented him from committing his wide knowledge to print.

1Quate: B. P. Bishop Museum, Honolulu, Hawaii; Thompson: State Board of
Health, Raleigh, N.C.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

It was on the recommendation of Mr. Lane that in 1955 the senior
author began a revision of North American Melanotus. Five years
ago he was joined by the junior author. During most of this time,
the work has been done during spare time by both.

The new species were recognized early in the study by the senior
author and authorships are to be credited to him.

We have arrived at the following conclusions concerning the tax-
onomy of the genus: the features distinguishing one species from the
other do not differ greatly; typical specimens of each species are
identifiable on the basis of external characters; some specimens of
each species vary enough to obscure the gap between the taxa; positive
identification can be made by the study of the genitalia, which is
specific in both sexes; and a natural division of the genus into sub-
generic taxa is not apparent to us at this time.

One of the most striking features of the zoogeography of North
American Melanotus is that the species are concentrated east of the
100th meridian. Melanotus in North America is predominantly an
eastern genus (fig. 1). It is not within the scope of this paper to
dwell at length on possible causes of this distributional pattern, but
we may discuss it briefly. The apparent relationship of New and
Old World faunas suggests that there have been migrations over a
Bering Straits land bridge. We may further assume that the Mela-
notus click beetles which came from Eurasia and crossed the bridge
stayed on the eastern side of the Rocky Mountains. This might
have been the origin of the distributional pattern we see today.
Adaptation to the more humid climate and edaphic conditions pre-
vailing in the East may be responsible for the maintenance of the
pattern.

In western North America there are only a few Melanotus and
most of these are in the Southwest. There appears to be a group of
species—longulus, hamatus, lanceatus, beamert, and concisus—which
are derived from a single ancestor. Perhaps it was an offshoot from
the main group in the East. At any rate, speciation has occurred
in the Southwest where we now find the five species, and one of these
has spread northward through the Great Basin and along the West
Coast to Canada. In the northern part of its range it has further
differentiated into a separate subspecies.

The two most distinctive species of North American Melanotus also
are found in the Southwest. These are cribricollis and chiricahuae,
and they are so distinct from others that we presume they are a
Neotropical element.

The practical effect of these distributional patterns is that iden-
tification of Melanotus in the Far West is simple since there is only a
single species with two subspecies; in the Southwest there are only a

NO. 3568 MELANOTUS—QUATE AND THOMPSON 3

few species with which to contend, so identification of that Melanotus
fauna is not difficult. The bulk of the Melanotus are in the eastern
half of the United States, and this is where the greatest problems of
identification will be encountered.

It appears that nearly all American species of Melanotus north of
Mexico are now known. Supporting this conclusion is the fact that
of the 46 Nearctic Melanotus only nine (20 percent) are named in
this paper or have been named in the last few years, and half of these
have been recognized for at least 20 years. What few species may
remain undiscovered are probably in the Southwest.

We have made an artificial division of Melanotus into three groups.
The first two, containing nearly all the species, are based on the

Ficure 1.—Distribution of Melanotus in the United States (figures indicate the number
of species recorded in each state or area).

presence or absence of mandibular pits. This is an obvious character
and it simplifies identification; however, we do not feel that this
single character indicates natural units, and the taxa based on it are
merely utilitarian. The third group, composed of only cribricollis
and chiricahuae, is characterized by large pronotal punctures and
reduced parameres in the male genitalia; it is probably monophyletic.

A key has been prepared which will simplify identification of most
specimens. The greatest difficulty will be individual variants that
do not seem to quite fit either part of a couplet. Many revisions have
failed to eliminate all deficiencies, but for the majority of specimens
the key should provide a satisfactory shortcut to their proper
identification.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

The authors have been able to study all but six of the extant types.
The locations of the types have been indicated in the appropriate
places. Museums have been abbreviated thus: U.S. National Mu-
seum (USNM); Museum of Comparative Zoology, Harvard Univer-
sity (MCZ); Purdue University (PU); Cornell University (CU); Uni-
versity of Kansas (KU); California Academy of Sciences (CAS);
British Museum (Natural History) (BMNH); Zoologisches Museum
der Humboldt-Universitat zu Berlin (ZM).

In addition to the types, over 13,000 specimens were examined by
us during the course of this study.

Ficure 2.—Distribution of the three types of female bursae of Melanotus similis.
© normal © intermediate © spiny

Morphological terms used by us are common ones, but those which
might be unclear are defined. The ‘front’ is the flat area between
and above the eyes, bounded anteriorly by the “frontal margin.”” Be-
low the frontal margin and occupying the central area between the
antennae is the ‘“‘nasale.’”’ On either side of the nasale and between
it and the antennal bases is a pair of pits, the ‘“‘parantennal foveae.”
Within some of the larger foveae is an elevated part that gives them
an earlike appearance; the raised part is termed the “foveal tragus.”’
The ‘mandibular pit” is a clearly marked depression of varying size
on the lateral, exposed margin of the mandible.

The shape and measurements of the pronotum are important tax-
onomic features. The length is measured along the midline and does

NO. 3568 MELANOTUS—QUATE AND THOMPSON 5

not include the projection of posterolateral “hind angles.” The width
is measured as the widest part of the pronotum before the hind angles
and is not affected by their expansion if they are wider than the rest
of the prothorax. The relation of the antennal length to the pronotum
used in the descriptions refers to the distance before or beyond the tip
of the hind angles, which the antenna reaches when stretched along
and parallel to the side of the pronotum.

As an indication of size, we have measured only the length of the
elytra. This is more accurately measured than the whole length of
the insect and is as useful as an indication of its overall dimension.

The genitalia of both sexes are important taxonomically, but the
parts are simple and easily defined. Following the terminology of
Snodgrass (1957), the male genitalia consist of a central ‘‘aedeagus”
(median lobe) which is flanked by a pair of ‘‘parameres” (lateral lobes,

Figure 3.—Distribution of species.
O Melanotus longulus longulus
@ M. I. oregonensis

gonostyli). These are supported by the ‘phallobase.” The para-
mere may be plain at the tip or expanded into an “apical blade.” The
taxonomically important parts of the female genitalia (fig. 5) are the
large membranous sac, the ‘‘bursa,’’ which contains spines of various
shapes and number, and the “accessory gland” and ‘“‘spermathecal
duct,’ which arise from the bursa. A more detailed morphological
description was given by Becker (1956); he also described the tech-
niques used in dissecting the female genitalia. It is only necessary to
add that light reflected from the substage mirror of a stereoptic micro-
scope is helpful in seeing the delicate spermathecal duct, its divertic-
ulum, and accessory gland.

Bibliographic references are intended to be complete; catalog refer-
ences (Leng, 1920; Schenkling, 1927) are given only when new infor-
mation is involved.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Because of the large number of records involved, distributional
data have been reduced to listing states and counties (or locales and
provinces) with the earliest and latest months of collections for each
species. Only complete collecting data are given for the new species
and the two subspecies.

M. C. Lane, Collaborator, Agricultural Research Service, USDA,
has given invaluable assistance to this study. He permitted us to
study specimens in his collection, allowed the use of his notes of
type specimens, loaned numerous identified specimens, and identified
many of the species in the early stages of this work.

Grants from the National Academy of Sciences (Marsh Fund Grant
no. 130, 1960) and the American Philosophical Society (Grant 2614,
Penrose Fund, 1960) permitted the senior author to study types in the
Zoologisches Museum der Humboldt-Universitét zu Berlin and the
British Museum (Natural History). Another grant by the National
Academy of Sciences (Marsh Fund) enabled the junior author to
study Melanotus types at Purdue University and the Museum of
Comparative Zoology, Harvard University. While a member of the
faculty, the senior author was provided funds by the University of
Nebraska Research Council to study the Lane Collection in Walla
Walla, Wash.

Through the kindness of Prof. C. H. Lindroth and K. J. Hedquist,
we were able to receive specimens of M. communis from the Gyllenhal
Collection at the Swedish Museum of Natural History, Stockholm.
While studying type specimens at various times, we enjoyed the help
and hospitality of Dr. K. Delkeskamp, Humboldt-Universitat, Miss
C. M. F. von Hayek, British University; Dr. P. J. Darlington, Jr.,
Museum of Comparative Zoology; and Dr. Leland Chandler, Purdue
University.

The large collections of the U.S. National Museum have been vital
to our studies. We are indebted to the officials of the Smithsonian
Institution and T. J. Spilman, USDA, for the privilege of studying
the material.

We also thank the following for the loan of specimens: F. G. Werner,
University of Arizona; H. B. Leech, California Academy of Sciences;
P. D. Hurd, Jr., California Insect Survey; E. C. Becker, Canadian
National Collection; H. Dietrich, Cornell University; H. O. Lund,
University of Georgia; M. W. Sanderson and L. K. Gloyd, Illinois
State Natural History Survey; G. W. Byers, University of Kansas;
W. Enns, University of Missouri; D. A. Young, North Carolina
State College; W. C. Stehr, Ohio State University; H. R. Burke,
Texas Agricultural Experiment Station; and C. D. Fronk, University
of Wyoming.

NO. 3568 MELANOTUS—QUATE AND THOMPSON

List of North American Melanotus

SIMILIS GROUP
(Mandible without pit)

1. similis (Kirby) 9. verberans (LeConte)

2. spadix (Krichson) 10. emissus (LeConte)

3. decumanus (Erichson) 11. lanei, new species

4. castanipes (Paykull) 12. pilosus Blatchley

5. communis (Gyllenhal) 13. opacicollis (LeConte)
6. indistinctus, new species 14. clandestinus (Erichson)
7. dietrichi, new species 15. ignobilis Melsheimer
8. miscellus, new species

AMERICANUS GROUP
(Mandible with pit)

16. depressus (Melsheimer) 31. parallelus Blatchley
17. morosus Candéze 32. americanus (Herbst)
18. cribulosus (LeConte) 33. cribriventris Blatchley
19. corticinus (Say) 34. obscuratus Blatchley
20. sagittarius (LeConte) 35. beameri, new species
21. hyslopi van Zwaluwenburg 36. concisus Knull

22. prasinus Blatchley 37. lanceatus, new species

23. piceatus Blatchley 28) hamatis Kull
Baa ie Elrenle, pS eR EO

25. macer (LeConte)
26. testaceus (Melsheimer)
27. trapezoideus (LeConte)

39. longulus longulus (LeConte)
40. Ll. oregonensis (LeConte)
41. gradatus (LeConte)

28. tenax (Say) 42. insipiens (Say)
29. pertinax (Say) 43. leonardi (LeConte)
30. infaustus (LeConte) 44. taenicollis (LeConte)

CRIBRICOLLIS GROUP

45. cribricollis Candéze 46. chiricahuae Knull

List of Synonymical Names in North American Melanotus

abdominalis (Erichson) = 2. spadiz (Erichson)
angustatus (Erichson) =32. americanus (Herbst)
blatchleyi Leng=33. cribriventris Blatchley
canadensis Candéze=3. decumanus (Erichson)
carinus Blatchley= 20. sagittarius (LeConte)
cuneatus (LeConte) =3. decumanus (Erichson)
debilis Blatchley=17. morosus Candéze
divarcarinus Blatchley=16. depressus (Melsheimer)
dubius (LeConte) = 26. testaceus (Melsheimer)
exuberans (LeConte) =1. similis (Kirby)

fissilis (Say) =1. similis (Kirby)

fransiscanus Van Dyke=40. longulus oregonensis (LeConte)
glandicolor (Melsheimer) = 19. corticinus (Say)
inaequalis (LeConte) =4. castanipes (Paykull)

8 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

incertus (LeConte) =3. decumanus (Erichson)

laticollis (Erichson) =1. similis (Kirby)

lixus Blatchley=16. depressus (Melsheimer)

longicornis Blatchley=33. cribriventris Blatchley
longulus (LeConte) =17. morosus Candéze
ochraceipennis Melsheimer=1. stmilis (Kirby)

paganus Candéze=5. communis (Gyllenhal)

paradoxus (Melsheimer) =4. castanipes (Paykull)
parumpunctatus (Melsheimer) = 16. depressus (Melsheimer)
peninsularis Candéze= 14. clandestinus (Erichson)
perplexus Blatchley=14. clandestinus (Erichson)
prolizus (Erichson)=1. similis (Kirby)

scrobricollis (LeConte) =4. castanipes (Paykull)
secretus (LeConte) =15. ignobilis Melsheimer

simulans Blatchley=17. morosus Candéze

sphenoidalis (Melsheimer) = 1. similis (Kirby)

tenellus (Erichson) = 42. insipiens (Say)

tecanus Candéze=4. castanipes (Paykull)

variolatus LeConte=40. longulus oregonensis (LeConte)
vetulus (Erichson) =19. corticinus (Say)

Melanotus Eschscholtz

Melanotus Eschscholtz, 1829, p. 32. [For additional references, see Schenkling,
1927, p. 271.]

Perimecus Stephens, 1830, p. 263 [Type: Elater fulvipes Herbst, by monotypy.]

Ctenonychus Stephens, 1830, p. 272 [Type: Ctenonychus hirsutus Stephens, by
monotypy.]

Cratonychus Dejean, 1833, p. 87. [Type: Elater obscurus Oliver, designated by
Blanchard, 1845, p. 76.]

Type species: Hlater fulvipes Herbst, designated by Westwood,
1840, p. 26.

Adult (partly after Lane, in litt.): Color usually uniformly yel-
lowish brown to dark reddish brown, only 2 species bicolored.
Head with front coarsely punctate, punctures hexagonal and sep-
arated by less than own diameter; frontal margin complete, carinate
(sometimes obsolescent); parantennal fovea and nasale variously
developed, but well defined (nasale indistinct in few species). An-
tenna 11-segmented; segment 1 elongate and cylindrical, 2 spherical,
3 variable from spherical and equal size of 2 to much longer and
triangular, 4 and following triangular; male only with dense, erect
hairs on mesal margin. Palpus 3-segmented, light reddish brown
and lighter than rest of body.

Pronotal punctures present, but size and distribution variable;
hind angles well developed and carinate. Elytron with 9 striae
formed by rows of punctures and rows of quadrate, subcuticular
spots under striae. No prothoracic groove for receiving antenna;
meso- and metasternum not meeting to enclose mesocoxal cavities,
mesocoxae separated, metasternum truncate anteriorly. Tibiae with

NO. 3568 MELANOTUS—QUATE AND THOMPSON 9

apical spurs; tarsal claws pectinate; tarsi simple, without lobes.
Punctures on venter of body teardrop shaped.

Key to North American Species of Melanotus

1. Pronotum and elytron of different colors, pronovum partly or completely
redaish orange, and ely tron: DIACK jn) sen io sey co a as eye ee oe 2
Pronotum and elytron of same color ... . dst bo

2(1). Pronotum almost entirely orange and nodomont ouisnelle pea antenna
of o& usually exceeding tip of hind angle by 1 segment or less, 9
antenna ending about at tip... . . .. . . 43. leonardi
Pronotum black in center and orange ones and abdominal venter

partly or entirely orange; antenna of o ending before tip of hind

angle by about 1 segment and lessinQ. . ... . . 44. taenicollis

3(1). Antennal segments 4 and following much longer than wide; antenna
generally extends to or beyond hind angle; not with following combi-

ANOMS 1G oc ch eSahoh ster sae
Antennal mene 4 aad meas as pollowine iden than iene and lobes
lighter in color than rest of antenna; antenna ends 2 or more segments

before tip of hind angle; frontal eee slightly indented in center,

when viewed from front curves down, and hence nasale is only nar-

row space between frontal margin and clypeus; frontal and pronotal

punctures very large; Arizona to Baja California . . 45. cribricollis
4(3). Mandible without pit on lateral, exposed margin. ........ 5
Mandible, with pit om lateral margin...) 2 ee OL

Simitis Group
(Mandible without pit on lateral margin)

5(4). Antennal segment 3 small, ise to 2 or at least much nearer size of
Zthanvay Gee. s Pee, SG
Antennal segment 3 infemmetitie in size petwedd 2 ao i. clearly larger

aM aay rs oxy. te, eee hee Mota, Rakle: respi cr nia cee Os

6(5). Antennal segments beyond 3 intone same color as body, triangular,
erect male hairs well developed. . .. . Bait puies oeatd)
Antennal segments beyond 3 testaceous, Genter: in Peolon than rest of

body, subquadrate, erect male hairs poorly developed; sides of pro-

notum with marked concavity before hind angles, angles rather small

and not widely divergent; @ paramere incurved near center and ta-

pering to point, without apical blade ...... . . 15. ignobilis

7(6). Smaller species, elytron less than 8.0 mm; frontal margin thin and
jutting over nasale; nasale and parantennal fovea obsolescent; sides

of pronotum with concavity before hind angles, angles small; ¢@
paramere with apical blade. ... . .... . 14. clandestinus

Large species, elytron more than 9.0 mm; coftonbal margin not strongly
jutting over nasale; nasale and op tan sane fovea poorly developed,

but usually present; sides of pronotum often straight and divergent
posteriorly with nak hind angles; co paramere with apical
bladet.anew 2). is i .... . 4 castanipes

8(5). Frontal and prenntal prn a reed enlarged; punctures on
venter of abdomen oblong and lateral ones but little larger than

central ones; co paramere little shorter than aedeagus and with

AP Cig i acle nrc al ot eee par tae eC a iirats ve nsldideathatl ee we, woe O

10

9(8).

10(9).

11(10).

12(11).

13(11).

14(13).

15(14).

16(15).

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Frontal and pronotal punctures very large (fig. 10/); punctures on
venter of abdomen teardrop shaped and lateral ones several times
larger than central ones; co’ paramere about two-thirds length of
aedeagus and abruptly tapering to sbarp, recurved apex; New Mexico
to: Baja California «0. s):éekio 2 his oe, Ys A eharicahuae

Pronotum wider than long . . . Aeeat neater tered O)

Pronotum narrow, longer than wide (gssareas in te bit of hind angles) ;
frontal margin scoop shaped, jutting and a little upturned.

31. parallelus (part)

Pronotal punctures not so thick as to appear granulose; vestiture of
thorax and elytra of about same length and density. ...... Il

Pronotal punctures very dense and appearing granulose; vestiture of
pronotum short and fine, nearly velvety, considerably denser than
Onielytraw en mane ior, b meen Os «. % tel oskODaclCOlls

Pronotal punctures smaller tae figure on frons, dense and nearly con-
tiguous on all sides; small species, elytron length less than 7.6mm . 12

Pronotal punctures sparser and ee most ae with elytron length
Oo) OT MOre ne. ss aos te veiw

Vestiture of thorax rather sparse, tet on tee Se sparser than on dorsum;
antennal segment 3 intermediate in size between 2 and 4; frontal
margin not upturned. .... Sure ae 1. lanei

Vestiture of thorax long and einer oe on Co as long and thick
as on dorsum; antennal segment 3 shorter, closer to length of 2 than

4; frontal margin thin, shelflike and alittle upturned . . . 12. pilosus
Frontal margin not strongly protuberant, usually without a depression
before margin. . . ad pesca alee aA

Frontal margin prone pred aoe ened om side extends
out in shelflike extension, marked depression behind margin; ? bursa
without spines or with only patch of 5-8 small spines on each side
near base; o' paramere with margins before apical blade straight
and without indentation; large species, elytron 10-12mm.

3. decumanus

Frontal margin well defined, dark brown or black, not weakened in
center; elytron rarely as long as 13 mm and most species considerably
leSSixite) 222 4 sieehos <eee, 10

Frontal margin Senne weneoned’ in acaniee when viewed directly in
front appears obliterated and without black margin in center; large
species with elytron 11.4-14.9 mm (aver. 13 mm); ? bursa with spines
in large basal packet and small, separated distal patch (fig. 5f).

4. castanipes, females

Antennal segment 4 about 1% times as long as wide and considerably wider
than 3; elytron often less than 10 mmlong ... . ke! vice ibe eO

econ segment 4 usually slender, about two times as lone as wide and
only a little wider than 3; average elytron length 10.8 mm; & gono-
style with thick patch of 30-40 long, soft hairs on apical Bide (fig. le);
? bursa with dense packet of long, close-set spines arranged nearly in
trapezoid: (fig. 5¢) -. 3% 2 kik ‘ ‘ ee a SpaGix

Pronotal punctures smaller than Pee on raconte ieee | in center generally
separated by at least own diameter; elytron usually less than 8 mm
long; o aedeagus parallel sided until near apex and sharply tapering
tocacute tip’. |. 122%0)L) Ge) eS se ers ts eee

NO. 3568

17(16).

18(16).

19(18).

20(18).

21(4).

22(21).

23(22).

24(22).

25(24).

26(25).

MELANOTUS—QUATE AND THOMPSON ri

Pronotal punctures subequal in size to those on front; elytron usually
more than 8 mm; < apne often with convergent sides and tapering
tottipyies “la esate RLS

Antenna same pole as peed end eee e aanenmers ae tapering to
apical blade; outer margin of apical blade nearly straight . 9. verberans

Antenna usually lighter in color than head and thorax; @ paramere
with marked constriction at base of blade; outer margin of blade
rounded 75)... - Sine 1-7 LOS emissuis

Male aedeagus prone antl, near eee and hen abruptly tapering to
apex; spines of 9 bursa confined to basal third ........ 19

Male aedeagus slender and evenly tapering from base to apex; spines of
© bursa in center... . ba ath t Cae 20

Pronotal punctures large al nee ny, Tea al own diameter;
dark reddish brown to black species; spines of 9 bursa without eee
base (fig. 8b); o@ genitalia asinfigs.4a-d...... . . 1. similis

Pronotal punctures smaller and central punctures separated by own
diameter; reddish-brown species; spines of 9 bursa with large, plaque-

like bases (fig. 9c); o genitalia as in oe 4j--..... 8. miscellus
Male genitalia asin fig.47...... a si che sy vey O- COMMUNIS
Male genitalia asin fig. 4k ........... . . 6. indistinctus
Male genitalia gasping figs) 4). estes aay Cyloteh oct. Gietrichi

AMERICANUS GROUP
(Mandible with pit on lateral margin)

Antennal segment 3 small, subequal to 2 (in doubtful cases of some
29, 3 less than 1% length of 2)..... Sb Sacks, ae ee
Antennal segment 3 intermediate in size peneeen 2 ead a clearly larger
than 2 (in doubtful cases of some 99, more than 1% length

Of 2)). ae ; MEE er nen ce ee Te
Small eeeuee: Sion ees fone 6 AD a Th Weer eee emesis ee Man Ne e233
Larger species, elytron more than6mm .. . a irrew ian soto

Pronotum wider than long; pronotal punctures preter than those on
front and sparse, widely separated, coe size throughout; East
Coast to Nebraska... . . ... . . 382, americanus

Pronotal punctures coarse, larger hen on ieront on anterior two-thirds
and fine, sparse on basal one-third in o&; Southwest . 36. concisus

Antenna exceeds tip of hind angle by 1-4 segments; Bar weioiat, fovea
large; flagellar segments elongate ... . Mais Nae tile 43)

Antenna of o@ barely exceeds tip of hind srry 9° anvenna shorter;
parantennal fovea, clearly smaller than antennal segment 2; flagellar
segments triangular, not elongate; pronotal punctures large and dense;
o paramere with peculiar, winglike flaps near center; Texas.

38. hamatus

Elytron length 6-8 mm. . . . ee Care oe)

Large species, elytron 8.5-11.5 mm; mate eee poone as wide as
high; antenna exceeds tip of hind pee by about 2 segments; o para-
mere with definite apical blade and antennal hairs long and dense.

19. corticinus

Antenna very long, exceeds tip of hind angle by 2-4 segments; sides of
pronotum divergent posteriorly; nasal narrow, higher than wide.

33. cribriventris

12

27(21).

28(27).

29(28).

30(29).

31(29).

32(31).

33(32).

34(31).

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 12%

Antenna shorter, exceeds tip of hind angle by no more than 1 segment;
sides of pronotum svubparallel; nasale much wider than high.
34. obscuratus

Elytron more than 5 mm .. . BDI... 3A a8
Very small species, elytron less than as a mm; Srasale flat: frontal margin
rounded, moderately thin and jutting; Sorel paneeaes nearly as
large as those on front; parantennal and mandibular foveae relatively

large; southeast U.S. to Texas... . ... . . 42. insipiens
Pronotal punctures and vestiture aioe prone usual small increase
in density and size along sides. . . . . Nee 28

Pronotal punctures change from rather ace nonionly to smaller and
denser along posterior border, vestiture also becomes fine over same
area; antenna of co exceeds tip of hind angle by no more than one-half
segment, 9 antenna shorter; co aedeagus broad; paramere with small
apical blade; elytron about 8.5 mmlong...... . 41. gradatus

Parantennal fovea very large, as large or larger than antennal segment 3;
nasale large and protuberant, especially in o&; frontal margin strongly
projecting, angular with blunt apex when viewed from above; elytron
8 mm or more; sides of pronotum straight behind anterior curvature

and divergent posteriorly, pronotum wider than long. . .. . 30
Parantennal fovea clearly smaller than antennal segment 3; not with
BbOVECOMpPIBATION.: 7.; cus -sksues Since eee dae eins et ce en eer 31

Frontal margin scarcely overhanging protruding nasale, not extended
in angular apex; pronotal punctures as large as those on front; @
paramere with apical blade; elytron 8.9-11.2 mm. . 20. sagittarius

Frontal margin overhanging nasale and usually curved down on sides
above nasale, margin extends out sharply and ends in truncated,
angular apex; pronotal punctures smaller than those on front; @
paramere without apical blade; elytron 8.2-9.3 mm. . . 21. hyslopi

Sides of pronotum straight behind small anterior curvature; hind angles
large and expanded, beginning at about center of pronotum and thence
noticeably divergent, carina usually long and distinct. . . . . 32

Sides of pronotum concave or parallel from center to base of hind angles,
which arise well behind center; not with above combination of char-
acters ..<. . Ss JAS eae

Parantennal foe or BECnITS ieee ana an deccioped panera about as
high as wide; antenna not greatly elongate; smaller species, elytron
ienennoniece oor. cute iat yietiel PimOo

Parantennal fovea and nace) meee Pree wider nan high; antenna
very long, exceeds tip of hind angle by 3-4 segments, flagellar segments
elongate and slender, 4 about twice as long as wide; pronotal punctures
about as large as those on front and separated by own diameter; apex
of last visible sternite slightly inflated; elytron 8.5-10 mm.

24. difficilis

Pronotal punctures smaller than those on front; frontal margin often
rounded in center; o’ paramere oes at base, abruptly rounded
RUAUU yee a oo 2 22> testaceus

Pronotal punches Tents same size as noes on front (but widely
separated); frontal margin often coming to blunt point in center;
co’ paramere slender, gently tapering to tip. . . . 27. trapezoideus

Frontal margin thin, shelflike, and extending in front of nasale by
several times width of black rim, front with depression behind margin;
parantennal fovea small . . 5 .4( fos sce ee eee

NO. 3568

35(34).

36(34).

37(36).

38 (37).

39(38).

40(39).

41(39).

42(41).

43 (42).

44(41).

45(44).

46(44).

MELANOTUS—QUATE AND THOMPSON 12

Frontal margin not strongly protruding .. . . 36
Pronotal punctures separated by no more faa own neier Ayeaiien
segments elongate, o antenna exceeds ce of hind angle eo 3-4 seg-
ments; elytron about 7mm ... Pa jdicernes) co. macer
Pronotal punctures sparse, eepaiied aa more than own diameter;
flagellar segments triangular, antenna exceeds hind angle by no more
than 2 segments; mandibular pit very small; elytron 6-7 mm.
31. parallelus
Antenna extending to or beyond hind margin of pronotum ... . 37
Antenna short, ending before hind margin of pronotum in both sexes;
nasale flat, shallow; co paramere slender, aedeagus broad until near
QDEX). Salma. ... . . 23. piceatus
Nasale clearly Heanedt nor fopeoleeeent BOLO Seridiah brownor darker. 38
Nasale obsolescent; color brownish yellow, o' paramere without apical
blade; pronotal punctures large and compact; vestiture rather thick;
small species, elytron 5.0-6.5 mm; Texas. . ... . . 35. beameri
Antenna, legs, and body of same calor Baa ye Pe oO
Antenna and legs reddish brown and body black ester specimens
with body dark brown but darker than appendages); pronotum
convex and rounded dorsally, rounded and concave at base of hind
angles; central pronotal punctures small and sparse; co paramere

without apical blade. . .. . oo» « 29. pertinax
Western U.S., not occurring east of Nee Morice Sat ec eee oO,
Eastern U.S. and as far west as western Texas, Kansas, etc. . . . 41

Reddish-brown species; southwestern U.S. from south of Tehachapi
Mts., California to New Mexico, elytron 6.5-9.8 mm.
39. longulus longulus
Dark reddish-brown to black species; California north of Tehachapi
Mts., Great Basin, Pacific Northwest; elytron 7.5-10.9 mm.
40. longulus oregonensis

Larger species, elytron more than 7.5 mm........... 42
Smaller species, elytron less than 7.5mm... . is. Peer A.A:
Parantennal fovea well developed; one pitievaies well sepa-

ratedisn si) mast hs ra SS Geeta

Parantennal fovea eee ene flat; nronGial nee very close
together; o’ paramere "Nags eal lateral margin straight; color
usually reddish brown. . . ; . .. . . 18. cribulosus

Sides of pronotum parallel or concave over aaa half; pronotal punc-
tures usually smaller than those on front; color often reddish brown.

17. morosus

Sides of pronotum divergent posteriorly; pronotal punctures usually as

large as those on front; color aes dark reddish brown, darker than

morosus .. . ae . .. . . 16. depressus
Nasale well aeeeed hots as ae as ee sleet Kabel Ean a SO
Nasale flat, not protuberant, wider than high . . ....... . 46
Pronotal punctures about as large as those on front; pronotum flattened
and not rounded... . . LS AB 27. trapezoideus (part)
Pronotal punctures small, eaelice than those on front; pronotum
TOUNGEA) erste ie j . . . . . 82, americanus (part)
Side of pronotum rent nithoun curvature before hind angle . . 47

Side of pronotum with slight curvature before hind angle; frontal margin
broad and nearly straight when viewed from above. . 22. prasinus

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

47(46). Male paramere with definite apical blade; thorax about as long as wide;

distributed in central and south central US ......... 48

Male paramere without apical blade; thorax wider than long; largely
distributed along east coast. . .. . (Mies 28, tenax

48(47). Male paramere not unusually slender, pase of oreo blade angled at
abouti9Osm Faesen 2). pathy 80. infaustas

Male paramere very slendett pate of eapioal ielarle gently rounded and

not strongly angulate.. (jaw... Ss a Ss ho ameeator

The similis Group
(Mandible with pit on lateral margin)
1. Melanotus similis (Kirby)
Fiaures 4a—d, 8b, c; PLATE la

Perimecus similis Kirby, 1837, p. 149.

Melanotus similis —Erichson, 1842, p. 116.—Candéze, 1860, p. 363.

Elater fissilis Say, 1839, p. 183. [New synonymy.]

Cratonychus fissilis—LeConte, 1853, p. 477.

Melanotus fissilis—Candeéze, 1860, p. 352.—Blatchley, 1910, p. 750.—Van
Zwaluwenburg, 1922, p. 12—Thomas, 1941, p. 259.— Dietrich, 1945, p. 57.—
Severin, 1949, p. 16—Brooks, 1960, p. 40.

Cratonychus laticollis—Erichson, 1842, p. 102.

Ctenonychus ochraceipennis Melsheimer, 1846, p. 150.

Melanotus ochraceipennis—Thomas, 1941, p. 259.

Ctenonychus sphenoidalis Melsheimer, 1846, p. 150.

Melanotus sphenoidalis.—Thomas, 1941, p. 260.

Cratonychus exuberans LeConte, 1853, p. 477. [New synonymy.]

Melanotus exuberans.—Candéze, 1860, p. 354.—Thomas, 1941, p. 260.

Mauz.—Body color dark reddish brown, covered with yellowish
hairs evenly distributed over body.

Head: Front coarsely punctate, most punctures hexagonal shaped,
distance between punctures less than half puncture diameter, center
slightly protuberant; clypeus lightly punctate, parantennal fovea
shallow, width of nasale 1% to 2 times height. Mandible without pit;
palpi yellowish brown. Antenna exceeds tip of hind angle by 2 seg-
ments, average ratio of segments 2:3:4=4:8:12; erect male hairs
moderately dense, often sparse enough to be inconspicuous; third
segment 1% to 2 times longer than wide; outer angles of flagellar
segments often lighter in color than rest of segment, usually testaceous.

Pronotum often moderately flattened and expanded; wider than
long, ratio of width length=1.18 (1.06-1.25); punctures moderately
coarse at center, about size of those of front, separated by distance
equal to less than puncture diameter, punctures little more dense on
sides and front, oval shaped except those on anterior angles hexagonal
shaped as on front; hind angles divergent, often with 2 carinae; side
concave before hind angle.

NO. 3668 MELANOTUS—QUATE AND THOMPSON 15

Elytron with punctures of striae moderately deep, separated by
distance equal to less than own diameter, interstrial area at center of
elytron about 3 times as wide as puncture diameter. Venter of meta-
thorax and abdomen with evenly distributed, elongate punctures.
Metacoxal plate evenly tapering to apex. Genitalia as figured;
paramere with well-defined angle at tip, with about 8 hairs along
outer angle of apical blade.

Elytron: 9.4+0.5 mm (7.9-11.4). 180 spec.

FEMALE.—Similar to male, larger, antenna shorter, extending to tip
of hind angle. Genitalia as figured, bursa with few to large number of
spines, spines short, about as long as diameter of base, accessory gland
very long and slender, as long as spermathecal duct and one-half as
wide, diverticulum branching near center of sperm, duct without
visible blind pouch apically in usual position near spermatheca.

Elytron: 10.2-+0.9 mm (8.4—12.8). 120 spec.

Typres.—Lectotype of similis: Male, “Amer. Bor./ex Mus. Murray/
Frey Coll. 1905-100” (BMNH). Selected by Quate, 1960.

Types of fissilis: Specimens lost.

Lectotype of Jlaticollis: Male, ‘“‘Amer. Sept., Norwish”’ (ZM).
Lectotype selected by Quate, 1960, from cotypes of 1 male and 5
females.

Lectotype of ochraceipennis: Female, ‘‘Pa./Ziegler’” (MCZ). Se-
lected by Quate, 1962. First specimen of this species in Melsheimer
Collection too badly broken for positive identification and therefore
not selected as lectotype.

Type of sphenoidalis: Male. ‘Pa.”” (MCZ). Only specimen in
Melsheimer Collection.

Lectotype of eruberans: Male, probably Santa Fe, N. Mex. (MCZ).
First specimen in type series selected by Quate, 1962. Second speci-
men of type series is a female belonging to the communis complex and
is not exuberans.

DistRrBuTION.—Southern Canada and U.S. east of Montana, Utah,
and Arizona; southwest to southern California andMexico.

AuaBAMA: Mobile; February, June. Arizona: Apache, Cochise, Coconino,
Gila, Graham, Madera, Maricopa, Pinal, Pima, Santa Cruz, Yavapai, Yuma;
April, October. ArKANsAS: Hempstead, Lawrence, Polk, Washington; March,
June. Cauirornia: Riverside (Blythe); May, July. Cotorapo: El Paso,
Montezuma (Berkeley); July, August. Connecticut: Fairfield, Hartford,
Litchfield (January, reared?), Middlesex, New Haven, Tolland; May—November.
Fiorwa: Brevard, Hendry, Manatee, Orange, Osceola, Palm Beach, Seminole,
Volusia; February, August. Grorara: Chatham, Floyd, Fulton, Peach, Thomas,
Warren, Okefenoke Swamp; March, September. Ittrnors: Boone, Champaign,
Cook, DeKalb, Edgar, Iroquois, Jackson, Jo Daviess, Johnson, Kanakakee, Lake
McHenry, McLean, Marion, Marshall, Mason, Peoria, Putnam, Union, Vermilion;

March, September. Inprana: Clark, DeKalb, Greene, Knox, Lake, Marion,
Orange, Porter, Pulaski, Stark, Steuben, Tippecanoe, White; April, October.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Iowa: Buchanan, Clayton, Fayette, Floyd (reared), Franklin, Harrison, Henry,
Jasper, Johnson, Muscatine, Pottawattamie, Poweshiek, Story, Woodbury; May,
October. Kansas: Atchinson, Bourbon, Clark, Decatur, Dickinson, Doniphan,
Douglas, Gove, Gray, Johnson, McPherson, Miami, Norton, Osborne, Pottawat-
omie, Rawlins, Reno, Riley, Scott, Sedgwick, Shawnee, Sheridan, Sumner; April,
August. Kentucky: Jefferson, Ohio: June, August. Lovurstana: Lafourche,
Orleans, Plaquemines, St. Landry, St. Tammany; January, September. MaIne:
Penobscot, Washington; June, July. Maryzianp: Anne Arundel, Calvert, Har-
ford, Montgomery, Plummers Isl., Prince Georges, Queen Annes, Washington;
April, December. Massacuuserts: Berkshire, Essex, Franklin, Hampden,
Hampshire, Middlesex, Norfolk, Plymouth, Suffolk, Worcester; April, September.
Micuigan: Calhoun, Cheboygan, Huron, Monroe, Oakland, Ottawa, St. Clair,
Sanilac, Tuscola, Wayne; April, August. Minnesota: Big Stone, Dakota, Henne-
pin, Pope, Ramsey, St. Louis, Winona, Wright; May, August. Mississippi:
George, Greene, Lauderdale, Oktibbeha; January, September. Mussourt:
Boone, Buchanan, Callaway, Cape Girardeau, Lawrence, Pike, St. Louis, Wayne;
April, October. Montana: Custer; August. NrBraska: Dakota, Dawes,
Dodge, Holt, Lancaster, Lincoln, Otoe, Red Willow, Scotts Bluff, Sioux, Washing-
ton; May, August. New Hampsnrre: Coos, Strafford; May, June. New
Jersry: Atlantic, Bergen, Burlington, Gloucester, Middlesex, Morris, Ocean;
April, November. New Mexico: Bernalillo, Dona Ana, Grant, Lincoln, Sando-
val, San Miguel, Santa Fe, Socorro; June, August. New York: Albany, Cattarau-
gus, Cayuga, Clinton, Columbia, Cortland, Delaware, Dutchess, Erie, Essex,
Franklin, Greene, Herkimer, Jefferson, Madison, Monroe, Nassau, New York,
Niagara, Oneida, Ontario, Orange, Orleans, Oswego, Queens, St. Lawrence, Suf-
folk, Sullivan, Tompkins, Ulster, Warren, Wayne, Westchester, Wyoming, Yates;
March, October. Norra Caroiina: Buncombe, Chowan, Iredell, Pasquotank,
Polk, Richmond, Swain, Wake, Watauga, Wilson; April, July. Norra Daxora:
Cass, Grand Forks; June. Onto: Ashtabula, Athens, Delaware, Fairfield, Frank-
lin, Fulton, Greene, Hamilton, Hocking, Holmes, Lucas, Pike, Preble, Summit,
Warren; May, October. OxLAHoMA: Atoka, Cleveland, Latimer, Oklahoma,
Tulsa; March, July. PrEnNsytvanta: Adams, Allegheny, Bradford, Dauphin,
Delaware, Erie, Franklin, Lehigh, Lycoming, Montgomery, Northampton,
Philadelphia, Sullivan, Susquehanna, Westmoreland, York; March, October
(Janaury and December, reared or in hibernation?). RuopEr Istanp: Washing-
ton; June, July. Soura Carorina: Oconee; April, August. SournH Daxorta:
Brookings, Codington, Sanborn, Union; June, July. TENNESSEE: Davidson,
Knox, Morgan, Sevier, Smith; January, August. Texas: Anderson, Bexar,
Brazos, Brewster, Cameron, Colemen, Dallas, Goliad, Jeff Davis, Jim Wells,
Lamar, Lubbock, Palo Pinto, Presidio, Smith, Shelby, Travis, Victoria, Wallers,
Wharton, Wilbarger, Williamson; February, September. Utan: Cache, San
Juan, Utah, Washington, Zion Canyon; June, July. Vermont: Orange, Wind-
ham; May. Vrrernra: Fairfax, Montgomery, Nelson; June. WASHINGTON,
D.C.: March, December. West Virarnra: Greenbrier, Lewis, Preston; July,
August. Wisconsin: Bayfield, Columbia, Dane, Dodge, Grant, Jefferson,
Lafayette, Rock, Sauk, Trempealeau, Waukesha; June, November. MANITOBA:
Aweme, Killarney, Winnipeg; June, August. Nova Scotra: Annapolis Royal;
June. Ontario: Belleville, Muskoka, Ottawa, Prince Edward, Ridgeway,
Sudbury, Toronto; May, July. Qursrc: Aylmer, Montreal, Mt. St. Hilaire,
Rigaud, St. Johns; June, August.

It is regrettable that the widely used name fissilis has to be replaced
by the unfamiliar similis; however, both Mr. Lane and the senior

NO. 3568 MELANOTUS—QUATE AND THOMPSON 17

author have examined types of similis, and there is no doubt that that
species is the same as the one known as fissilis.

Melanotus similis exhibits a good deal of variation throughout its
range. Variations in the ratio of the thoracic width to length, in the
size of the pronotal punctures, in the shape of the distal enlargement
of the male paramere, and the spines of the female bursa have been
observed. Some of the variation appears in a regular pattern with
an east-west cline and some is oriented in a north-south pattern in an
irregular manner. The two patterns are noncorrelated and inde-
pendent of each other (fig. 2).

The eastern populations of M. similis have a relatively wider
pronotum which generally narrows to the west and finally the narrow-
est pronotum is found in the Southwest. There is an approximate
east-west clinal variation, but this is irregular through the Midwest.
Table 1 shows the ratios of the pronotal length/width. There is no
significant difference between adjacent populations, but, as shown in
the chart, there are significant differences between nonadjacent popu-
lations. All measurements were made of males with dissected
genitalia. Females are not included because samples of uniform
sizes were not available in dissected specimens; however, less ex-
tensive analysis of females indicates that a similar pattern also exists
in that sex.

The male genitalia show variation roughly correlated with the
pronotal width/leneth ratio. No satisfactory means of expressing
the difference qualitatively was found, but the differences in various
parts of the range are illustrated in figures 5a-e. Generally, the
distal enlargement of the paramere is longer and the subapical in-
dentation is more abrupt in eastern populations, but there is less
constancy than in the pronotal variation and only a weak clinal
trend is noted. From a taxonomic viewpoint, the differences are not
ereat enough to cause difficulty with identification, and the male
genitalia are the most reliable criteria for recognition of the species.

The female genitalia display marked differences in the number of
spines on the bursa. Females can be sorted roughly into three
classes on the basis of the bursal spines. In the northern and western
part of its range, the three types are found with a little dominance of
the ‘normal’’ type, which has few spines; in the southeastern United
States there is almost exclusively the ‘spiny’ type, which has a
large number of spines (fig. 2). There are intermediates to the two
types which unite them in a gradient series, and no formal taxonomic
discrimination is warranted.

Ratio of width/length of pronotum of Melanotus similis (analysis
based on 20 males from each area, total of 140 specimens):

221-532—67—__2

18 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121

Iowa Lincoln, North Platte,
New York Illinois Missouri Kansas Nebraska Nebraska Arizona
120 1.19 i a7) 1. 16 ele Z; 1.14 1. 12

LSD 0.01=0.024

In the southwestern United States there is a form which has been
described under the name eruberans. Many specimens of this form
appear quite different from typical similis—the punctures of the
pronotum are more coarse and dense, the pronotum is narrower, the
distal enlargement of the paramere is smaller, and the indentation is
gradual and not abrupt. Viewed apart, this form does appear to be
a distinct species, but the complete picture shows it is connected with
other similis forms by an intergrading series. The shape of the
pronotum and genitalia appears to be merely an extreme extension
of the clinal trend described above, and southwestern specimens are
more similar to specimens from western Nebraska, Kansas, Texas,
and eastern Colorado than to eastern populations. The narrower
pronotum and smaller genitalia are characteristic of the western popu-
lations; furthermore, in the same locality one finds typical ‘“ezxu-
berans”’ types intermediated with clear intergrades to typical similis.
Specimens have been seen with the coarse pronotal punctation of
“eruberans,”’ typical similis genitalia, and others with the opposite
combination. Some are intermediate between the two forms in both
features. Females possess the “spiny” bursa and are quite like other
similis females except that generally they have coarser pronotal punc-
tation. It therefore appears that “eruberans”’ is a synonym of similis
and represents a geographical variant not defined well enough to
justify segregating it as a separate taxon. (A similar, though less
well documented, trend to larger pronotal punctures and a narrower
pronotum was also noted in communis from the Southwest.)

Most specimens identified by Mr. Lane or by us will be labelled
in collections as M. fissilis Say.

2. Melanotus spadix (Erichson)
FiguREs 4e, 8e

Cratonychus spadix Erichson, 1842, p. 103.
Melanotus spadix.—Candéze, 1860, p. 354.
Melanotus abdominalis of authors, not Erichson, 1842.
Melanotus decumanus.—Dietrich (not Erichson), 1945, p. 55.
Maue.—Large species, body color reddish brown to dark reddish
brown, covered with yellowish hairs evenly distributed over body.
Head: Front coarsely punctate, most punctures hexagonal shaped,
distance between punctures less than half puncture diameter, surface
sometimes with slight, broad depression anteriorly, frontal margin
evenly convex, not strongly protuberant; parantennal fovea very

NO. 3568 MELANOTUS—QUATE AND THOMPSON 19

shallow. Mandible without pit; palpi reddish to yellowish brown.
Antenna exceeds tip of hind angle by about 2 segments, erect male
hairs usually very dense and readily observed; average ratio of seg-
ments 2:3:4=4:8:13, segment 4 slender, about twice as long as
maximum width (10:6), outer parts of flagellar segment lighter in
color than rest of segment.

Pronotum wider than long, ratio of width/length=1.2; punctures
about same size as those of front, at center separated by distance
equal to puncture diameter or less, little more dense on sides and
front, oval in shape except those on anterior angles hexagonal shaped
as on front; hind angles divergent, with single, strong, dark carina;
sides divergent and nearly straight to about two-thirds distance from
front, rather sharply angled before concavity in front of hind angles.

Elytron and venter of thorax and abdomen as in similis. Genitalia
as figured; paramere with well-defined angle at tip, large cluster of
30-40 long hairs along outer part of apical blade.

Elytron: 10.8+0.97 mm (8.5-13.0). 30 spec.

FEMALE.—Similar to male; larger in size, antenna shorter. Geni-
talia as figured; bursa nearly trapezoidal with numerous spines com-
pacted to extent that bursa appears sclerotized; spermathecal duct
without diverticulum.

Elytron: 11.3+0.84 mm (10.8-12.6). 12 spec.

Typr.—Holotype of spadix: Male, “Amer. Sept.’”? (ZM 17035).
Only specimen in type series.

Distripution.—Eastern U.S. west to Nebraska and Arkansas.

ARKANSAS: Washington; July. FuLoripa: Manatee; March. Geroreia:
Clarke, Fulton, Habersbam; April, July. Inurvors: Piatt, Whiteside; June.
Inp1ana: Clark, Lake; June, July, December. Marynanp: Anne Arundel,
Baltimore, Montgomery, Plummer’s Is.; May, June. Mricuiagan: Marquette,
Oceana; June, July. Montana: Barry; June. Nepraska: Washington; June.
New Jersey: Atlantic, Burlingtion, Middlesex, Ocean; May—August. Nrw
York: Erie, Queens, Suffolk; May-July. Norra Caronina: Moore, Polk;
May. PENNSYLVANIA: Dauphin; August. Sourn Caronina: Oconee; June.
TENNESSEE: Sevier; June. Virainta: Gloucester, Nelson; May, June.

M. spadix is closely allied to M. similis but is usually larger and
lighter in color, and antennal segment 4 is more slender. For posi-
tive identifications, however, it is usually necessary to examine the
dissected genitalia. The male genitalia of the two species are quite
dissimilar, and spadiz males are easily recognized by the large tuft
of hairs covering the whole surface of the lateral part of the gono-
style apical lobe. The female genitalia are also conspicuously dif-
ferent, and the densely spined bursa of spadiz bears little resemblance
to that of similis.

Many specimens identified by Mr. Lane and by us will be labelled
as M. abdominalis since this name is believed to be the correct one

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

for spadiz, but study of the types of abdominalis reveals it is an un-
familiar species which apparently does not occur in North America.

3. Melanotus decumanus (Erichson)
Ficures 4f-h, 8d; PLATE 1b

Cratonychus decumanus Erichson, 1842, p. 104.—LeConte, 1853, p. 474.

Melanotus decumanus.—Candéze, 1860, p. 341.—Blatchley, 1910, p. 749.—
Thomas, 1941, p. 258.—Severin, 1949, p. 15.—Fattig, 1951, p. 20.

Cratonychus cuneatus LeConte, 1853, p. 473.

Melanotus cuneatus.—Thomas, 1941, p. 258.

Cratonychus incertus LeConte, 1853, p. 474.

Melanotus canadensis Candéze, 1860, p. 342.—Blatchley, 1910, p. 751.—Thomas,
1941, p. 258.—Dietrich, 1945, p. 55.

Mate.—Large species, body color dark reddish brown, covered
with yellowish hairs evenly distributed over body.

Head: Front coarsely punctate, most punctures hexagonal shaped,
distance between punctures less than half puncture diameter, surface
with broad depression anteriorly, margin strongly jutting and forming
noticeable shelf above clypeus; parantennal fovea very shallow,
usually teardrop shaped. Mandible without pit; palpi reddish to
yellowish brown. Antenna about exceeds tip of hind angle by 1%
segments, erect male hairs dense and easily observed, average ratio
of segments 2:3:4 varies from 5:10:20 to 5:38:19, segments 4 and
following longer and more slender than similis.

Pronotum wider than long, ratio of width/length =1.2, a little
flattened dorsally, punctures about same size as those on front, at
center separated by distance equal to puncture diameter or less, little
more dense on sides and front, oval in shape except those on anterior
angles hexagonal shaped as on front; hind angles divergent, with
single, strong, black carina, and often a second smaller one; sides
divergent and nearly straight to about two-thirds distance from base,
rather sharply angled before concavity in front of hind angles.

Elytron and venter of metathorax and abdomen as in similis.
Genitalia as figured, paramere straight or with gently sloping sides
basad of apical blade, blade with 8 hairs.

Elytron: 11.2+0.6 mm (10.0-12.4). 33 spec.

FremaLe.—Similar to male. Larger, antenna little shorter. Gen-
italia as figured; bursa completely devoid of spines or sometimes with
patch of 5-8 small spines (like those of similis) on either side of
bursa near base.

Elytron: 12.2+0.8 mm (10.0-14.1). 47 spec.

Typrs.—Holotype of decumanus: Female, ‘Amer. Sept.’’? (ZM).
Only specimen in type series.

No. 3568 MELANOTUS—QUATE AND THOMPSON oh

Lectotype of cuneatus: Male, labelled with orange circle (Southern
States). (MCZ). First specimen of type series selected as lectotype
by authors in 1962.

Lectotype of incertus: Male, labelled with green circle ‘Middle
States.” (MCZ). First specimen to type series selected as lectotype
by authors in 1962.

Holotype of canadensis: Male, “Canada” (BMNH).

DiIsTRIBUTION.—EHastern U.S. and Canada west to South Dakota,
Colorado, and New Mexico.

AuaBaMA: Mobile; May. Connecticut: Fairfield, Hartford, New Haven;
March, August. Grorata: Bibb, Clarke, Crawford, DeKalb, Fayette, Fulton,
Richmond, Walton; April, June. INnprana: Greene, Vigo; June, October. ILui-
NoIs: Champaign, Marshall; May, June. Iowa: Dickinson, Woodbury; July.
Kansas: Douglas, Greenwood, Leavenworth, McPherson, Reno; May, June.
Kentucky: no data. Marynanp: Montgomery; June. MassacHusETTSs:
Hampshire, Middlesex; June. Micutcan: Cheboygan; July. NEBRASKA:
Cuming, Dakota, Lancaster, Lincoln, Otoe, Thomas; May, June. New Hamp-
SHIRE: Strafford; June. New Jerssy: Atlantic, Bergen, Essex, Monmouth,
Montclair, Morris, Ocean, Passaic; June, July. New Mexico: Bernalillo,
Sandoval; May, June. New York: Cattaraugus, Greene, Kings, Nassau,
Tompkins, Westchester; June, July. Norra Carouina: Beaufort, Brunswick,
Buncombe, Columbus, Duplin, Hyde, Jackson, Johnston, Montgomery, Wake;
May, June. Onto: Clark, Erie, Hamilton; May, July. PENNsyLvantia: Dela-
ware, Northampton, Philadelphia; June, September. Ruopr IsutaANp: Wash-
ington; June, July. SourH Carouina: Aiken, Dorchester, Edgefield, Florence,
Oconee; May, June. SoutnH Dakota: Brookings, Union; June. TENNESSEE:
Smith, Vrreinra: Alexandria, Dinwiddie, Essex, Faquier, Nelson, Nottaway,
Prince Edward; May, June. Nova Scotra: Annapolis; June.

The large, jutting frons of M. decumanus is usually sufficient to dis-
tinguish this species from M. similis, spadix, and castanipes; it is vari-
able, however, and there are specimens that have a reduced frontal
margin which might cause them to be confused with the other species.
The females often have a smaller margin than males. The male geni-
talia differ from similis in the gradual expansion of the paramere basad
of the apical blade but can be confused with some similis in which the
enlargement is not abrupt. The female bursa is strikingly different
from that of the allied species with the complete lack of spines.

4. Melanotus castanipes (Paykull)
FIGuRES 41, 8f; PLATE lc

Elater castanipes Paykull, 1800, p. 23.

Cratonychus castanipes.—Erichson, 1842, p. 95.

Melanotus castanipes—Redtenbacher, 1849, p. 249.—Schwartz, 1892, p. 152
[as a synonym of rufipes]—Putzeys, 1908, p. 294 (larva).—Blatchley, 1910,
p. 746.—Binaghi, 1939, p. 224.—Thomas, 1941, p. 257.—Dietrich, 1945, p.
54.—Severin, 1949, p. 15.—Fattig, 1951, p. 19.—Jeuniaux, 1955, p. 234.

D2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Melanotus paradoxus Melsheimer, 1846, p. 152.

Cratonychus paradoxus.—LeConte, 1853, p. 480.

Cratonychus inaequalis LeConte, 1853, p. 476.

Melanotus inaequalis —Leng, 1920, p. 174.—Thomas, 1941, p. 258.

Cratonychus scrobicollis LeConte, 1853, p. 476.

Melanotus scrobicollis—Leng, 1920, p. 174.—Thomas, 1941, p. 258.—Fattig,
1951, p. 19.

Melanotus texanus Candéze, 1860, p. 351.

Mauer.—Large species, body color reddish to dark reddish brown,
covered with yellowish hairs evenly distributed over body.

Head: Front coarsely punctate, most punctures hexagonal shaped,
distance between punctures less than half puncture diameter, surface
often with shallow depression on disc, margin not protuberant, ex-
tending little beyond level of clypeus, rounded, weakened in center
and when viewed from directly in front appears obliterated; paran-
tennal fovea very shallow. Mandible without pit; palpi yellowish
brown. Antenna exceeds tip of hind angle by about 3 segments, seg-
ments 2 and 3 small, much smaller than 4, ratio of 2:3:4=5:7:17.

Pronotum wider than long, punctures about same size as those of
front, little deeper than in similis, at center separated by distance
equal to puncture diameter or less, little more dense on sides and front,
oval in shape except those on anterior angles hexagonal shaped as on
front; hind angles divergent, with single, strong, black carina; sides
markedly divergent beyond center, appear thinner than usual.

Genitalia as figured; paramere with large apical blade, side nearly
straight basad of blade.

Elytron: 11.2+0.7 mm (9.9-12.6). 50 spec.

FrmMaLe.—Similar to male, but larger. Antennal segments 2 and 3
not as small, ratio of 2:3:4=7:9:17. Pronotum is fuller and does
not have thin appearance toward margin. Genitalia as figured; bursa
with numerous, long spines on basal % arranged so densely to give
bursa sclerotized, striated appearance, apical % membranous except
small patch of spines; accessory gland distinctly clavate.

Elytron: 12.7+0.7 mm (10.8-14.1). 50 spec.

Typres.—Types of castanipes: 3 males, 2 females; 1 labelled with
red and green tags and ‘213/61,’ others only with green tag (Riks-
museum, Stockholm). These are presumed to be Paykull’s types, but
no lectotype has been selected because we feel that the validity of
these types may be questioned.

Lectotype of paradozus: Male, no collection data (MCZ). Only
specimen in type series.

Lectotype of inaequalis: Female, labelled with pale blue circle
‘Lake Superior’ (MCZ). First specimen in type series selected as
lectotype by authors, 1962.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 23

Lectotype of scrobicollis: Male, labelled with pink circle ‘‘Middle
States” (MCZ). Third specimen in type series selected as lectotype
by authors in 1962. First specimen with broken antennae and second
specimen, apparently male, doubtfully referred to this species.

Type of texanus: Female, ‘Texas, Jansen Coll. ex Candéze 1903,
130,” (BMNH).

DistriputTion.—Holarctic. Southeastern Canada, eastern U.S.
west to Utah, Arizona, southern California, Mexico.

Arizona: Coconino, Pima; June, July. Catirornia: ‘S. Cal.’”? CoLtorapo:
Larimer, “Waldo Canon’; June, July. Inimnors: McHenry; May. Maine:
Penobscot; June, July. Massacuuserts: Suffolk; May. Muicuigan: Alpena,
Crawford, Marquette, Oakland; June, July. Minnesora: St. Louis. NEw
HampsuireE: Carroll, Cheshire, Coos; April, August. New Mexico: San Miguel.
New York: Cattaraugus, Cortland, Erie, Essex, Niagara, Oswego, Tompkins,
Warren; March, September. Norra Carouina: Wake; June. Onio: Summit;
May. PrNNsyLvania: Carbon; June. RuopE Is~tanp: Washington; August.
Urau: Juab, Utah; January, July. Wisconsin: Dodge, Milwaukee; May, July.
Nova Scotia: Bridgetown; July. Ontario: Prince Edward; May. QurEBEc:
Mt. St. Hilaire, Regaud, Deparquest, Hull, Aylmer, Hudson; May, July. Mrx-
1co: El Salto Diego, June.

With its large size and small second and third antennal segments,
the male of castanipes is not difficult to recognize; it is easily sepa-
rable from ignobilis by the shape of the flagellar segments. The
females are more difficult to identify, since the third antennal seg-
ment is larger and more nearly intermediate between the second and
fourth, which makes it similar to spadiz or similis. The weakened
frontal margin, when viewed from directly in front, is the most useful
external character for separating castanipes, but in doubtful cases
only the dissected genitalia will provide a reliable answer.

M. castanipes is a widely distributed and variable species. Large
series from throughout its range might show it divides into subspecies,
but, from our rather limited material, we can only point out the pat-
tern of variation we observe without applying names to any of the
variants. ‘Throughout the species there is sexual dimorphism with
the females larger, the edges of the pronotum less flattened laterally,
the frontal margin less jutting, and the third antennal segment longer.
Specimens from the eastern United States are observably different
from those in the Southwest and the Great Plains. The western
specimens are smaller (male average elytron length is 10.7 mm com-
pared to overall average elytron length of 11.2 mm), and the pronotum
is more strongly flattened laterally and often more strongly divergent
posteriorly. The most conspicuous variation of the western speci-
mens is the thin, jutting frontal margin which often contrasts sharply
with the flattened or rounded, obsolescent margin in eastern specimens.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

These differences are more strongly marked in males than in females,
but the trend is parallel in the two sexes. The distinctive genitalic
characters in both the male and female, however, are constant through-
out the range of the species.

Two females from Europe have been dissected and examined. No
observable differences are noted between these specimens and Ameri-
can ones and, externally, they are similar to the eastern forms de-
scribed above. The female internal genitalia is precisely as the
American specimens. The bursa bears a dense patch of spines cen-
trally and a smaller, separated patch distally. The accessory gland
is enlarged, clublike apically and the spermathecal duct branches as
shown in the illustration.

Specimens examined: 1 9, Skane, Sweden, ‘Stromberg Det.”;
I.N.H.S. 1 9, Germany (?), Andreas Bolter Collection; I.N.H.S.

Melanotus rufipes (Herbst) is a European species closely related to
castanipes, and the synonymizing of the two species as proposed long
ago by Schwartz (1892) would seem warranted; however, we accept
the decision of recent students of the Kuropean fauna who regard the
two species as distinct (Binaghi, 1939; Jeuniaux, 1955).

5. Melanotus communis (Gyllenhal)
Fiegures 47, 9a,b; PuatTs 1d

Elater communis Gyllenhal, 1817, p. 188.—Say, 1839, p. 184.

Perimecus communis.—Kirby, 1837, p. 148.

Cratonychus communis.—Erichson, 1842, p. 102.—LeConte, 18538, p. 477.

Melanotus communis—Emmons, 1854, p. 88.—Candéze, 1860, p. 353.—Comstock
and Slingerland, 1891, p. 262.—Blatchley, 1910, p. 750.—Hyslop, 1916, p.
5.—van Zwaluwenburg, 1922, p. 12—Fenton, 1926, p. 502—Thomas, 1941,
p. 259.—Dietrich, 1945, p. 57.— Jewett, 1946, p. 10.—Fattig, 1951, p. 20.

Melanotus paganus Candéze, 1860, p. 359.

Mauze.—Body color reddish brown, covered with white or yellow
vestiture evenly distributed over body, specimens in Southwest
often darker.

Head: Front coarsely punctate, punctures rounded hexagonal,
separated by distance equal to less than own diameter, surface with
transverse depression anteriorly, sometimes very weak or reduced to
pair of shallow, anterolateral depressions, margin black or dark
reddish brown, not strongly protuberant above clypeus; clypeus
lightly punctate, parantennal fovea shallow, comma shaped, width
of interfoveal area 14-2 times height. Mandible without pit; palpi
yellowish brown. Antenna 1 or 2 segments longer than pronotum,
reddish brown, usually outer angles of flagellar segments testaceous
and usually with median, dark brown, longitudinal streak on each
side of flagellar segments; ratio of segments 2:3:4=4:7:9, segment 4

NO. 3568 MELANOTUS—QUATE AND THOMPSON Bs)

about 1% times as long as maximum width; erect male hairs short
and rather sparse.

Pronotum wider than long, ratio of width/length=1.22 (1.16-1.30,
S.D.=0.03, 100 spec.); punctures at center usually separated by
distance equal to more than own diameter, denser on margins, punc-
tures often larger in specimens from Southwest; hind angles divergent,
often with two carinae, lateral carina extending little anterior of base
of hind angle, median carina, when present, shorter and weaker than
lateral; side concave before hind angle.

Genitalia as figured; paramere with small indentation marking
apical blade, aedeagus evenly tapering to acute apex.

Elytron: 9.0+0.5 mm (8.1—10.1). 50 spec.

FEMALE: Similar to male. Genitalia as figured; accessory gland,
spermathecal duct, and diverticulum originate close together near
apex of bursa, bursal spines tacklike, large, conspicuous.

Elytron: 9.0+0.7 mm (7.1-10.6 mm). 40 spec.

Typrs.—Lectotype of communis: Male, “Amer. Bor. ??—401/62”
(Schéenherr Collection, Riksmuseum, Stockholm). First of two
specimens in type series selected as lectotype by Quate, 1962.

Lectotype of paganus: Female, “Amer. Bor., Coll. Janson, Ex
Dejean” (BMNH). First of two specimens in type series selected
as lectotype by Quate, 1960.

Two females from the Zoologiska Instit., Uppsala, were sent to us
when we requested the loan of communis types. These belong to the
species morosus and we believe they are not the type specimens of
Gyllenhal.

Distripution.—Kastern U.S. west to South Dakota, Oklahoma,
and Texas.

AuABAMA: Mobile; March. Arizona: Graham; August. ArKANsAS: Hemp-
stead, Washington; April, June. Connecticut: Hartford, Litchfield, Middiesex,
New Haven, Tolland; (Jan.), March-November. Conorapo: Pingree Pk.
(County?); August. FLortma: Dade, Palm Beach; May, July. Gerorata:
Fulton, Harris; June. ILtinois: Champaign, Cook, Gallatin, Lake, Piatt, Pike,
Pope, Union, Vermilion, Wabash; March-September. INp1aNA: Clark, Greene,
Pulaski, Spencer, Starke, Tippecanoe; April-September. Iowa: Decatur, Henry,
Scott; April-May. Kansas: Barton, Douglas, Gray, Reno, Republic; June-
July. Kentucky: Jefferson; March. Lovrstana: East Baton Rouge, Saint
Tammany; June. Maryann: Calvert, Harford, Prince Georges, Washington;
February-November. Massacuusrtts: Essex, Franklin, Hampden, Hamp-
shire; April-September. Micnigan: Kent, Monroe, Ottawa, Washtenaw;
April-June. Minnesora: St. Louis, no date. Mussrssrrp1: George; April-May.
Missouri: Boone, Callaway, Mississippi, Phelps, St. Louis, Stoddard; March-
November. Nesraska: Cherry, Lancaster, Lincoln, Sarpy; April-July. New
JERSEY: Bergen, Burleigh, Burlington, Camden, Essex, Middlesex, Morris,
Ocean, Union; February-September. New Yorx: Dutchess, Erie, Nassau,

26 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Niagara, Orange, Rockland. Schenectady, Tompkins, Wayne, Westchester;
March—December. NortruH Carouina: Wake (Mills River); June, September.
Onto: Athens, Franklin, Hocking, Richland; March, November. OKLAHOMA:
Woodward; June. PENNsyLvANIA: Allegheny, Bucks, Philadelphia, York;
March-September. SoutTH Carouina: Oconee; September, December. SoutTuH
Daxota: Bon Homme, Brookings, Union; June-July. TENNESSEE: Montgomery;
April. Texas: Harrison, Gonzales; April, June. Vireinta: Alexandria, Canal,
Fairfax, Loudoun, Nottaway, Spotsylvania; March-December. WaAsHINGTON,
D.C.: March-September. West Vireinia: Marion; no date.

6. Melanotus indistinctus Quate, new species

Fiaurss 4k, 9f,g

Maue.—Same as communis, but average size smaller, a little darker
in color, frontal margin little wider than black border, parantennal
fovea small but distinct, and pronotal punctures somewhat larger and
denser. Genitalia with slender, tapering aedeagus, paramere shorter
and thicker in communis, emargination before blade deeper and more
marked, blade broader and shorter.

Elytron: Holotype 8.2 mm; paratypes 8.3+0.5 mm (7.49.4). 50
spec.

Frmate.—Not clearly separable from communis (see discussion
below).

DistRIBUTION.—Virginia west to Kansas, Oklahoma, and Texas.

Holotype, male (USNM): Pittsfield, Pike Co., Ill., July 6, 1946, A. T. McClay.

Paratypes, 122 males (USNM, CU, ILL, U. Mo., U. Nebr., TEX AM, CAS,
Lane Coll’n): ILuinots: same as holotype, July 5-9, 1946; same, June 24, July 1,
7, 9, 1947, B. Cadwell; La Clede, Fayette Co., July 7, 1950, Ross and Sanderson;
Shawneetown, Gallatin Co., June 14, 1934, Ross and DeLong; Havana, Mason
Co., July 8, 1910, at light; Harrisburg, Saline Co., June 25, 1932, Ross, Dozier,
Park; Dubois, Washington Co., July 2, 1909, at light; Fox Ridge St. Pk., July 9,
1944. Virearnta: Holland, Nansemond Co., April 28, 1947, W. D. Fronk.
KeENntTuckKy: no other data. Mussisstppi1: Lincoln Co., May 18, 1938, peach
tree, W. F. Turner. Mussourtr: St. Louis, August 12, 1935, June 22, 1937,
USDA traps; Webster Groves, St. Louis, May 31, 1919, June 17, 1919, Satter-
thwait; Charleston, Mississippi Co., June 12, 1917, Satterthwait; Columbia, Boone
Co., May 4, 1940, W. R. Enns; same, June 15, 1907, R. H. Wolcott; same, July 14,
1941, H. E. Brown; Lathrop, Clinton Co., July 3, 1948, E. C. Becker. Lovuistana:
Shreveport, Caddo Co., June 2, 1949, J. H. Robinson; Olivier, Iberia Co., April
1904, E. S. G. Titus; Leesville, Vernon Co., May 1948, H.S. Fitch. ARKANSAS:
Clark Co., May 12, 1939, M. W. Sanderson; Hope, Hempstead Co., April 30, 1925,
April 23, 1926, May 17, 1926, July 5, 1926, June 25, 1931, L. Knobel; Hempstead
Co., May 8, 1939, Sanderson and Van Dyke; Miller Co., May 8, 1939, Sanderson;
Fayetteville, Washington Co., May 31, 1942, at light, Sanderson; Washington Co.,
May 30, 1938, Sanderson and C. Cameron. Kansas: Medora, Reno Co., July 3;
same, July 14, 1926, J. W. McColloch. Oxtanoma: Muse, LeFlore Co., May 7,
1939, Sanderson. Trxas: Luling, Gonzales Co., May 2, 1953, B. J. Adelson;
Tyler, Smith Co., May 20, 1942, L. D. Christenson, Bronson, Sabine Co., May 30,
1945, J. H. Robinson; Tyler, Smith Co., March 24, 1939, L. 8. Jones. ‘“‘So.
McAlester, Ind. T., June 11, Wickham.’? Cherokee Co., May 5, 1952, June 1,

NO. 3568 MELANOTUS—QUATE AND THOMPSON 27

1952, light trap; Smith Co., April 6, 14, 1956, April 24, 1937, May 2, 1956; Nacog-
doches, Nacogdoches Co., March 28, 1940.

Other specimens: TENNESSEE: Milan, Gibson Co., June 2. Texas: Orange,
Orange Co., June 20, 1949, W. C. Stehr.

Only from Pittsfield, Ill., do we have a good series of females
associated with males. In this series there appear to be two types:
one is indistinguishable from communis, and the other has large
bursa spines; the spermathecal duct branches farther from the bursa,
and the bursa surface is coarsely reticulate. It is possible that the
second type represents the female of indistinctus, but, since the evi-
dence is not unequivocal, we have not definitely identified any of the
females nor included them in the type series.

This species will be found in some collections identified by Lane as
‘species 20.”

7. Melanotus dietrichi Quate, new species

Fiaures 41, 9d,e
Melanotus communis var. A.—Dietrich, 1945, p. 57.—Severin, 1949, p. 16.

MALE.—Same as communis in all features except male genitalia.
Male genitalia with very narrow aedeagus and paramere, paramere
with elongate blade at tip and several hairs.

Elytron: Holotype 8.7 mm; paratypes 9.9+0.4 mm (8.8-10.6).
60 spec.

Femate.—Unknown (or inseparable from communis).

DistrisuTion.—Connecticut to Pennsylvania, west to Iowa.

Holotype, male (CU): Fairmont, Marion Co., W. Va., March 24, 1930, P. N.
Musgrave.

Paratypes, 67 males (USN M, CU, ILL, CIS, U. Mo., Ohio U., Lane Coll’n): New
Yorx: Poughkeepsie, Dutchess Co., June 19, 1933, P. J. Chapman; Staatsburg,
Dutchess Co., June 23, 1934, H. Dietrich; Rhinebeck, Dutchess Co., Nov. 5,
1934, Dietrich; Yonkers, Westchester Co., Nov. 21, 1940; Kensico,? Co., April
27, 1935, J. A. Angell. New Jersey: Teaneck, Bergen Co., October 1923;
Brookdale, Essex Co., April 24, 1916, E. R. Kolmbach. Connecticut: New
Canaan, Fairfield Co., Sept. 24, 1919, M. P. Zappe. Wasninaton, D.C.: Rock
Creek, Dec. 30, 1906; same, April 21, 1907, C. E. Burden. Marynanp: Cumber-
land, Allegany Co., Nov. 4, 1915, W. F. Pennington; Woliville,? Co., May 2,
1913. West VIRGINIA: same as holotype. Onto: Columbus, H. H. P. Severin;
Athens, Athens Co., Sept. 25, 1949, P. J. Spangler; same, June 26, 1935, A. Sintic;
same, April, June, Oct., Nov., Dec., W. C. Stehr; Ames Twp., Athens Co., April
24, 1934, Stehr; Delaware Co., April 1921, D. J. & J. N. Knull. Inprana:
Beverly Shores, Porter Co., Aug. 6, 1932. In.itNor1s: Summit, Cook Co., April
17, 1908; Vermilion Co., April 25, 1926, K. F. Auden. Iowa: Ames, Story Co.,
June 19, 1926; same, July 2, 1931, W. C. Stehr. Lovuistana: Baton Rouge,
June 20, 1916, T. H. Jones.

Other specimens: Connecticut: Middlesex Co., April. New Yorx: Colum-
bia Co., June, Queens Co. PENNSYLVANIA: Westmoreland Co., June, July.
Marytanp: Allegany Co., Nov. Onto: Athens Co., June, July. WEstT

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

VIRGINIA: Fayette Co., TENNESSEE: Knox Co., May. Iowa: Henry Co.,
April.

8. Melanotus miscellus Quate, new species
Fiaures 4n, 9c
Melanotus communis var. B.—Dietrich, 1945, p. 57.—Severin, 1949, p. 16.

MaAtEe.—Same as communis, except averages little smaller in size,
often darker reddish brown, and genitalia differ in being shorter,
aedeagus wider, paramere thicker, and with broader and shorter
apical blade, blade with dense cluster of hairs. Ratio of pronotal
width/length=1.2 (1.1-1.3).

Elytron, holotype 8.9 mm; paratypes 8.5+0.4 mm (7.6-9.7).
40 spec.

FEMALE.—Similar to male except sexual differences. Genitalia as
figured; differs from communis in that spines of bursa nearer apex,
spermathecal duct, spermathecal duct diverticulum and accessory
gland arise independently from bursa, accessory gland base far re-
moved from base of spermathecal duct.

Elytron: Allotype 8.7 mm; paratypes, 8.5+0.4 mm (7.8-9.5).
19 spec.

DistrisuTion.—Ontario, eastern U.S. west to Illinois.

Holotype, male and allotype, female (USNM): Chapel Hill, Orange Co., N.C.,
Feb. 3, 1935.

Paratypes, 52 males, 41 females (USNM, CU, ILL, NU, CAS): same data as
holotype and allotype. Marynianp: Sparrows Pt., Baltimore, July 3, 1932,
J. W. Green; Edgewood, Harford Co., Nov. 3, 1918, H. Dietrich. Virernia:
Pennington Gap, Lee Co., Hubbard and Schwarz. Fuioripa: Apalachicola,
Franklin Co., Taxodium distishum, W. F. Fiske. Grorerta: Savannah, Chatham
Co., March 9, 1940, Van Dyke. Mu£ssissippr: Lucedale, George Co., Feb. 27,
1931, March 22, 1932, H. Dietrich; New Augusta, Perry Co., Feb. 12, 1931, H.
Dietrich. Inp1ana: LaPorte Co., June 9, 1953, wheat, K. Pruess. ILuiINots:
Homer, Champaign Co., March 30, 1907, Hart and Hood; Mahomet, Champaign
Co., Sept. 13, 1933, H. H. Ross; St. Joseph, Champaign Co., May 4, 1913; Dec.
24, 1944, J. L. C. Rapp; Riverside Wds., Cook Co., Sept. 13, 1949, W. Tietz;
Western Springs, Cook Co., June 16, 1949, Ross and Stannard; Willow Springs,
Cook Co., May 12, 1921; Havana, Mason Co., July 11, 1948, Sanderson, Stannard,
Becker; White Heath, Piatt Co., March 20, 1942, Ross and Riegel; April 6, 1926,
K. F. Auden; Allendale, Wabash Co., April 30, 1950, Smith and Stannard;
Benson, Woodford Co., Oct. 11, 1933, Frison and Ross. Ontario: Prince Edward
Co., April 16, June 12, July 15, 1922.

This species appears intermediate between similis and communis.
External characters of color and pronotal punctures resemble com-
munis, but the male genitalia more closely resemble similis; the
position of spines of the female bursa is as in similis, but the shape of
the spines is like that of communis. The genitalia structures, however,
are distinct and although not easily recognized, seem constant. ‘The
larvae, according to Lane (in litt.), are also distinctive.

No. 3568 MELANOTUS—QUATE AND THOMPSON 29

This species will be found in some collections identified by Lane as
“species 2.”

9. Melanotus verberans (LeConte)

Ficures 4m, 97; PLATE le

Cratonychus verberans LeConte, 1853, p. 478.
Melanotus verberans.—Candéze, 1860, p. 356.—Blatchley, 1910, p. 753.—Thomas,
1941, p. 260.

MatEe.—Same as communis, except generally smaller in size and
usually reddish brown, but some specimens darker, antenna and
thorax usually concolorous, erect male hairs in antenna dense and
conspicuous, with smaller and sparser pronotal punctures, genitalia
with aedeagus nearly parallel sided and abruptly tapering to apex,
paramere rather broad and with little curvature basad of apical blade.
Ratio of pronotal width/length=1.2 (1.1-1.3).

Elytron: 7.7+0.45 mm (6.9-8.3). 9 spec.

FEMALE.—Similar to male except sexual differences. Genitalia as
figured; differs from communis in that apex of bursa more bulbous and
truncate, spermathecal duct diverticulum branches at or near base of
duct.

Elytron: 6.8-7.4 mm.

Typrs.—Lectotype of verberans: Male, labelled with pink circle
“Md.”’ (MCZ). Only specimen now in type series, selected as lecto-
type by authors in 1962; location of second type unknown.

DistrRiBuTIoN.—Maryland to Florida and west to South Dakota and
Texas.

ALABAMA: Locality?; January. ARKANsAs: Washington; May. Ftoripa:
Pinellas; March, May. Groraia: Ribb, Peach; April, June. I tuinots: Mason;
July. Inprana: Tippecanoe; April. Kansas: Nodata. Marynanp: Baltimore;
July. Mussissrprr: Forrest, George; January, April. NrpBrasKxa: Lancaster;
May. New Jersry: Middlesex; April. New York: Essex, Niagara; May,
November. Norra Carouina: Duplin, Moore, Orange, Wake, Wayne; April,
May, October. Onto: Athens; November. Sourn Carouina: Bamberg,
Oconee; May, July, December. Souta Daxota: Bon Homme; June. TrExas:
Bowie, Victoria; March. Vrrernta: Fairfax; April.

10. Melanotus emissus (LeConte)

Fiaurss 40, 9k; Puatrr 1h

Cratonychus emissus LeConte, 1853, p. 478.
Melanotus emissus.—Candéze, 1860, p. 372.—Thomas, 1941, p. 260.

Matz.—Similar to verberans; antenna reddish brown and lighter
than rest of body, which is generally dark reddish brown, erect male
hairs of antenna dense and conspicuous, hind angles of pronotum

sometimes lighter in color than rest of pronotum, pronotal punctures

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

denser. Genitalia differ in being shorter, aedeagus wider (but not as
wide as verberans), apical blade broader, and bearing four bristles.

Elytron: 6.60.6 mm (5.9-8.4). 32 spec.

FErmMALe.—Not positively identified (see discussion below).

Typr.—Holotype of emissus: Male, labelled with orange circle
“Ga.”’ (MCZ). Damaged specimen only one in type series.

DistriBuTIon.—Maryland and North Carolina west to South
Dakota and Kansas.

ALABAMA: Mobile; April. Iowa: Story; June. Kansas: Douglas; no date.
Maryuanp: Baltimore; July. NErsBRaAsKA: Lancaster; July. Norra CARo.uina:
Bladen, New Hanover, Pender; May. Soura Dakota: Union, Yankton; June,
July. Virarnra: Elizabeth City; May.

In the material studied there are two different females associated
with the males on the basis of similarity in appearance and locality.
One seems to be the same as the female of verberans and the other a
distinct form (fig. 94). The latter, from Ash, N.C., is tentatively
assigned to emissus but not with certainty. This also poses the
question as to the proper association of sexes in verberans, but that
cannot be answered at this time.

ll. Melanotus lanei Quate, new species

Ficures 5a, 9h; PuateE If

Mae.—Same as communis, but smaller in size, generally darker
in color; antenna as in communis, ratio of segments 2:3:4=5:9:12,
segment 4 about 1) times as long as wide; pronotum narrower, ratio
of width/length=1.0-1.1, vestiture thicker, pronotal punctures
smaller and denser; genitalia shorter, aedeagus evenly tapering from
base to apex.

Elytron: Holotype 6.6 mm; paratypes 6.7+0.6 mm (5.9-8.2).
17 spec.

FEMALE.—Same as male, but larger. Genitalia as communis with
diverticulum branching from spermathecal duct, differs from communis —
in that bursa has fewer spines, from 15 to 20 (allotype with 17). |

Elytron: Allotype 8.0 mm; paratypes 7.6+0.5 mm (6.7-8.7). 21
spec.

DistRIBUTION.—Pennsylvania south to Florida and west to South
Dakota and Kansas.

Holotype, male, allotype, female (USNM): Lincoln, Nebraska, July 2, 1957,
black light trap.

Paratypes, 9 males, 13 females (USNM, NU, CAS, MCL). Washington, D.C.,
May 22, Hubbard and Schwarz. PrNNsyLvANIA: Darby, Delaware Co., July 4.
Iowa Ames, Story Co., June 6, 1931, J. F. Glawe; same, June 10, 1925. Sour
Daxorta: Brookings, June 22, 1943, H. C. Severin; Kimball, Brule Co., June 17,
1947, Severin; Turner Co., June 15, 1929, Severin. NrsBrasKa: Lincoln, June 27,
1909, F. H. Shoemaker; Fairmont, Fillmore Co., June 17, 1914, G. W. Denning;

NO. 3568 MELANOTUS—QUATE AND THOMPSON 31

Bradshaw, York Co., July 9, 1917, Amaranthus, EH. J. Yates. Kansas: Douglas
Co., J. C. Bridwell; Topeka, Popenoe. ArKkansas: Hope, Hempstead Co.,
May 19, 1931, June 11, 1954, J. W. Green; “Vinita, Ind. T., June 7-8, ’99, Wick-
ham.” FrLoripa: Lake City, Columbia Co., 1933, Wickham.

It is a pleasure to dedicate this species to M. C. Lane in recognition
of his long and diligent studies of the Elateridae that are greatly
advancing the taxonomic knowledge of this large family.

12. Melanotus pilosus Blatchley

Fiaures 5b, 91; PLatE 1g

Melanotus pilosus Blatchley, 1910, p. 751.—Fenton, 1926, p. 502.—Blatchley,
1930, p. 35.—Dietrich, 1945, p. 59.

Mate.—Similar to communis, but smaller and more densely covered
with whitish or gray vestiture; antenna with erect hairs very dense,
segment 3 short, ratio of segments 2:3:4=4:7:12, segment 4 broad,
twice as long as wide; frontal margin very thin, upturned and nearly
straight on center rather than rounded; pronotal punctures smaller
and denser than communis; paramere with short, broad apical blade
and deep indentation basad of blade.

Elytron: 6.9+0.4 mm (6.4-8.0). 24 spec.

FEMALE.—Similar to male, bursa small, with about 30 spines,
spines with large bases; spermathecal duct and diverticulum arising
separately from bursa; accessory gland far basad of spermathecal
duct.

Elytron: 6.9 mm (6.5-—7.2).

Typr.— Holotype: Male, Posey Co., Ind., June 6, 1904, Blatchley
(PU).

DistripuTion.— Central U.S.

ARKANSAS: Washington; May. I.uinois: Wabash; June. INpIANA: Bar-
tholomew; Tippecanoe; June. Iowa: Linn; June. Kansas: Riley, Sedgwick ;

June. Montana: Big Springs St. Pk.; June. Nersprasxa: Antelope, Chase,
Custer, Keith, Lancaster, Lincoln, Scotts Bluff; June. Oxuanoma: Hayes; July.

13. Melanotus opacicollis LeConte
Fiaurss 5d,e, 97; PLats 17
Melanotus opacicollis LeConte, 1866, p. 390.—Blatchley, 1910, p. 751.

Matz.—Similar to communis but vestiture of pronotum very dense,
velvety, pronotal punctures numerous, dense, and so close together
as to appear granulose; frontal margin thin and protruding. Geni-
talia with evenly tapering aedeagus, paramere with marked inden-
tation below blade, slightly sinuous before center.

Elytron: 7.2 mm (6.9-7.5). 4 spec.

Frmate.—Similar to male. Bursa with 6 to 11 spines, much less
numerous than communis, spermathecal duct diverticulum branches
from spermathecal duct at distance clearly distal of base of duct.

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Elytron: 8.0+0.5 mm (7.1-8.8). 18 spec.

Typrs.— Lectotype: Female, “TIll./477” (MCZ). First specimen
in type series selected as lectotype by authors in 1962.

DistRIBUTION.— Indiana and Mississippi, west to Kansas.

ARKANSAS: Hempstead; May. I tuinots: Greene, Marion, Rock Is.; May,
June. Inprana: Owen; July. Iowa: no other data. Kansas: Linn, Mont-

gomery, Riley; June. Louistana; ? Co.; May. M&ussissippr: George, Greene;
May. Monrana: Boone, Callaway, Clinton, Jackson; May, July.

14. Melanotus clandestinus (Erichson)
Fiaure 5f

Cratonychus clandestinus Erichson, 1842, p. 112.—LeConte, 1853, p. 474.

Melanotus clandestinus——Candéze, 1860, p. 343.—Thomas, 1941, p. 258.—
Dietrich, 1945, p. 55.—Fattig, 1951, p. 20.

Melanotus peninsularis Candéze, 1889, p. 46.—Fattig, 1951, p. 22. [New
synonymy.]

Melanotus perplexus Blatchley, 1920, p. 46; 1930, p. 44.—Fattig, 1951, p. 22.

Maue.—Body color reddish brown, head and pronotum usually
darker than appendages and elytra, covered with white (sometimes
yellowish) vestiture evenly distributed over body.

Head: Front with prominent anterior transverse depression, margin
strongly protuberant above clypeus, a little angulate and blunt across
center when viewed from above; parantennal fovea obsolescent,
nasale not at all developed. Mandible without pit. Antenna ex-
ceeds pronotum by 1-1} segments, ratio of segments 2:3:4=2:2:7,
segment 4 about 1% as long as maximum width; erect male hairs
long and very dense.

Pronotum about as long as wide (variable and apparently Texas
specimens usually narrower), punctures very dense and subequal to
those on front, vestiture little finer than on elytra; sides rounded,
with slight concavity at base of hind angles; hind angle subparallel,
each with single carina extending slightly beyond base of hind angle.

Genitalia as figured; paramere without apical blade.

Elytron: 5.9+0.5 mm (5.0-6.9). 47 spec.

Frmate.— Unknown to us.

Typres.—Lectotype of clandestinus: Male, ‘Amer. Sept.” No.
17056 (ZM). Selected by Quate, 1960. One other specimen in type
series.

Lectotype of peninsularis: Male, “Florida, ex. coll. Morrison”’
(Roy. Mus. Nat. Hist., Brussels). Selected by Quate, 1960. Eight
other males in type series.

Holotype of perplexus: Male, Dunedin, Fla., July 5, 1915, W.S.B.
(Plu):

DistRIBuTION.— New Jersey south to Florida and west to Texas.

|
|
|
|

NO. 3568 MELANOTUS—QUATE AND THOMPSON 33

ALABAMA: Mobile; July. Fuoripa: Highlands, Hillsborough, Lee, Levy,
Palm Beach, Pinellas, Suwannee, Volusia; May, July. Mzussissippr: George,
Jackson, Pearl River; June. New Jersey: Atlantic, Burlington, Cape May,
Ocean; June, August. Norra Carouina: Columbus, Johnston, Moore; June,
July. Sours Carouina: Charleston; May. Texas: Bexar; July.

M. clandestinus is one of the few, small species with the elytra less
than 7 mm. It is easily recognized by the dense pronotal punctures,
small third antennal segment, lack of parantennal fovea and mandibu-
lar pit, in addition to the small size.

15. Melanotus ignobilis Melsheimer
Ficures 5g, 10a

Melanotus ignobilis Melsheimer, 1846, p. 152.—Candéze, 1860, p. 371.—Blatchley,
1910, p. 746 —Thomas, 1941, p. 258.—Dietrich, 1945, p. 55—Jewett, 1946,
p. 9.—Severin, 1949, p. 16.

Cratonychus ignobilis—LeConte, 1853, p. 474.

Cratonychus secretus LeConte, 1853, p. 474. [New synonymy.]

Melanotus secretus—Candéze, 1860, p. 344-—Thomas, 1941, p. 258.—Deen
and Cuthbert, 1955, p. 193.—Fattig, 1951, p. 20.

Mauzs.—Body color dark reddish brown, but some specimens
lighter, covered with yellowish vestiture evenly distributed over body.

Head: Front with shallow anterior transverse depression, margin
very narrow, not strongly protuberant above clypeus, evenly rounded
or anteriorly flattened when viewed from above; parantennal fovea
small and shallow, nasale flattened, width usually twice height.
Mandible without pit. Antenna exceeds tip of hind angle by 1-2
segments, largely testaceous with reddish-brown streak on either
side, lighter in color than rest of body; segments 2 and 3 subequal,
ratio of 2:3:4=2:3:6, flagellar segments nearly quadrangular;
segment 4 about twice as long as maximum width; erect male hairs
short but dense.

Pronotum usually wider than long, ratio of width/length=1.0-1.2;
punctures at center usually separated by distance equal to less than
own diameter, subequal to those on front, denser on margins; sides of
pronotum rounded with slight concavity at base of hind angles, hind
angles subparallel with one strong carina extending well cephalad
of base. Genitalia as figured; paramere without apical blade.

Elytron: 7.2 mm (5.2-9.3). 13 spec.

FremaLe.—Antenna short of pronotal base by about 1 segment;
segment 3 about 1% times 2, but about % times 4; internal genitalia
as figured, bursa with 3 groups of dense, setiform spines near apex;
accessory gland short and clavate, spermathecal duct very short,
diverticulum arising near center of duct.

Elytron: 7.6 mm (5.2-10.0). 24 spec.

221-532—67——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Typrs.—Lectotype of ignobilis: Male, no data (MCZ). Un-
labelled specimen in LeConte Collection, assumed to be type, agrees
with our interpretation of the species; to forestall possible confusion,
specimen selected as lectotype by authors in 1962.

Lectotype of secretus: Male, labelled with orange circle “Southern
States” (MOCZ 2510). First specimen of type series selected as
lectotype by authors in 1962.

Distripution.—New York south to Florida and west to South
Dakota and Texas.

AvapamMa: Colbert, Mobile; May, June. Arkansas: Washington; June, July.
Fiorma: Highlands, Alachua, Putnam, Volusia; April, May. Gzoreia:
Charlton, Fulton; July. Inurnors: Marshall, Putnam; June, July. INDraANa:
Clark; June. Lovurstana: Madison; June, July. Mussissrppr: Greene, Jack-
son, Perry; May, July. Montana: Phelps, St. Louis; June. NErBRaska:
Lancaster; July. New Jersey: Ocean; June, July. New Yorx: Niagara,
Tompkins; July. Norra Caroxina: Wake; June. PENNSYLVANIA: Dau-
phin; June. Sours Caronina: Charleston, Oconee; March, July. Sourn
Dakota: Brookings; July. TENNEssEE: Morgan; June, August. TExas:
McCulloch; July.

M. ignobilis is a distinctive species easily recognized by the short
third antennal segment, quadrate yellowish flagellar segments, lack
of mandibular pit, and dense pronotal punctures. The female in-
ternal genitalia are quite different from any other North American
Melanotus, and the male genitalia are also markedly different from
other species.

We can find no significant differences between secretus and ignobilis.
The former name was applied to smaller specimens of ignobilis which
often have the prothorax as long as wide and are usually lighter in
color. The male and female genitalia of these smaller, lighter colored
specimens do not differ from ignobilis, and we consider them as a part
of the variation range of that species; hence, secretus is synonymized
with ignobilis.

The americanus Group
(Mandible with pit)
16. Melanotus depressus (Melsheimer)
Fieures 57, 106; PLatE 1j

Ctenonychus depressus Melsheimer, 1846, p. 151.

Cratonychus depressus.—LeConte, 1853, p. 475.

Melanotus depressus.—Candeéze, 1860, p. 345.—Blatchley, 1910, p. 755.—Thomas,
1941, p. 258.—Dietrich, 1945, p. 55.—Fattig, 1951, p. 20.

Ctenonychus parumpunctatus Melsheimer, 1846, p. 151.

Cratonychus parumpunctatus.—LeConte, 1853, p. 478.

Melanotus parumpunctatus.—Candeze, 1860, p. 355.—Blatchley, 1910, p. 754.—
Thomas, 1941, p. 260.—Dietrich, 1945, p. 57.—Fattig, 1951, p. Bile

NO. 3568 MELANOTUS—QUATE AND THOMPSON 35

Melanotus divarcarinus Blatchley, 1910, p. 754.—Thomas, 1941, p. 259.—Dietrich,
1945, p. 56.—Severin, 1949, p. 16.—Deen and Cuthbert, 1955, p. 193.
[New synonymy.]

Melanotus lixus Blatchley, 1910, p. 754. [New synonymy.]

Mauz.—Body color dark reddish brown, covered with white or
yellow hairs evenly distributed over body.

Head: Front with pair of shallow anterolateral depressions not
strongly protuberant above clypeus, evenly rounded or a little angu-
late and flattened when viewed from above, parantennal fovea
crescent shaped, small, but moderately deep and well defined; nasale
obsolescent. Mandible with shallow, slitlike pit. Antenna exceeds
tip of hind angle by 1 to 1% segments; reddish brown, ratio of seg-
ments 2:3:4=3:5:6, segment 4 about twice as long as maximum
width; erect male hairs short but dense.

Pronotum wider than long; punctures at center usually separated
by distance equal to little less or little more than own diameter,
equal to or little smaller than those on front; sides of pronotum straight
behind small anterior curvature, divergent posteriorly; hind angles
subparallel, strong carina extending cephalad of base of hind angle
Genitalia as figured; paramere without apical blade.

Elytron: 8.740.6 mm (6.5-10.2). 85 spec.

FEMALE.—Similar to male. Pronotum with sides often more
strongly divergent; antenna exceeds pronotum by %-1 segment;
internal genitalia as figured, bursa long and coiled in loop, with
scattered, peglike spines nearly throughout entire length, accessory
gland originates near base, clavate, spermathecal duct branches near
base.

Elytron: 9.30.7 mm (8.2-10.2). 44 spec.

Typres.—Lectotype of depressus: Male, labelled with red label
“PA.” (MCZ). First species of type series selected as lectotype by
authors in 1962.

Lectotype of parumpunctatus: Female, labelled with red label
“Pa.” (MCZ). First specimen of type series selected as lectotype by
authors in 1962. Second specimen is ignobilis.

Holotype of dwarcarinus: Male, Posey Co., Ind., June 2, 1909,
W.S. B. (PU).

Holotype of lizus: Female, Posey Co., Ind., July 6, 1912, W.S. B.
(PU):

DistrisutTion.—Connecticut to North Carolina, west to Nebraska
and Texas.

ARKANSAS: Benton, Hempstead, Pike, Washington; May, July. CoNnNeEcTI-
cut: Fairfield, New Hampshire; June. Groraia: Catoosa, Fulton, Gwinnett,

Lumpkin, Moran, White, Worth; May, July. Inuinoi1s: Champaign, Clay,
Jackson, Johnson, Kane, Marion, McLean, Peoria, Pope, Putnam, Rock Island,

221-532—67——4

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Scott, Union, Vermilion, Washington; April, July. INnprana: Spencer, Tippe-
canoe; June. Iowa: Henry, Story; June, August. Kansas: Chautaugua,
Cowley, Douglas, Miami, Montgomery, Riley; May, June. Kentucky: Jeffer-
son; June. Lovuistana: Madison; June. Marynianp: Anne Arundel, Washing-
ton; June, July. Mussissippi: George, Greene; April, May. Montana: Calla-
way, Clinton, Jefferson, Lawrence, St. Louis, Webster; April, July. NEBRASKA:
Cass, Dakota, Lancaster; June. New Jersey: Lakehurst, Morris, Ocean,
Warren; June, August. Nrw Yorx: Dutchess, Essex, Seneca, Tompkins; May,
July. Norta Carouina: Guilford; June. Onto: Clifton, Delaware, Erie,
Franklin, Greene, Hocking, Pickaway, Ross; May, July. OxLaHomaA: Payne,
Sequoyah; July. PrENNsytvania: Adams, Allegheny, Northampton, Phila-
delphia; May, July. TENNEssEE: Sevier; June. Trxas: Bexar, Brazos, Brew-
ster, Dallas, Kerr, Madison, Val Verde; April, July. Vireainta: Fairfax, Loudoun;
May, July. Wasuineton, D.C.: June. West Virarnia: Greenbrier; July.

Melanotus depressus is subject to considerable variation. Most
conspicuous is the range in overall size. The variation in elytron
length from 6.5-10.2 mm is about as large as any North American
Melanotus. There appears to exist a cline in the males with size of
the elytron diminishing to the west and south, as shown in the fol-
lowing chart. Curiously, the female does not appear to vary in
the same way.

Variation in elytron length of M. depressus (average length in mm):

East Coast to Arkansas,
Indiana Illinois Missouri Kansas Nebraska Teras
o 2 a ? a g a
93 9.1 8.8 9.3 8. 5 9.3 7.9

Nonclinal variation exists in the size of the parantennal fovea and
mandibular pit. The pronotal punctures vary in size and density
but are almost always as large as those on the front, which are larger
than in morosus, and never as compact as in cribulosus. The sides of
the pronotum are usually markedly divergent and are a useful feature
in identifying the species when one is familiar with it. Few speci-
mens have the sides subparallel and lack the characteristic wedge
shape.

Other than a broader than usual aedeagus seen in some northern
specimens, the male genitalia are constant. The female genitalia
also show little variation and are characteristic of the species. We
have found the genitalia to be reliable features of identification, and
dissected specimens can be identified positively with little difficulty.

The bulk of specimens of the ‘americanus group” which we have
seen belong to either morosus or depressus. We have dissected a large
number of specimens to verify our identifications. We concluded
that there are only two species in this section of the genus. We can
find nothing to support the recognition of the third species. The
type of parumpunctatus appears to us to be depressus, and it is within

No. 3668 MELANOTUS—QUATE AND THOMPSON 37

the size range of that species. MJ. depressus has page priority over
parumpunctatus and is the preferred name.

17. Melanotus morosus Candéze
Fiaures 5h, 10e
Cratonychus longulus LeConte, 1853, p. 480. [Not LeConte, 1853, p. 473.]
Melanotus morosus Candéze, 1860, p. 346.—Fattig, 1951, p. 22.
Melanotus debilis Blatchley, 1910, p. 754; 1930, p. 36. [New synonymy.]
Melanotus simulans Blatchley, 1927, p. 140; 1930, p. 48.—Fattig, 1951, p. 22.

Matz.—Body color reddish brown, covered with white hairs, evenly
distributed over body.

Head: Front with shallow or no anterior depression, margin not
strongly protuberant above clypeus, evenly rounded or flattened
anteriorly; parantennal fovea moderately large and deep, circular;
nasale about 1% times as wide as high. Mandible with deep teardrop-
shaped pit; palpus light reddish brown. Antenna exceeds tip of hind
angle by 1-2 segments, reddish brown, ratio of segments 2:3:4=3:5:6,
segment 4 about twice as long as maximum width; erect male hairs
short but dense and easily seen.

Pronotum wider than long, ratio of width/length=1.04-1.18;
punctures at center usually separated by distance equal to more
than own diameter, smaller than punctures on front; sides of pronotum
usually straight behind anterior curvature and subparallel, but often
rounded with concavity at base of hind angles; hind angles slightly
divergent, carina usually extending cephalad of base of hind angle.
Genitalia as figured; paramere without apical blade.

Elytron: 7.8+0.6 mm (6.3-8.8). 40 spec.

Frmate.— Antenna extends to or beyond base of pronotum by 1
segment; genitalia as figured, bursa with very dense, shaggy appearing,
quadrate patch of setiform spines and few peglike spines on apical
extension, accessory gland very slender, spermathecal duct branches
near center.

Elytron: 8.1+0.6 mm (7.0-9.0). 20 spec.

Typrs.— Lectotype of longulus: Male, labelled with orange circle
“Southern States” (MCZ 2524). First specimen of type series selected
as lectotype by authors in 1962. Second and 8rd specimens not
true types, labelled ““Tex.’’; 4th specimen apparently infaustus.

Holotype of debilis: Female, Marshall Co., Ind., June 26, 1904,
Mis. b. (PU).

Lectotype of simulans: Male, Dunedin, Fla., Mar. 15, 1918, W.S.B.
(PU).

In the Institut Royal des Science Naturelles, Brussels, there is a
male from Iowa City, Iowa, labelled as the type of morosus. In the
British Museum (Natural History) there is another female from
Georgia labelled as the type of morosus. Neither of these specimens

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

are true types, since morosus is a substitute name for the secondary
homonym longulus, and the types of morosus Candéze are those
selected by LeConte for longulus (1853, p. 480, not 1853, p. 473).

DisTRIBUTION.—Quebec to Florida and west to Nebraska and
Texas.

ALABAMA: Mobile; June. ARKANSAS: Clark, Hempstead, Lawrence; April,
June. Connecticut: Tolland; May. FLorimpa: Manatee, Osceola, Pinellas;
March. GeroreGta: Catoosa, Clarke, Cobb, Fulton, Hall, Rabun, Seminole,
Thomas; March, June. ILLinois: Champaign, Cook, Knox, Putnam, Scott,
Washington; May, July (Nov.).. Maryztanp: Anne Arundel, Baltimore, Har-
ford, Prince Georges; June, October. Massacuusertts: Barnstable, Middlesex,
Nantucket, Worcester; May, July. Myussissrppi: George, Greene, Harrison,
Lamar; April, May. Montana: Jefferson, Lawrence, St. Louis; April, July.
Nespraska: Douglas; June. New Hampsuire: Carroll; July. New Jersey:
Atlantic, Camden, Ocean; May, August. New York: Orange, Suffolk, Tompkins,
Washington; June, August. Norra Caroxiina: Buncombe, Duplin, Hyde,
Moore, Sampson, Swain, Wake, Wayne; April, July. Onto: Adams, Delaware,
Fairfield, Franklin, Greene, Hocking, Scioto; June, August. PENNSYLVANIA:
Allegheny, Indiana, Monroe, Northampton, Philadelphia; April, July. Sours
Carouina: Oconee; April. TENNESSEE: Carter, Sevier; June, September.
Texas: Bastrop; April. Virainia: Fairfax, Fauquier, Prince Edward; May.
WasHINGTON, D.C.: April, June. Ontario: “Go Home Bay”; June. QuEBEC:
Windsor Co.; July.

M. morosus is most likely to be confused with depressus. In general,
morosus may be separated from that species by the small pronotal
punctures, subparallel sides of the pronotum with pronounced anterior
curve, and a more slender, jutting frontal margin. Less frequently,
morosus is lighter in color than depressus. Typical specimens are not
difficult to identify when one is familiar with the characters, but small
variations (artificial or natural) make it necessary to dissect the
genitalia for positive identification. The thick mat of spines in the
bursa easily identifies the females. The male genitalia are charac-
terized by the angulate paramere, rather than evenly rounded as in
depressus, the moderately dense hairs at the apex of the paramere,
and the rather broad, tapering aedeagus.

Specimens from Florida are usually smaller and the pronotal width/
length ratio is more variable, but they do not seem to differ signifi-
cantly from other morosus. M. simulans is regarded as a synonym of
morosus.

18. Melanotus cribulosus (LeConte)
Fiaures 5j, 10d

Cratonychus cribulosus LeConte, 1853, p. 478.
Melanotus cribulosus.—Candéze, 1860, p. 357.—Hyslop, 1915, p. 17.—Thomas,
1941, p. 260.—Severin, 1949, p. 17.—Srivastava, 1958, p. 87.
Matre.—Body color reddish brown, covered with fairly dense whitish
hairs evenly distributed over body.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 39

Head: Front with shallow depression, margin not strongly pro-
tuberant above clypeus, rounded to flat when viewed from above;
parantennal fovea small, but well defined; nasale obsolescent. Man-
dible with pit variable from ovoid to slit shaped. Antenna exceeds
tip of hind angle by % to 1% segments, reddish brown, ratio of seg-
ments 2:3:4=2.5:4:5, segment 4 about 1} times as long as maximum
width; erect male hairs very short and inconspicuous.

Pronotum wider than long, ratio of width/length=1.05-1.13; punc-
tures at center usually separated by distance equal to less than own
diameter, as large as those on front; sides of pronotum mostly straight
behind moderate anterior curvature, with slight concavity at base of
hind angles; hind angles divergent, carina extending cephalad of
base of hind angle. Genitalia as figured; paramere without apical
blade.

Elytron: 7.5+0.4 mm (6.4-8.5). 24 spec.

FEMALE.—Similar to male, except antenna slightly shorter in com-
parison to pronotum; genitalia as figured, bursa with few naillike
spines with large bases, accessory gland short and clavate, sperma-
thecal duct branches near base.

Elytron: 7.2-7.3 mm.

Typrs.—Lectotype of cribulosus: Male, labelled with green circle
“Nebr. Terr.’”’ (MCZ 2519). First specimen of type series selected as
lectotype by authors in 1962.

DistripuTIon.—Central U.S.

Inuino1s: Champaign, Kane, Mason, McHenry, Warren; June, July. Kansas:
Cowley, Douglas, Gove, Greenwood, Gray, Kingman; June, July. NEBRASKA:
Lancaster, Saline; June, July. Ox uanoma: Cleveland; April, May. Sovurn
Daxora: Haakon; June.

In superficial appearance, cribulosus looks like a member of the
communis complex; however, the mandibular pit removes it from that
group. The shallow parantennal fovea and dense pronotal punctures
distinguish it from other members of the americanus group.

19. Melanotus corticinus (Say)
Fiaures 5n, 107

Elater corticinus Say, 1823, p. 174; 1839, p. 183.

Cratonychus corticinus.—LeConte, 1853, p. 473.

Cratonychus vetulus Erichson, 1842, p. 105. [New synonymy.]

Melanotus vetulus—Leng, 1920, p. 175.

Melanotus glandicolor Melsheimer, 1846, p. 152.—Emmons, 1854, p. 88.—Can-
déze, 1860, p. 371.—Blatchley, 1910, p. 745.—Thomas, 1941, p. 259.

Cratonychus glandicolor—LeConte, 1853, p. 477.

Matz.—Body color reddish brown, covered with whitish vestiture,
evenly distributed over body.

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Head: Front with shallow depressions, margin thick, dark reddish
brown, not strongly protuberant above clypeus, angulate and blunt
when viewed from above; parantennal fovea large, deep and rounded,
foveal tragus weakly developed; nasale usually as wide as or a little
less than height, well developed. Mandible with large slitlike pit;
palpus reddish brown, antenna exceeds tip of hind angle by 2 or 3
segments, segments 2 and 8 subequal, flagellar segments elongate
and slender, segment 4 about twice as long as maximum width, erect
male hairs sparse, but long and easily seen.

Pronotum wider than long; punctures in center usually separated
by distance equal to less than own diameter, equal to size of punc-
tures on front; sides of pronotum straight and divergent posteriorly
with slight anterior curvature, hind angles divergent with carina
usually extending little cephalad of base. Genitalia as figured;
paramere with apical blade.

Elytron: 10.6 mm (10.1-11.8). 11 spec.

FremMaLe.—Similar to male; antenna 3 a little larger than 2, but
still much smaller than 4, so not intermediate in size between 2 and 4;
genitalia as figured, accessory gland slender, bursa with moderate
number of tacklike spines.

Elytron: 11.8 mm (11.0-12.7). 7 spec.

TypEs.—Types of corticinus: ‘United States,’”’ specimens lost.

Lectotype of glandicolor: Female, no locality data, but stated to
be Pa. (MCZ). First specimen of type series selected as lectotype
by authors in 1962, but other 2 specimens probably not true types.

Holotype of vetulus: Female, ‘America Sept.” (ZM). Only
specimen in type series.

DistriBpuTION.—Ontario, eastern U.S. west to Illinois and Missouri.

Iuuinois: Kankakee, Vermilion; June, July. INp1ana: Clark; June. Mary-
LAND: Baltimore; June. Montana: Montgomery; May. New JERSEY:
Warren; June. Norra Carouina: Buncombe; June. PENNsyLVANIA: Adams;
May. TENNESSEE: Sevier; June. Vireinrta: Norfolk; May. Onrario: Pr.
Edward.

Melanotus corticinus is one of the more easily identified species;
the small third antennal segment, large parantennal fovea, and large
size make its recognition relatively easy. Females with a larger
third antennal segment may cause a little trouble in running through
the key, but even though the third segment is larger than the second,
it is not intermediate in size between the second and fourth and should
not be taken out in the wrong part of the couplet.

20. Melanotus sagittarius (LeConte)
Figures 5m, 10c; Puats 1k

Cratonychus sagittarius LeConte, 1853, p. 480.

No. 3568 MELANOTUS—QUATE AND THOMPSON 41

Melanotus sagittarius.—Candéze, 1860, p. 547.—Blatchley, 1910, p. 751.—Thomas,
1941, p. 260.—Dietrich, 1945, p. 58.
Melanotus carinus Blatchley, 1910, p. 752; 1930, p. 35. [New synonymy,]

Matzr.—Body color reddish brown, covered with yellowish vesti-
ture, evenly distributed over body.

Head: Front with pair of shallow anterolateral or an anterior
transverse depression, margin angulate with blunt apex when viewed
from above, not strongly protuberant above clypeus; parantennal
fovea large and deep, as long as antennal segment 2, opening rounded,
foveal tragus weakly developed; nasale protuberant, width 1 to 1%
times height. Mandible with deep, teardrop-shaped pit. Antenna
exceeds tip of hind angle by 2 to 2% segments, reddish brown, ratio of
segments 2:3:4=3:6:9, flagellar segments rather elongate, segment
4 about twice as long as maximum width; erect male hairs long,
rather dense, and easily seen.

Pronotum wider than long, ratio of width/length=1.14(1.05-1.22);
punctures at center usually separated by distance equal to about
own diameter, subequal to size of those on front; sides of pronotum
nearly straight beyond small anterior curvature, sometimes with
slight concavity at base of hind angle. Genitalia as figured; paramere
with apical blade.

Elytron: 9.90+0.54 mm (8.70-10.80). 50 spec.

Frema.Le.—Antenna exceeds tip of pronotum by 1 to 1% segments;
genitalia as figured, bursa with moderate number of peglike spines;
accessory gland little longer than bursa, arises well before sperm duct;
sperm duct branches near base.

Elytron: 10.2+0.9 mm (8.5-12.0). 31 spec.

Typrs.—Lectotype of sagittarius: Male, labelled with white circle
“Hastern States & Canada” (MCZ). First specimen in type series
selected as lectotype by authors in 1962. Second specimen labelled
with pink circle “Middle States” is probably the type mentioned in
the description but is damaged and cannot be identified definitely,
so has not been selected. There appears no question that LeConte
studied the first specimen and that it agrees with current interpre-
tation of sagittarius, and its selection as the lectotype will fix the name
of the species without disturbing the present nomenclature.

Holotype of carinus: Male, Lake Co., Ind., June 4, 1905 (PU).

DistriBuTion.—New York to Florida and west to Illinois.

Fioripa: Pinellas; February, April. Inuinors: Putnam; July. INpraNa:
Marion; May. Maryann: Baltimore; July. Massacnuusrerts: Hampshire,
Middlesex; June, July. Mussissrpp1: George; April. Nrw Jersry: Middlesex;
July. New Yor: Dutchess, Rockland, Tompkins, Westchester; June, July.

Ox1o: Athens; June. Prnnsytvanra: Allegheny, Bradford, Monroe; May,
July. Vireinia: Page; July. West Virernia: Hardy; July.

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

21. Melanotus hyslopi Van Zwaluwenburg
Fiaurss 51, 10f

Melanotus hyslopi Van Zwaluwenburg, 1921, p. 210.—Thomas, 1941, p. 261.—
Dietrich, 1945, p. 59.—Fattig, 1951, p. 22.

Mae.—Body color reddish brown, covered with white or yellowish
vestiture, evenly distributed over body.

Head: Front with pair of shallow anterolateral depressions, not
strongly protuberant above clypeus, angularly produced into trun-
cated projection when viewed from above; parantennal fovea large
and deep, excavated as deep as length of antennal segment 2, crescent
shaped, foveal tragus weakly developed; nasale strongly protuberant,
as wide as high. Mandible with deep, rounded, or ovate pit. An-
tenna exceeds tip of hind angle by 2-3 segments, segment 3 inter-
mediate to 2 and 4, reddish brown, flagellar segments rather elongate,
segment 4 about twice as long as maximum width; erect male hairs
long, but sparse.

Pronotum wider than long, ratio of width/length=1.05-1.18;
punctures at center usually separated by distance equal to more than
twice own diameter, smaller than those on front; sides of pronotum
straight behind anterior curvature and divergent posteriorly; hind
angles divergent, with carina usually extending well cephalad of base
of hind angle. Genitalia as figured; paramere without apical blade.

Elytron: 8.5+0.3 (7.6-9.1). 60 spec.

FremMa.e.—Antenna exceeds tip of hind angle by 1-2 segments;
internal genitalia as figured, bursa with large swelling at base of ac-
cessory gland, gland clavate, elongate apex of bursa looped, sper-
mathecal duct branches near base, diverticulum very short and
curled.

Elytron: 8.7+0.4 mm (7.6-9.4). 55 spec.

Typr.—Holotype: Male, South Mountain, Md. (USNM 24561).
Not studied by authors.

DistrisuTion.—New Hampshire south to North Carolina and
west to Wisconsin and Illinois.

Connecticut: Litchfield; June. I~Ltino1is: Champaign, McHenry; July, Sep-
tember. Maryianp: Anne Arundel, Baltimore, Frederick; June, July. Mas-—
SACHUSETTs: Barnstable, Bristol, Essex, Hampshire, Middlesex; June, August.
Micuiecan: Oceana; August. Mussissrppi: Forrest; April. New HampsuHirReE:
Carroll; August. Nrw Jersry: Atlantic, Bergen, Camden, Ocean; May, July.
New York: Erie, Essex, Niagara, Rockland, Suffolk, Sullivan, Tompkins; May,
September. Nortu Carouina: Buncombe, Macon; July. Onto: Delaware,
Fairfield, Hocking, Summit; May, July. Prnnsytvanra: Dauphin, Monroe,
Northampton, Philadelphia; May, July. TENNxEssEE: Sevier; June, July.
Vireinia: Giles, Rockbridge; July, August. Wisconsin: No other data.
Ontario: Pr. Edward Co.; June.

No. 3568 MELANOTUS—QUATE AND THOMPSON 43

M. hyslopi and sagittarius have larger parantennal fovea than any
other North American Melanotus and can be distinguished by that
character in addition to more usual features of ordinary size, mandib-
ular pit, and intermediate sized third antennal segment. They may
be separated from each other by hyslopi generally being smaller,
having sparser pronotal punctures, longer antennae, larger paranten-
nal fovea, and a more angulate frontal margin.

22. Melanotus prasinus Blatchley
Fiaures 6a, 10g,h
Melanotus prasinus Blatchley, 1910, p. 752; 1930, p. 35.—Thomas, 1941, p. 258.

Matze.—Body color reddish brown, covered with white vestiture
evenly distributed over body.

Head: Front with pair of shallow anterolateral transverse depres-
sions, margin dark reddish brown, sometimes strongly protuberant
above clypeus and turned up in front, fairly thin, parantennal fovea
moderately small, but well defined, are shaped, with small foveal
tragus; nasale poorly developed 1%-2 times as wide as high. Man-
dible with deep slit or teardrop-shaped pit. Antenna exceeds tip of
hind angle by 1-2 segments, reddish brown, ratio of segments 2:3:4=
3:4:5, segment 4 about twice as long as maximum width; erect male
hairs short and rather sparse but easily seen.

Pronotum usually wider than long, ratio of width/length=1.0-1.07;
punctures at center usually separated by distance equal to about
twice own diameter, slightly denser on margins, smaller than punc-
tures on front; sides of pronotum gently rounded and divergent
posteriorly on anterior half, with concavity at base of hind angles;
hind angles divergent, carina weak, extending little cephalad of base
of hind angle. Genitalia as figured; paramere without apical blade.

Elytron: 6.8 mm (6.2—7.3). 14 spec.

FrmMALe.—Similar to male; bursa with moderate number of pointed,
plaquelike spines; accessory gland originates near apex of bursa.

Elytron: 7.5mm. 1 spec.

Typr.—Holotype of parasinus: Male, Vermilion Co., Ind., June
15, 1904, W. S. B. (PU).

DistrisuTiIon.—Connecticut to North Carolina, west to Illinois and
Missouri.

Connecticut: Windham; August. ItitNois: Champaign; July. MAaryLAND:
Plummers Isl.; April, June. Massacuusretts: Barnstable, Middlesex; July.
Montana: Carter; June. New Hampsaire: Hillsboro; no date. New JerRsEy:
Bergen, Cape May; June, July. New York: Kings; July. Norra Caro.ina:

Buncombe; July. Onto: Hocking, Lucas; June. PENNsyLVANIA: ‘Estes’;
July, TENNESSEE: Smith; no date.

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

The few specimens of prasinus available to us do not give a satis-
factory picture of this species, and we know little of its variation or
geographical distribution. We have three more males and two fe-
males that might belong to prasinus, but they differ in outline and
shorter, less jutting frontal margin and are questionably identified as
prasinus and are not included in the description.

Probably M. prasinus will most likely be confused with small speci-
mens of depressus from which prasinus differs in having smaller and
sparser pronotal punctures, a better developed nasale, and a more
strongly jutting frontal margin.

23. Melanotus piceatus Blatchley
Fiaures 5k, 107
Melanotus piceatus Blatchley, 1927, p. 141; 1930, p. 48.—Fattig, 1951, p. 22.

Mate.—Body color dark reddish brown to black, covered with
sparse yellowish vestiture, evenly distributed over body.

Head: Front with pair of shallow to marked depressions, margin
variable from thick and barely protruding beyond clypeus to thin and
extending beyond clypeus by little more than own width, rounded or
angulate and flattened anteriorly when viewed from above; par-
antennal fovea small, shallow, arc shaped; nasale flat, width about
1% times height. Mandible pit slit to teardrop shaped; palpus light
reddish brown. Antenna short of tip of hind angle by about 1 seg-
ment, reddish brown, segment 3 nearly as long as 4,4 about 1% times
maximum width, erect male hairs very short and evident only at base
of segments.

Pronotum wider than long; punctures at center usually separated
by distance equal to 1-2 times own diameter, subequal to those on
front; sides gently curved over anterior one-third or one-half, sub-
parallel or convergent posteriorly with small concavity; hind angles
slightly divergent with carina usually extending cephalad of base of
hind angles. Genitalia as figured; paramere without apical blade.

Elytron: 8.4 mm (7.5-10.2). 10 spec.

Frma.e.—Antenna short of tip of hind angle by 1-3 segments;
internal genitalia as figured, bursa large, with 50-100 scattered,
peglike spines.

Elytron: 8.6 mm (7.7-9.2). 7 spec.

Typr.—Holotype of piceatus: Male, Dunedin, Fla., March 17, 1922,
W.S.B. (PU).

DistTRIBUTION.—Southeastern U.S.

AutaBAMA: Colbert, Mobile; May, June. Fiorma: Brevard, Charlotte,
Highlands, Hillsborough, Pinellas, Volusia; February, May. Grora1a: Chatham;

March. SourH Carouina: Charleston, Georgetown, Pickens; May. TENNES-
SEE: Great Smoky Mts. Nat. Pk.; June.

No. 3568 MELANOTUS—QUATE AND THOMPSON 45

The male genitalia of piceatus are somewhat similar to cribulosus.
However, it is readily distinguished from that and other species by
the short antennae and other characters. It apparently has no close
relatives in North America and may be of Neotropical origin as
suggested by its distribution.

24. Melanotus difficilis Blatchley

Fiaure 69

Melanotus difficilis Blatchley, 1910, p. 751; 1930, p. 35.—McClure, 1933, p. 145.—
Dietrich, 1945, p. 59.

Maure.—Body color reddish brown, covered with whitish vestiture,
evenly distributed over body and longer than usual.

Head: Front without marked, transverse depressions but sometimes
with pair of shallow anterolateral ones, margin extending as thick
shelf in front of clypeus usually twice width of rim, anteriorly rounded
when viewed from above; parantennal fovea lacking or obsolescent;
nasale not developed. Mandible with small teardrop-shaped pit.
Antenna reddish brown, exceeds tip of hind angle by 3 to 3% segments,
segment 3 intermediate to 2 and 4, flagellar segments long and slender,
segment 4 twice as long as maximum width, following more slender;
erect male hairs very short but dense.

Pronotum flattened dorsally, wider than long; punctures at center
usually separated by distance equal to about own diameter, subequal
to those on front; sides of pronotum divergent posteriorly on anterior
half and parallel or a little convergent over remainder, hind angles
divergent, with carina usually extending about to base of hind angle.

Abdomen with last visible sternite usually inflated to give tip
enlarged appearance. Genitalia as figured; paramere with apical
blade.

Elytron: 7.5-9.7 mm.

FrmMate.—Unknown to us.

Typr.—Holotype of difficilis: Male, Posey Co., Ind., July 8, 1903,
Woo CEU):

DistrisuTion.—Central U.S.

Kansas: Douglas; June. Ixuinors: Alexander; June. Mussissrppi1: George;
May. Oxuanoma: Delaware; June. Texas: Austin, Dallas, Navarro; May.

25. Melanotus macer (LeConte)

Fiaures 6¢,f, 101

Cratonychus macer LeConte, 1853, p. 473.
Melanotus macer.—Candéze, 1860, p. 339.—Blatchley, 1910, p. 752.—Dietrich,
1945, p. 54.—Fattig, 1951, p. 20.

Mauz.—Body color reddish brown, covered with whitish hairs,
evenly distributed over body and longer than usual.

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Head: Front with conspicuous anterior, transverse depression,
margin extends as a thin shelve in front of clypeus by distance equal
to 2-3 times width of rim, angulate and flattened in front when viewed
from above and often with small, median notch; parantennal fovea
small but well defined, crescent shaped, opening much longer than
wide; nasale not developed. Mandible with small, slit-shaped pit.
Antenna long, exceeds tip of hind angle by 3-4 segments, reddish
brown, segment 3 intermediate to 2 and 4, flagellar segments slender,
nearly parallel sided, not serrate, segment 4 twice as long as maximum
width, following more slender; erect male hairs short, sparse, and
inconspicuous.

Pronotum wider than long; punctures at center usually separated
by distance equal to little less than own diameter, subequal to those
on front; sides of pronotum straight with little or almost no anterior
curvature and slightly divergent posteriorly; hind angles almost
parallel with carina, usually extending cephalad of base of hind angles.
Genitalia as figured; paramere with apical blade.

Elytron: 7.2-7.6 mm.

Frmate.— Unknown to us.

Typrs.—Lectotype of macer: Male, labelled with pink circle ‘Middle
States, incl. N.Y.” (MCZ 2507). First specimen of type series selected
as lectotype by authors in 1962.

Distrisution.—North Carolina west to Kansas.

Iowa: Story; June. Kansas: Cherokee, Riley; June, July. MUssissippi:
George; May. New York: Nassau; July. Norra Carouina: Moore, Wake;
May.

The two species difficilis and macer can be separated from other
North American Melanotus by the long, slender, nonserrate antennae
greatly extending beyond the pronotum, the obsolescent nasale, and
poorly developed or vestigial parantennal fovea. The two can be
distinguished without difficulty by characters cited in the key.

26. Melanotus testaceus (Melsheimer)
Fieures 6), 11ce,d

Ctenonychus testaceus Melsheimer, 1846, p. 151.

Melanotus testaceus—Thomas, 1941, p. 258.

Cratonychus dubius LeConte, 1853, p. 479. [New synonymy.]

Melanotus dubius.—Candéze, 1860, p. 372.—Thomas, 1941, p. 260.—Fattig, 1951,
p. 21.

Melanotus angustatus of authors, not Erichson, 1842.

Ma.z.—Body color reddish brown, covered with white vestiture,
evenly distributed over body.

Head: Front with pair of shallow anterolateral transverse depres-
sions, margin not strongly protuberant above clypeus, evenly rounded
or angulate and flattened across center; parantennal fovea small, deep

NO. 3668 MELANOTUS—QUATE AND THOMPSON 47

and well defined, semicircular; nasale protuberant, width 1 to 1% times
height. Mandible with deep, oval pit. Antenna exceeds tip of hind
angle by 1 to 1% segments along sides, reddish brown, ratio of segments
2:3:4=3:4.5:5, segment 4 about 1% times as long as maximum width,
erect male hairs short but easily seen.

Pronotum usually as wide as long, ratio of width/length=1.0-1.07;
punctures at center usually separated by distance equal to about twice
own diameter, smaller than punctures on front; sides of pronotum
behind small anterior curvature nearly straight except slight concavity
near center and markedly divergent posteriorly; hind angles divergent,
carina usually extending cephalad of base of hind angle. Genitalia
as figured; paramere slender, tapering, without apical blade.

Elytron: 7.0+0.5 mm (5.2-7.8). 29 spec.

FEMALE.—Similar to male; bursa with moderate number of scat-
tered, plaquelike spines, accessory gland originates near center of
bursa, spermathecal duct diverticulum originates not far from base of
duct, short and capitate.

Elytron: 6.9+0.3 mm (6.3-7.5). 23 spec.

Typrs.—Type of testaceus: No specimen found at MCZ definitely
labelled and unquestionably the type. In LeConte Collection a
teneral (and hence testaceous) male labelled (by LeConte?) “angus-
tatus.’’ We suspect this specimen is the type of testaceus and think
LeConte placed it with angustatus after he decided the two species
were synonymous (1853, p. 475).

Type of dubius: Male, labelled with orange circle “Southern States”’
(MCZ 2520). Only specimen in type series; the orange label indicat-
ing locality in South does not correspond with LeConte’s (1853, p. 479)
listing of N.Y., but nonetheless is regarded as the type.

Distripution.—New York to Florida, west to Missouri and Texas.

ARKANSAS: Hempstead, Washington; April, August. FLoripa: Pinellas;
April. Groraia: Fulton; June. I uinors: Alexander, Lawrence; April, July.
Montana: Callaway; May. New Jersey: Ocean; June. NEw York:
Suffolk, Tompkins; June, August. Norra Carouina: Wayne; April. Onto:
Delaware, Greene, Hocking, Scioto; May, June. Trxas: no data. VIRGINIA:
Nottoway; May.

This species has been identified as angustatus, and nearly all of our
identifications will bear a label with that name. However, the type of
angustatus is a specimen of americanus, and for a long time the species
has been misidentified in North America.

The name testaceus is used for this species, although it is based on
tenuous grounds, as there appears to be no unequivocal type specimen;
however, this seems to be the name most aptly applied to the species.

Undissected specimens will often be confused with trapezoideus,
although the characters in the key will be useful in separating the two

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

most of the time. It is advisable to dissect the genitalia from at least
a part of the collection being identified.

27. Melanotus trapezoideus (LeConte)
FIGuRES 6c, 116

Cratonychus trapezoideus LeConte, 1853, p. 475.

Melanotus trapezoideus.—Candéze, 1860, p. 348.—Blatchley, 1910, p. 752.—
Thomas, 1941, p. 259.—Dietrich, 1945, p. 55.— Jewett, 1946, p. 9.—Fattig,
1951, p. 20.

Mate.—Body color reddish to dark reddish brown, covered with
white or yellowish vestiture, evenly distributed over body.

Head: Front usually with pair of shallow anterolateral depressions;
margin dark reddish brown, thinner in center, angulate and flattened
when viewed from above, weakly protuberant in front; parantennal
fovea semicircular or circular, well defined; nasale raised, as wide as
high. Mandible with small teardrop-shaped or oval pit. Antennae
exceeds tip of hind angle by 1 to 14 segments, reddish brown, ratio of
segments 2:3:4=5:8:10, segment 4 about 1% times maximum
width, erect male hairs of moderate length and dense.

Pronotum wider than long, ratio of width/length=1.07-1.16; punc-
tures at center usually separated by distance equal to little more
than own diameter, equal or subequal to those on front; sides of
pronotum straight behind small anterior curvature, diverging pos-
teriorly; hind angles divergent, with carina usually extending cephalad
of base of hind angles. Genitalia as figured; paramere without apical
blade.

Elytron: 6.10.5 mm (5.2-7.3). 35 spec.

FreMALE.—Antenna exceeds tip of hind angle by about one-half
segment; internal genitalia as figured, bursa not elongate or enlarged
at base of accessory gland and with more than 50 peglike spines.

Elytron: 5.1-6.9 mm. 6 spec.

Typrs.—Lectotype of trapezoideus: Male, labelled with pink circle
“Middle States, incl. N.Y.’”’ (MCZ 2511). First specimen of type
series selected as lectotype by authors in 1962.

DistrRiBuTION.—Ontario to North Carolina, west to Minnesota,
Illinois, and Arkansas.

ARKANSAS: Garland, Washington; June. Connecticut: Litchfield, New
Haven; June, July. Itimnois: Champaign, Knox, McHenry, Putnam, Rock
Island; May, July. Massacuusrerts: Berkshire, Hampshire, Middlesex, Nor-
folk; June, July. Minnesota: Hennepin; July. Mussissippr: George; June.
New Hampsuire: Carroll; August. New Jrersry: Bergen, Gloucester; June.
New York: Erie, Essex, Oswego, Rockland, Suffolk, Tompkins, Westchester;
April, September. NortH Carouina: Macon; June. PENNSYLVANIA: Adams,
Dauphin, Indiana, Philadelphia; May, July (November). West VIRGINIA:
Marion, Pendleton; August. Ontario: Ridgeway; August.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 49

28. Melanotus tenax (Say)

FicursEs 6d, 1la; Puats 1m

Elater tenax Say, 1839, p. 185.

Cratonychus tenaz.—LeConte, 1853, p. 479.

Melanotus tenaz.—Candéze, 1860, p. 360.—Blatchley, 1910, p. 755.—Thomas,
1941, p. 260.—Dietrich, 1945, p. 58.—Fattig, 1951, p. 21.

Mauz.— Body color reddish to dark reddish brown, covered with
white or yellow vestiture, evenly distributed over body.

Head: Front with pair of shallow anterolateral transverse de-
pressions, margin dark reddish brown, not strongly protuberant above
clypeus, thinner in center, evenly rounded or a little flattened an-
teriorly when viewed from above; parantennal fovea smaller than
trapezoideus, well defined, circular; nasale slightly raised, width equal
to 1% times height. Mandible with deep, teardrop-shaped pit.
Antenna exceeds tip of hind angle by one-half segment, reddish
brown, ratio of segments 2:3:4=4:6:9, segment 4 about twice as
long as maximum width; erect male hairs short and dense.

Pronotum as wide as or little wider than long; punctures at center
usually separated by distance equal to own diameter, subequal to
those on front; sides of pronotum straight, subparallel; hind angles
slightly divergent, carina usually extending well cephalad of base of
hind angle. Genitalia as figured; paramere without apical blade.

Elytron: 5.5+0.3 mm (5.2-6.3). 15 spec.

Frema.e.—Antenna not extending to tip of hind angle; internal
genitalia as figured, bursa with about 50 peglike spines, elongate and
enlarged at base of accessory gland, gland short and clavate.

Elytron: 5.0-6.2 mm. 4 spec.

Typrs.— Types of tenaz: ‘“‘Mass.’’ Specimens lost.

DistriputTion.—New York south to Georgia and west to Illinois.

GroraiA: Houston; June. Ituinors: Greene; June. Maryianpb: Prince
Georges; June. Massacuusretts: Middlesex, Nantucket; June. New York:
Albany; June. Norra Carouina: Craven; May. Sout Carouina: Berkeley;
April.

M. trapezoideus and the tenax can be distinguished from other
species of the americanus group by their small size, large pronotal
punctures, and large parantennal fovea. The two are separable by
characters in the key. Also, the pronotum of trapezoideus is usually
flatter and the sides more markedly divergent than in tenar. The
female internal genitalia are dissimilar; the chief differences are the
unusual swelling of the bursa at the base of the accessory gland and
the midlateral origin of the spermathecal duct of tenaz, but not in
trapezoideus. Unlike most species of Melanotus, the male genitalia
of these species appear indistinguishable and are not of value in
separating the two.

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

29. Melanotus pertinax (Say)
Fiaures 61, 10k

Elater pertinax Say, 1839, p. 185.

Cratonychus pertinaz.—LeConte, 1853, p. 479.

Melanotus pertinax.—Candéze, 1860, p. 359.—Blatchley, 1910, p. 755.—Thomas,
1941, p. 260.—Dietrich, 1945, p. 58.—Severin, 1949, p. 17.—Fattig, 1951,
p. 21.

Mauze.— Body color black or dark reddish brown and appendages
lighter reddish brown (light colored teneral specimens also have
appendages paler than body), covered with white vestiture evenly
distributed over body.

Head: Front with pair of shallow anterolateral or transverse
depressions, margin narrow and evenly rounded when viewed from
above, thick and not strongly protuberant above clypeus; parantennal
fovea small but moderately deep circular or semicircular, foveal
tragus seldom developed; nasale flat, width about 2 times height.
Mandible with deep, teardrop-shaped pit. Antenna exceeds tip of
hind angle by about 1 segment, reddish brown, ratio of segments
2:3:4=3:4:5, segment 4 about 1% times as long as maximum width;
erect male hairs short and dense.

Pronotum wider than long, ratio of width/length=1.06—-1.20;
punctures at center usually separated by distance equal to more than
twice own diameter, smaller than those on front, sides of pronotum
rounded and concave at base of hind angles; hind angles divergent,
carina weak, usually not extending cephalad of base of hind angle.
Genitalia as figured; paramere without blade.

Elytron: 6.4+0.4 mm (6.0-7.3). 50 spec.

FremMALeE.—Antennal tip short of pronotum base by 1 segment;
internal genitalia as figured, bursa with sparse, scattered peglike
spines, accessory gland very short and clavate.

Elytron: 7.2+0.4 mm (6.6-8.1). 27 spec.

Typres.—Types of pertinaz: ‘Mass.’ Specimens lost.

DistriButTion.—Maine to North Carolina, west to Wisconsin and
Illinois.

Connecticut: Litchfield, New Haven; May, July. Groraia: Clarke; May.
Inuino1s: Champaign, Knox, McHenry, McLean; May, July. Maine: York;
July. Massacuuserts: Berkshire, Bristol, Hampshire, Middlesex, Suffolk,
Worcester; April, August. Minnesota: Hennepin, St. Louis; May, June. NEw
HampsHIRE: Hillsboro, Strafford; May, July. New York: Columbia, Erie,
Putnam, Tompkins, Washington, Wyoming; May, August. Norra Caro.ina:
Buncombe; June. PENNSYLVANIA: Lycoming; July. SoutH Carouina: Aiken;
June. VERMONT: No data. Wisconsin: Milwaukee; June.

That the appendages are of a different color than the body sets
pertinax apart from other American Melanotus, since no other species
has this coloration.

etn

NO, 3568 MELANOTUS—QUATE AND THOMPSON yl

30. Melanotus infaustus (LeConte)
FicureEs 67, 117,7; PLATE lo

Cratonychus infaustus LeConte, 1853, p. 478.
Melanotus infaustus—Candéze, 1860, p. 357.—Thomas, 1941, p. 260.—Severin,
1949, p. 17.—Fattig, 1951, p. 21.

Mate.—Body color reddish brown, covered with whitish vestiture
evenly distributed over body.

Head: Front with pair of shallow, anterolateral, transverse depres-
sions, margin dark reddish brown, thin and strongly protuberant
above clypeus, flattened anteriorly and straight when viewed from
above; parantennal fovea small, but well defined, arc shaped, and
depth about equal to width of opening; nasale as wide as high, slightly
protuberant. Mandible with deep slitlike pit. Antenna exceeds tip
of hind angle by about 1)% segments, reddish brown, ratio of segments
2:3:4=3:6:9, segment 4 about 1% times as long as maximum width;
erect male hairs short and only moderately dense.

Pronotum about as long as wide, punctures at center usually
separated by distance equal to or less than own diameter, subequal to
those on front, sides of pronotum straight behind anterior curvature,
nearly parallel; hind angles divergent, carina usually extending slightly
cephalad of base of hind angle. Genitalia as figured; paramere with
apical blade.

Elytron: 5.5-7.6 mm. 10 spec.

FremaLe.—Antenna exceeds tip of hind angle by 1 or less segment;
internal genitalia as figured, bursa with large patch of spines anteriorly
and few, scattered ones posteriorly, spines teardrop shaped, spermathe-
cal duct and accessory gland originate close together on bursa.

Elytron: 6.5-7.5 mm. 3 spec.

Types.—Lectotype of infaustus: Male, labelled with orange circle
“Ga.” (MCZ 2518). First specimen of type series selected as lectotype
by authors in 1962.

DistriputTion.—Florida, Georgia, Indiana, Illinois, Kansas, South
Dakota, and Texas.

InpIANA: Tippecanoe; June. Ixiinois: Whiteside; May. Kansas: Douglas,
Reno; June. Sourn Daxora: Brookings, Lake; June, July. Texas: Big Bend
Nat. Pk., Bexar, Kerr; March, June.

M. infaustus appears to be a rare species seldom seen in collections.
It is difficult to identify positively unless the genitalia are dissected.
Undissected specimens may be confused with the more common tenaz,
trapezoideus, and prasinus in the Midwest and South or with lanceatus
in Texas. The slender thorax is helpful in separating it from other
species, except lanceatus, but positive identification requires a study
of the distinctive genitalia.

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

31. Melanotus parallelus Blatchley
Ficures 6h, 11g; PLatr 1n

Melanotus parallelus Blatchley, 1920, p. 45; 1930, p. 44.—Deen and Cuthbert,
1945, p. 193.—Fattig, 1951, p. 22.

Mauzr.—Body color reddish brown, covered with white or yellowish
vestiture, evenly distributed over body.

Head: Front with marked anterior depression, margin dark reddish
brown, strongly protruding in front of nasale by several times own
width, evenly rounded or angulate and flattened in center when viewed
from above; parantennal fovea small, shallow, are shaped or some-
times obsolescent; nasale flat, width equal to slightly more than height.
Mandible with shallow, slit-shaped pit (easily obscured if specimen is
dirty; thus, also keyed out in fissilis group). Antenna exceeds tip of
hind angle by 1 to 14 segments; reddish brown, segment 3 intermediate
in size between 2 and 4, 4 about 1% times as long as maximum width;
erect male hairs short and moderately thick.

Pronotum slender, longer than wide; punctures at center usually
separated by distance equal to own diameter or more, equal or
subequal to those on front, sides of pronotum nearly straight
behind anterior curvature and parallel or slightly convergent
posteriorly, hind angles divergent, carina usually extending just to
base of hind angle, close to and paralleling sides. Genitalia as
figured; paramere with apical blade.

Elytron: 5.9 mm (5.7-6.3). 4 spec.

Frma.LEe.—Antenna ends short of tip of hind angle by about 1 seg-
ment; internal genitalia as figured.

Elytron: 5.9 mm.

Typre.—Holotype of parallelus: Male, Dunedin, Fla., April 5,
1915, W.S. B. (PU).

DistrRiIBUTION.—Southern U.S.

ARKANSAS: No data. Fiorripa: Highlands, Marion, Pinellas; February, April.
Nonrtu Caroutina: Montgomery, Moore; April, May.

The relatively slender pronotum which is as long as or longer than
wide, and the thin, jutting frontal margin are the main features
separating parallelus from other species of the americanus group.
The outline of parallelus pronotum accentuates its narrowness and
makes it appear longer in relation to the width than it really is.
This is another species rarely seen in collections.

32. Melanotus americanus (Herbst)

FiGcurREs 6), lle,f

Elater americanus Herbst, 1806, p. 74.
Cratonychus americanus.—Erichson, 1842, p. 114—LeConte, 1853, p. 479.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 53

Melanotus americanus.—Candéze, 1860, p. 361.—Blatchley, 1910, p. 747.—
Fattig, 1951, p. 21.

Cratonychus angustatus Erichson, 1842, p. 113.—LeConte, 1853, p. 475. [New
synonymy.]

Melanotus angustatus.—Candéze, 1860, p. 345.—Blatchley, 1910, p. 753.—Thomas,
1941, p. 258.

Maxe.—Body color reddish brown, covered with white vestiture
evenly distributed over body. Small species.

Head: Front with pair of shallow, anterolateral, or single, curved
anterior depressions, margin dark reddish brown, protuberant above
clypeus, a little angulate or flat across center when viewed from
above; parantennal fovea large, semicircular, moderately deep, with
weak foveal tragus; nasale usually as wide as high, well developed.
Mandible with deep, teardrop-shaped pit. Antenna exceeds tip of
hind angle by 1%-2 segments; segment 3 short but intermediate
between 2 and 4, ratio of segments 2:3:4=2:3:5, segment 4 about
1% times as long as maximum width; erect male hairs long and
conspicuous.

Pronotum usually wider than long, ratio of width/length =1.00-1.17,
rounded in appearance when viewed from side; punctures small at
center and usually separated by distance equal to twice own diameter;
sides of pronotum rounded anteriorly and almost parallel behind.
Genitalia as figured; paramere enlarged apically, but without apical
blade.

Elytron: 5.4+0.4 mm (4.5-6.3). 50 spec.

Frmate.—Antenna extends to tip of hind angle or exceeds it by
about one-half segment; genitalia as figured, numerous spines in
anterior half, spines long and without flat bases, pouchlike swelling
at base of accessory gland, gland short and expanded beyond base,
diverticulum branching from spermathecal duct near center of duct.

Elytron: 5.7-+0.3 mm (4.8-6.7). 93 spec.

Typrs.—Lectotype of americanus: Male, ‘‘America Sept.”” (ZM).
Selected by Quate, 1960. Seven other males in type series.

Lectotype of angustatus: Female, ‘‘Pennsylvania’” (ZM 17057).
Selected by Quate, 1960; 4 other females in type series.

DistriputTion.—Kastern U.S. west to Indiana and Illinois.

Connecticut: Hartford, Litchfield; June. DELAWARE: Sussex; May.
GroraiA: DeKalb, Fulton, Rabun; May, June. Itimnots: McHenry, Putnam,
Rock Is.; June, July. Inprana: Marion; July. Maryitanp: Anne Arundel,
Baltimore, Frederick, Plummers Is.; June, July. Massacuuserts: Barn-
stable, Hampshire, Middlesex; June, September. NrBRasKa: Cass; May,
July. New Jersey: Bergen, Middlesex, Morris, Union; June, July. New
York: Putnam, Suffolk, Westchester; April, August. Norra Carourna: Bun-
combe, Cherokee, Cumberland, Moore, Montgomery, Pender, Pickens, Swain,
Wake; April, July. Onto: Delaware, Fairfield, Franklin, Greene, Hocking;
May, June. PrnNnsytvania: Allegheny, Dauphin, Northampton, Perry; June,

221-532 675

54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

July. Ruopr Isuanp: Washington; July. SourH CArouina: Oconee; March.
West Virarnia: Greenbrier; July.

Melanotus americanus is one of the smaller species of North American
Melanotus. ‘This feature, with the small third antennal segment and
high rounded pronotum bearing small, scattered punctures, makes
the species more readily recognized than many others.

Examination of the types of angustatus indicates that this species
has been misidentified for many years. The type series is mixed,
but the first is definitely americanus and has been selected as the
lectotype. This specimen has been dissected and the identification
has been confirmed by M. C. Lane.

33. Melanotus cribriventris Blatchley
Figures 6k, 11h

Melanotus cribriventris Blatchley, 1910, p. 747; 1930, p. 33.

Melanotus longicornis Blatchley, 1910, p. 746; 1930, p. 35—Thomas, 1941, p. 258.
[New synonymy.]

Melanotus blatchleyi Leng, 1918, p. 205. [New name for longicornis Blatchley,
not Candéze, 1860. New synonymy.]

Ma.re.—Body color reddish brown, covered with white or yellowish
vestiture, evenly distributed over body.

Head: Front with shallow to marked anterior depression, margin
dark reddish brown, not strongly protuberant above clypeus, angulate
and blunt when viewed from above; parantennal fovea large, deep
and well defined, nasale higher than wide, prominently raised. Man-
dible with deep, rounded pit. Antenna exceeds tip of hind angle by
3 to 3% segments, segments 2 and 3 subequal, flagellar segments
elongate and slender, segment 4 about twice as long as maximum
width; erect male hairs long and shaggy.

Pronotum wider than long; punctures at center separated by
distance equal to less than own diameter, subequal to size of punctures
on front; sides of pronotum straight, divergent posteriorly with
slight anterior curvature; hind angles divergent, with strong carina,
extending cephalad of base. Genitalia as figured; paramere without
apical blade.

Elvtron: 7.2-7.6 mm. 5 spec.

FremMALE.—Antenna exceeds tip of hind angle by 2 segments;
internal genitalia as figured, bursa with about 20 sharp, thornlike
spines, accessory gland long and very slender, spermathecal duct
branched near base.

Elytron: 7.8 mm (7.7-8.1). 8 spec.

Typrs.—Type of longicornis: Male, Orange Co., Ind., May 31,
1904. Not seen by us, but studied by M. C. Lane and determined to
be same as cribriventris.

NO. 3568 MELANOTUS—QUATE AND THOMPSON AD

Holotype of cribriventris: Female, Kosciusko Co., Ind., July 11,
1904, W. S. B. (PU).

Distrinution.—New Jersey west to Kansas.

Inuinots: Rock Is.; no date. Inprana: Allen; June. Kansas: Douglas; no
date. MaryLanp: Baltimore, Plummers Is.; April. New Jersgy: Warren;
June. Onro: Hamilton, Hocking; June. PENNsyLvania: Northampton; June.
TENNESSEE: Morgan; May.

The long antennae and short third antennal segment are distinctive
characteristics of cribriventris, and specimens go through the key with
little difficulty. The shaggy antennae are especially characteristic of
the male and provide an obvious clue to its identification.

Blatchley apparently believed the male and female belonged to two
different species. He first named the male longicornis, but this name
is preoccupied by longicornis Candéze. Leng observed the homonymy
and renamed Blatchley’s species blatchleyr. However, cribriventris is
merely the female of longicornis Blatchley and is therefore available
as a replacement name and has precedence over blatchleyi. Some
identifications will bear this latter name, but the correct one is
eribriventris.

34. Melanotus obscuratus Blatchley

Fiaures 7), 111
Melanotus obscuratus Blatchley, 1927, p. 141; 1930, p. 45.

Matr.—Body color dark reddish brown, vestiture white.

Head: Front with shallow, anterior depression, margin rounded,
not strongly protuberant; parantennal fovea small but distinct, nasale
wider than high, not well developed. Mandible with slitlike pit. An-
tenna exceeds tip of hind angle no more than 1 segment; segments 2
and 3 subequal, ratio of segments 2:3:4=2:3:6, segment 4 about
twice as long as maximum width; erect male hairs short and
inconspicuous.

Pronotum little longer than wide, ratio of width/length =0.9; punc-
tures at center separated by about own diameter, subequal to size of
punctures on front; sides of pronotum nearly parallel, hind angles a
little divergent.

Genitalia as figured; paramere with well-defined apical blade.

Elytron: 6.0-6.4 mm. 2 spec.

Fremate.—Antenna extends just to tip of hind angle; genitalia as
figured, bursa with few spines largely in distal half, accessory gland
originates well before apex, which leads to spermathecal duct.

Elytron: 7.4 mm. 1 spec.

Typr.—Holotype: Male, Ormond, Fla., April 13, 1913, W.S. B.

ePU):

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

DistTRIBUTION.—Southeastern U.S.
Norta Carouina: Brunswick; June. Fiorimpa: Volusia; April.

Three females in the type series belong to morosus as shown by their
long third antennal segment and the internal genitalia characteristic
of that species. However, the holotype and paratype males are dis-
tinct from other North American Melanotus. The female description
above is based on a specimen from North Carolina associated with a
male obscuratus and agreeing with that specimen in external characters.

35. Melanotus beameri Quate, new species

FIGURE 6m

Maue.—Body color light reddish brown, covered with yellowish
vestiture evenly distributed over body.

Head: Front with shallow transverse, anterior depression; margin
protruding in front of nasale by about twice width of rim, angulate
or sometimes rounded and flattened in center when viewed from above;
parantennal fovea small but well defined, crescent shaped; nasale
obsolescent. Mandible with shallow, small, slithke pit. Antenna
reddish brown, exceeds tip of hind angle by 1-2 segments; segment
3 intermediate in size between 2 and 4; erect male hairs dense and of
moderate length.

Pronotum slender, about as wide as long; punctures large and
dense, as large as, or larger than, those on front and separated by less
than own diameter; sides of pronotum with little or no anterior cur-
vature and nearly parallel but a little rounded; hind angles markedly
divergent, carina extends to base of hind angle, close to and paral-
leling side. Genitalia as figured; paramere without apical blade.

Elytron: 5.3+0.3 mm (4.6-6.0). 34 spec.

FrmaLe.—Unknown.

DistRIBUTION.— Texas.

Holotype, male (KU): 65 mi. south of Marathon, Brewster Co., Texas, July
10, 1938, R. H. Beamer.

Paratypes, 33 males (KU, USNM, CNC): same as holotype; near Dugout
Well, Big Bend Nat. Pk., Texas, April 17, 1953, B. J. Adelson; Panther Jct. and
Nine Pt. Draw, Big Bend Nat. Pk., Texas, May 20, 1959, 2,600—4,000 ft., Howden
and Becker; Oak Spring, Big Bend Nat. Pk., May 22, 1959, 4,000 ft., Howden
and Becker.

M. beameri is similar to the following species, but readily separated
from them by the undeveloped nasale, large and coarse pronotal
punctures, and male genitalia.

The species is named in honor of Dr. R. H. Beamer, the late Curator
of the Snow Collection, University of Kansas, who did much work in
the American Southwest and through his diligent efforts added
significantly to our knowledge of insect fauna in that region.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 57

36. Melanotus concisus Knull
Fieurss 7a, 12a
Melanotus concisus Knull, 1959, p. 281.

Ma te (after Knull).—Body color dark brown, legs lighter, vestiture
short.

Head: Frontal margin broadly rounded, projecting over nasale;
parantennal fovea deep. Mandible with deep pit. Antenna exceeds
tip of hind angle by part of 1 segment; segment 3 subequal to 2.

Pronotum little longer than wide; punctures dense, coarse, umbili-
cate on anterior two-thirds, fine and sparse on posterior one-third;
sides of pronotum rounded. Genitalia as figured; paramere with
apical blade.

FEMALE (?).—Frontal margin nearly straight across center, projects
in front of nasale by width of rim; nasale poorly developed, paran-
tennal fovea shallow but definite. Antenna fails to reach tip of
hind angle by about 1 segment. Pronotum wider than long. Geni-
talia as figured; bursa with dense patch of pointed plaquelike spines,
accessory gland originates near apex of bursa.

Elytron: 6.1 mm (5.8-6.3). 5 spec.

Typre.—Holotype: Male, Artesia (Eddy Co.), N. Mex. (Knull
Collection).

DistrrpuTion.—Arizona and New Mexico.

Arizona: Cochise, Santa Cruz; August. New Mexico: Sandoval; August.

The above females agree reasonably well with the male described
by Knull and tentatively we are assigning them to concisus. The
small third antennal segment, large pronotal punctures, small size,
and distribution are the main recognition characters of the species,
aside from the genitalia. Positive identification of the female cannot
be made, however, until associated with male in the field.

37. Melanotus lanceatus Quate, new species

FiauREsS 7c, 126

Maue.—Body color reddish to dark reddish brown, covered with
white vestiture evenly distributed over body.

Head: Front with pair of shallow anterolateral or no depressions,
margin scarcely protruding in front of nasale, angulate and flattened
in center when viewed from above; parantennal fovea moderately
large, deep and circular, or semicircular; nasale protuberant, as wide
as or wider than high. Mandible with slit-shaped or teardrop-
shaped slit. Antenna reddish brown, exceeds tip of hind angle by
2-3 segments; segment 3 intermediate in size between 2 and 4; erect
male hairs short but dense.

58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Pronotum as wide as or less wide than long; punctures at center
separated by about own diameter, smaller than those on front; sides of
pronotum straight behind small anterior curvature, parallel or diver-
gent posteriorly; hind angles divergent, carina extending to base of
hind angle, close to and paralleling side. Genitalia as figured, very
slender; paramere with weakly developed apical blade.

Elytron: 6.5+0.4 mm (5.7-7.2). 64 spec.

FremaLe.—Antenna exceeds tip of hind angle by about one-half
segment; pronotum with sides more rounded and not parallel ; internal
genitalia as figured, bursa with patch of close-set, plaquelike spines,
bases of accessory gland and spermathecal duct at apex of bursa,
gland not clavate.

Elytron: 6.5 mm (6.3-7.0). 12 spec.
DistrisutTion.—Arizona and Texas.

Holotype, male (USNM): Dimmit Co., Texas, May 21, 1934, light trap.
Allotype, female (USNM): same, June 7, 1933, H. J. Reinhard.

Paratypes, 99 males, 12 females (USNM, CAS, CIS, CU, INHS, U. Ariz.,
Texas A and M, CNC): Arizona. Cochise Co.: Huachuca Mts., July 19,
Knull; same, Aug. 8, 9, 1952, 5400 ft., Leech and Green. Santa Cruz Co.:
Canelo, July 10, 1957, G. A. Butler; Nogales, Aug. 12, 1906; Santa Rita Mts.,
July 20, 25, 1959, J. G. Franclemont. Texas. Bell Co.: Salado, April 10.
Bexar Co.: San Antonio, Mar. 17, 1953, B. J. Adelson. Brewster Co.: Alpine,
July 1-15, 1926, O. C. Poling; same, July 11, 1962, L. W. Hepner; Chisos Mts.,
June 10-12, 1908, Mitchell and Eichmann; Santa Elena, Big Bend Nat. Pk.,
May 4, 1959, 2,200 ft., Howden and Becker. Blanco Co.: Cypress Mills,
April 2. Burnet Co.: no locality, June. Cameron Co.: Brownsville, March
20, 1937, T. N. Freeman; same, May 17, 1904, H. 8. Barber; same, May 20, 1937,
May 30, 1932, June 3, 1932. Dimmit Co.: no locality, March 17, 1933, April 3,
30, 1933, June 7, 1933. Duval Co.: San Diego, April 27. Goliad Co.: Goliad,
April 18, E. A. Schwarz. Gonzales Co.: Luling, April 11, 1953, at light, B. J.
Adelson. Hidalgo Co.: Edinburg, no date. Jeff Davis Co.: Davis Mts., June
28, July 8, 1946. Jefferson Co.: Sabina, March, April 1910, F. C. Pratt. Kerr
Co.: Kerrville, April 2-18, 1959, Becker and Howden; same, April 11, 1906, June
1906, F.C. Pratt. Kleberg Co.: Kingsville, no date. LaSalle Co.: Cotulla, April 17,
1906, May 12, 1906, March 27, 1908, F. C. Pratt. Robertson Co.: Hearne,
June 7, 1936, K. L. Maehler. Randall Co.: 15 mi. east of Canyon, June 26, 1956,
R. E. Selander. Sutton Co.: Sonora, April. Terrell Co.: Sanderson, April 27,
1959, Becker and Howden. Val Verde Co.: Del Rio, April 13, 1949, Michener
and Beamer; 13 mi. south of Del Rio, April 10, 1950, Beamer et al.; Devil’s
River, May 5, 1907, E. A. Schwarz. Victoria Co.: Victoria, March 26, J. D.
Mitchell.

M. lanceatus is closely related to longulus. The male genitalia of
the two are similar, but in lanceatus are much more elongate and
slender; the female internal genitalia have the same type of bursal
spines, but the origin of the accessory gland is in a different position
in each. Externally, the smaller pronotal punctures in lanceatus are
the most noticeable difference.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 59

In view of the closeness of the two and their allopatric distribution,
it is debatable if lanceatus should be considered a subspecies of longulus.
We decided against that ranking on the basis that the differences be-
tween lanceatus and longulus are greater than between the two sub-
species of longulus and that the female internal genitalia indicate a
ereater divergence of the two than other characters. It seems to us
highly improbable that the two are capable of interbreeding with
female reproductive organs so dissimilar, and, therefore, they should
not be considered as belonging to the same species.

38. Melanotus hamatus Knull

Ficure 7d
Melanotus hamatus Knull, 1959, p. 280.

Mauer.—Body color reddish brown, covered with whitish vestiture.

Head: Front with small, transverse depression behind margin, mar-
gin angulate, coming to obtuse but noticeable point at midline,
protruding in front of nasale only by width of black rim; nasale not
strongly elevated but defined, wider than long; parantennal fovea
shallow, C-shaped, definite. Mandible with deep, elongate pit. An-
tenna exceeds tip of hind angle by about one-half segment; segment
3 small, much smaller than 4 but a little larger than 2.

Pronotum wider than long; punctures large as on front, compact,
separated by less than own diameter; sides rounded with concavity in
front of hind angles; hind angles small, divergent. Genitalia as
figured; paramere with peculiar winglike flap on venter near center
and without apical blade.

Elytron: 6.4 mm (5.8-6.9). 8 spec.

Frema.te.—Unknown to us.

Typr.—Holotype: Male, Chisos Mtns., Texas (Knull Collection).

DiIsTRIBUTION.—Texas.

Texas: Big Bend Nat. Pk., Jeff Davis; July .

The small size, small third antennal segment, large and dense
pronotal punctures, together with the limited distribution in the
Southwest, will readily identify this species. Of course, dissected
males show the unusual male paramere that is dissimilar to all other
American Melanotus.

38a. Melanotus longulus (LeConte)

Cratonychus longulus LeConte, 1853, p. 473.
Melanotus longulus.—Candéze, 1860, p. 339.

Maue.—Body color reddish brown to black, covered with yellowish
vestiture, evenly distributed over body.

Head: Front with shallow to marked anterior transverse depression,
margin projects in front of nasale by width of rim or less, rounded or

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

angulate and flattened in front when viewed from above, sometimes
depressed in center; parantennal fovea small, deep and crescent shaped,
foveal tragus sometimes well developed; nasale usually developed but
not strongly protuberant, wider than high. Mandible with small
teardrop- or slit-shaped pit. Antenna light to dark reddish brown and
lighter than body in dark specimens, exceeds tip of hind angle by 1
to 1% segments, ratio of segments 2:3:4=3:4:7, flagellum usually
strongly serrate, segment 4 about 1% times as long as maximum width;
erect male hairs short, variable from dense to sparse, but always
evident.

Pronotum as long as or little wider than long; punctures subequal or
clearly as large as those on front, separated by distance equal to less
or more than own diameter.

Frma.te.—Antenna exceeds tip of hind angle by about one-half
segment; sides of pronotum generally rounded; bursa with numerous,
plaquelike spines, spermathecal duct branching at apex of bursa.

39. Melanotus longulus longulus (LeConte)
Ficures 7e, 12c,d

Cratonychus longulus LeConte, 1853, p. 473.

Melanotus longulus—Candéze, 1860, p. 339.—Horn, 1874, p. 23.—Van Dyke,
1932, p. 331.—Fall, 1934, p. 23—Van Dyke, 1942, p. 51—Stone and How-
land, 1944, p. 1.

Reddish brown to black, lighter colored in Southwest. Antenna
flagellum usually moderately serrate. Pronotal punctures subequal
to those on front in Southwest and larger and denser in California,
sides of pronotum nearly straight behind small anterior curvature,
usually slightly convergent posteriorly.

Male genitalia as figured; apical blade sharply angulate with sharp
outer, basal corner, concavity at base of blade well developed in
Arizona and New Mexico and poorly developed in California.

Female internal genitalia as figured, accessory gland longer than
bursa and very slender.

Male elytron: 7.2+0.6 mm (6.5-8.8). 50 spec.

Female elytron: 7.8+0.6 mm (6.3-9.0). 24 spec.

Typres.—Lectotype of longulus: Male, labelled with gold circle
“San Diego, Calif.”” (MCZ 2506). First specimen of type series
selected as lectotype by authors in 1962.

DisTRIBUTION.—Southwestern U.S. south of Tehachapi Mtns.,
California, Great Basin, and Baja California.

Arizona. Cochise Co.: Chiricahua Mts., June 20, 1950; Douglas, Aug.: Hua-
chuca Mts., 1960 m, Aug. 9, 1952; Portal, 2420 m, July 20, 1944; Whetstone
Mts., SE, Aug. 10, 1952. Coconino Co.: Oak Creek Cyn., July 1941; Williams,
July 2, 1953. Graham Co.: Graham Mts., Aug. 1, 1957. Maricopa Co.:

NO. 3568 MELANOTUS—QUATE AND THOMPSON 61

Aguila, Aug. 21, 1927. Mojave Co.: Valentine, Aug. 26, 1952. Pima Co.:
Baboquivari Mts., west side, July 27, 1952; Santa Catalina Mts., July 2, 1954,
July 3, 1955. Pinal Co.: Magna, Aug. 18, 1921; Oracle, July 7, 1950; Tucson,
August 1935. Santa Cruz Co.: Mt. Washington, Nogales, 1829 m, July 13, 1919;
Patagonia, July 1936. Yavapai Co.: Congress Jetn., July; Prescott, July 1, 1919.
Yuma Co.: Yuma, March 7, 1879. Ca.irornia. Imperial Co.: Salton Sea,
Apr. 3, 1927. Inyo Co.: Lone Pine, May 12, 1937; Westgard Pass Plateau,
Apr. 24, 1937; Whitney Portal, July 3, 1953. Kern Co.: Tejon Cnyn., May 12,
1927. Los Angeles Co.: Camp Baldy, June 26, 1958; Claremont, Apr. 19, 1926,
May 138, 1927; Crystal Lake, June 29, 1950; La Canada, July 28, 1948; Lancaster,
May 1895; Los Angeles, Mar. 27, 1879; Mt. Wilson, Apr. 30, 1916, June 13, 1904;
Palmdale, June 9, 1918; Pasadena, no date; Santa Catalina Is., May 1932; South
Gate, Apr. 9, 1952; Tanbark Flat, June 20—July 26, 1952. Orange Co.: Newport
Bay, May 14, 1940. Riverside Co.: Andreas Cnyn., Apr. 3, 1927; Coachella,
May 19, 1927; Idyllwild, June 7, 1940; Indio, June 2, 1918; Murray Cnyn., Mar.
24, 1918; Olanche, May 14, 1917; Palm Springs, Apr. 19, 1924, May 10, 1927,
May 30, 1939. San Benito Co.: Idria, June 29, 1955. San Bernardino Co.:
Forest Home, June 27, 1924; Lytle Creek, June 7, 1928. San Diego Co.: Upland,
Dec. 4, 1952. Ventura Co.: Rincon Beach, July 4, 1923; Saticoy, June 11, 1927;
Ventura, Apr. 25, 1932, Apr. 28, 1932, May 10, 1941; Santa Catalina Is., May
27, 1932. Nervapa. Esmeralda Co.: Goldfield, Aug. 4, 1905. New Mexico.
Hidalgo Co.: Animas Mts., 1670 m, Aug. 15, 1952. Lincoln Co.: Ruidoso, June
26, 1940. Santa Fe Co.: Santa Fe, no date. Trxas. Brewster Co.: Alpine,
July 11, 1938. Uvran. Iron Co.: Cedar City, Coal Cnyn., 1890 m, June 25, 1919.
Juab Co.: Eureka, June 17, 1902, July 26, Aug. 14. Washington Co.: St. George,
May 28, 1935. Basa Cauirornia. 10 mi. south of Catavina, July 29, 1938;
17 mi. south of Ensenada, June 14, 1938.

There is a marked color variation with a geographical basis in
l. longulus. Western populations, particularly in and around Los
Angeles Co., Calif., are colored like J. oregonensis with a black or
nearly black body and reddish-brown appendages. Farther east in
Arizona and New Mexico, the coloration is more like that of lanceatus
and beameri with body and appendage reddish brown. Few specimens
have been seen from the intervening area of southeast California and
we don’t know if a cline of color characters exists. Genitalia and other
structural characters are quite constant, although a more detailed
analysis might reveal differences in some structures as the overall
size and shape of the pronotum.

40. Melanotus longulus oregonensis (LeConte)
Ficures 7f, 12e,f

Cratonychus oregonensis LeConte, 1853, p. 480.

Melanotus oregonensis—Candéze, 1860, p. 373.—Lane, 1952, p. 67.
Melanotus longulus oregonensis—Van Dyke, 1932, p. 331.

Melanotus variolatus LeConte, 1861, p. 377. [New synonymy.]

Melanotus longulus variolatus.—Van Dyke, 1932, p. 331.

Melanotus longulus franciscanus Van Dyke, 1932, p. 332. [New synonymy.]
Melanotus franciscanus.—F all, 1934, p. 24.

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Larger than /. longulus; frontal margin more broadly flattened than
in longulus. Usually black with lighter colored antennae. Antenna
flagellum often strongly serrate. Pronotal punctures large, as large
as those on front and separated by less than own diameter; sides of
pronotum often more strongly rounded than in longulus with marked
concavity at base of hind angle.

Male genitalia as figured; apical blade weakly developed, gently
rounded at base.

Female internal genitalia as figured, accessory gland shorter than
bursa and distinctly clavate.

Male elytron: 9.00.7 mm (7.3-10.3). 50 spec.

Female elytron: 9.1+0.9 mm (7.5-10.9). 50 spec.

Typres.—Lectotype of oregonensis: Male, labelled with dark blue
circle (Oregon) (MCZ 2522). First specimen of type series selected
as lectotype by authors in 1962.

Lectotype of variolatus: Male, labelled with gold circle (Calif.)
(MCZ 2522). Third specimen of type series selected as iectotype by
authors in 1962; first and second specimens are females, and males
are definitive for recognizing the form.

Holotype of franciscanus: Male, hills back of Oakland, ca. May 8,
1910 (CAS 3182).

Distrrpution.—Northwestern U.S., north of Tehachapi Mtns.,
California; overlaps with longulus in north-central Utah.

Catirornia. Alameda Co.: Berkeley, May 9, 1940, June 9, 1949; Castle
Rock Pk., May 25, 1932; Moraga, June 1, 1940; Niles, May 30, 1933; Pleasanton,
June 5, 1932. Butte Co.: Oroville, May 29, 1926. Calaveras Co.: Big Trees,
May 17, 1937; Murphys, May 23, 1936, Jan. 17, 1951. Contra Costa Co.:
Brentwood, May 19, 1949; Giant, May 24, 1932; Mount Diablo, May 30, 1930,
Apr. 20, 1942, May 3, 1957; Walnut Creek, May 30, 1913, June 1930. El Dorado
Co.: Camino, June 21-29, 1948; Pollock Pines, July 10, 1948. Fresno Co.:
Clovis, Apr. 21, 19381; Dalton Creek, Apr. 18, 1920; Mendota, Apr. 19, 1956.
Humboldt Co.: Blocksburg, June 3, 1935; Fort Seward, May 22, 1935. Kern
Co.: Bakersfield, May 5, 1931. Lake Co.: Anderson Springs, May 17, 1952.
Lassen Co.: Doyle, May 20, 1934. Madera Co.: Bass Lake, July 24, 1934;
Coarsegold, May 12, 1942; Madera, May 2, 1947; Northfork, Feb. 29—Mar.
30, 1920. Marin Co.: Fairfax, May 1922; Mill Valley, May 9, 1949. Mari-
posa Co.: Miami Ranger Sta., July 29, 1946. Merced Co.: Los Banos, Mar. 16,
1946; Panoche Hills, Apr. 23, 1921; Yosemite, May 29, 1937. Modoc Co.:
Cedar Pass, July 29, 1946. Mono Co.: Leavitt Meadows, June 26, 1937. Mon-
terey Co.: Carmel, Jan. 7, 1908. Napa Co.: Monticello, May 30, 1930. Nevada
Co.: Greenhorn, August 28, 1954; Tahoe, June 3-July 8, 1915. Placer Co.:
Auburn, May 1939; Newcastle, Apr. 23, 1949; Penryn, July 15, 1939. Plumas Co.:
Keddie, June 12, 1941. Sacramento Co.: Sacramento, Apr. 23, 1922. San Benito
Co.: June 24, 1933. San Joaquin Co.: Lodi, Apr. 23, 1931; Stockton, May 12,
1933; Tracy, May 4, 1933. San Mateo Co.: Halfmoon Bay, Mar. 16, 1952. Santa
Clara Co.: Morgan Hill, May 17, 1922; Mtn. View, May 18, 1941; Palo Alto,
May 7, 1920; San Jose, Mar. 2, 1941. Santa Cruz Co.: Santa Cruz Mtns., no
date. Shasta Co.: Hat Creek, June 25, 1949, July 25, 1953; Redlands, June 6,

NO. 3568 MELANOTUS—QUATE AND THOMPSON 63

1953. Siskiyou Co.: Yreka, May 10, 1932. Solano Co.: Green Valley Falls,
Apr. 26, 1941. Sonoma Co.: Santa Rosa, April 1942; Sobre Vista, May 24, 1910.
Stanislaus Co.: Westley, June 8, 1948. Tehama Co.: Red Bluff, Apr. 14, 1928,
May 11, 1949; Vina, May 11, 1920; Western, Oct. 30, 1920. Trinity Co.: Carr-
ville, May 17, 1934, May 30, 1934. Tulare Co.: Kaweah; Sequoia Nat. Pk.,
June 13, 1929, May 25, 1930; Visalia; Wood Lake, Mar. 28—May 3, 1947.
Tuolumne Co.: Hardin Flat, May 29, 1955. Yolo Co.: Winters, Apr. 23, 1950.
Yuba Co.: Marysville, June 5, 1933. Ipano. Bannock Co.: Lava Hot Springs.
Canyon Co.: Parma, May 18, 1930. Lincoln Co.: Shoshone, June 10, 1938.
Montana. Lewis and Clark Co.: Wolf Creek, July 4, 1938. Nervapa. No
other data. OrgGcon. Baker Co.: Richland, June 14, 1940. Benton Co.: Cor-
vallis, July 2, 1945. Douglas Co.: Drain, May 19, 1914. Harney Co.: June 22,
1912. Jackson Co.: McLeod St. Pk., May 22, 1960; Medford, June 12, 1915,
May 14, 1954; Talent, May 4, 1954. Lake Co.: Lakeview, May 27, 1957.
Yamhill Co.: June 1913, May 1934. Utan. Cache Co.: June 26, 1954; Lewis-
ton, May; Bear River, Logan, 1925. Juab Co.: Eureka, July 2 (with l. longulus).
Salt Lake Co.: Apr. 14, 1909; Ft. Douglas, June 7, 1926. Tooele Co.: Stockton,
June 22, 1917. Utah Co.: Dividend, June 1921; Provo, June 2, 1939; Provo
Cnyn., May 21, 1913. Weber Co.: Roy, July 14, 1957. Wasuineton. Kittitas
Co.: Yakima River, 8 mi. south of Ellensburg, July 22, 1959. Walla Walla
Co.: Walla Walla, June 1936. Whitman Co.: Pullman, May 13, 1930; Wawawai,
May 13, 1944. British CotumsBia. Oliver, May 22, 1924; Vernon, May 1,
1924; Victoria, June.

The subspecies longulus and oregonensis are closely related forms,
allopatric in distribution (fig. 3). Longulus differs from oregonensis
rather constantly in having a smaller average size, being lighter in
color, the nasale better developed, the fovea a little deeper, the
antennae less sharply serrate, and the flagellar segments smaller.
These features are subject to some variation and a small percentage of
specimens possesses characters of the other subspecies. The genitalic
characters, on the other hand, seem entirely constant within the ranges
of the respective subspecies. Specimens have not been seen which are
intermediate between the two subspecies in genitalic characters. A
specimen from northern Utah is intermediate in size and color but
oregonensis on genitalic characters. Perhaps these forms have reached
full species status and there is no interbreeding between populations.
This is unproven, however, and we have seen few specimens from the
borders of the two forms’ area. In our opinion, the close relationship
and allopatric distribution is best shown by the nomenclatorial rank
of subspecies.

Van Dyke (1932) recognized the affinity of longulus and oregonensis
and was the first to group them as subspecies of one species. At the
same time he recognized the third subspecies, franciscanus. He
presents a scheme, that our observations do not support, in which the
smaller and lighter longulus of southern California gets larger and
darker in populations to the north in the Sierra Nevada and the Great
Basin. The lowland populations in central California supposedly got

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

still larger and darker and constituted the subspecies franciscanus.
This suggests a north-south cline from southern California and east-
west cline through central California. Our studies show a sharp break
between the populations along a line in the Tehachapi Mountains,
California, and if there is a north-south cline, it is abrupt and not
eradual as outlined by Van Dyke. On the average, specimens from
the mountains are smaller and those in the lowlands larger. (Although
one of the smallest specimens of oregonensis in our material is from
Pleasanton, Alameda Co., Calif.) This is most likely a phenotypic
expression of a more vigorous climate, shorter summer season, and
perhaps less abundant food in the mountains, and we think has nothing
to do with the genotypic characters which distinguish the two sub-
species. The form franciscanus is regarded as merely a larger and
darker phenotype of oregonensis that enjoys a more favorable environ-
ment but does not warrant nomenclatorial recognition.

The species variolatus has been synonymized with longulus by Van
Dyke (1932) on the grounds that it was based on individual variants
and in this we concur.

41. Melanotus gradatus LeConte

FiGuRE 7g; PLATE 1p

Melanotus gradatus LeConte, 1866, p. 390.—Blatchley, 1910, p. 753.—Thomas,
1941, p. 260.

Mate.—Body color reddish brown, covered with white or yellowish
vestiture that grows denser and sometimes nearly velvety posteriorly
on pronotum.

Head: Front with pair of shallow anterolateral depressions; margin
not strongly protuberant above nasale, evenly rounded or flattened
in front when viewed from above; parantennal fovea moderately large,
deep, semicircular, foveal tragus small; nasale raised slightly, width
from 1% to 1% times height. Mandible with large teardrop-shaped
pit. Antenna reddish brown, exceeds tip of hind angle by one-half
segment or less, segment 3 nearly as large as 4, 4 about 1% times as
long as maximum width; erect male hairs very short, fine, rather
sparse, and easily overlooked.

Pronotum about as wide as long; punctures on anterior two-thirds
as large as those on front, and separated by distance equal to less
than own diameter, those on posterior one-third grow much smaller
and denser, often giving surface granulose appearance; sides of pro-
notum vary from straight and diverging posteriorly to curved with
slight concavity at base of hind angle; hind angles divergent, carina
usually extending cephalad of base of hind angle and weakened
anteriorly. Genitalia as figured; paramere with apical blade.

Elytron: 7.9-8.2 mm. 3 spec.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 65

Fremate.—Antenna extends just to tip of hind angle or short of tip
by one-half segment; internal genitalia as figured, bursa with 2 patches
of tacklike spines, accessory gland slender and shorter than bursa,
spermathecal duct branches near its base.

Elytron: 8.5-9.5 mm. 5 spec.

Typr.—Holotype of gradatus: Male, Maryland. Only specimen
in type series, not seen by us.

Distripution.—Maryland west to Kansas and Arkansas.

ARKANSAS: Lawrence, Washington; April, June. I uiNors: Gallatin, Pope,
Pulaski, St. Clair; May, June. Inprana: Clark; June. Kansas: Douglas;
June. Onto: Hamilton; June.

The fine vestiture and punctures on the posterior part of the pro-
notum, the short antennae, and moderately large size readily dis-
tinguish M. gradatus from other North American Melanotus.

42. Melanotus insipiens (Say)
Figures 7h, 12g
Elater insipiens Say, 1825, p. 267.—1839, p. 184.
Cratonychus insipiens.—LeConte, 1853, p. 480.
Melanotus insipiens.—Candeéze, 1860, p. 361.—Fattig, 1951, p. 21.
Cratonychus tenellus Erichson, 1842, p. 114——LeConte, 1853, p. 480.
Melanotus tenellus.—Candéze, 1860, p. 362.—Fattig, 1951, p. 22.

Mauzr.—Very small, light to dark reddish brown, species with
white or yellowish vestiture.

Head: Front with shallow, narrow anterior depression or lacking
entirely; margin dark reddish brown, not strongly protuberant over
clypeus, evenly rounded when viewed from above, slightly upturned;
parantennal fovea deep, well defined, semicircular; nasale wider than
high. Mandible with deep, rounded, or elongate pit. Antenna
exceeds tip of hind angle by 2 to 2% segments, segment 3 intermediate
in size between 2 and 4, 4 about 1% times as long as maximum width;
erect male hairs moderately long and sparse.

Pronotum a little wider than long; punctures at center usually
separated by distance equal to about own diameter, slightly smaller
than punctures on front; sides of pronotum straight behind small
anterior curvature and divergent posteriorly; hind angles divergent,
carina extending cephalad of angle base, close to and paralleling sides.

Last visible sternite of abdomen with punctures on posterior half
round and considerably larger than on rest of abdominal venter.
Genitalia as figured; paramere with apical blade.

Elytron: 3.5+0.2 mm (3.1-4.0). 50 spec.

Female. Antenna exceeds tip of hind angle by one-half segment;
internal genitalia as figured, bursa with few tacklike spines, accessory
gland and spermathecal duct originate close together at apex of bursa,
duct diverticulum very short.

66 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Elytron: 3.60.2 mm (3.0-4.3). 50 spec.

Typrs.—Type of insipiens: Specimens lost.

Lectotype of tenellus: Female, ‘Amer. Sept.’”? (ZM). Lectotype
selected by Quate in 1960.

Distrisution.—New Jersey to Florida, west to Texas.

ALABAMA: Mobile; June. ARKANSAS: Hempstead; June. FLoripa: Columbia,
Pinellas, Osceola, Volusia, Wakulla; April, May. Grorera: Charlton, Houston;
May. New Jersey: Burlington, July. Norra Carouina: Bladen, Johnston,
Lenoir, Moore, Pender, Richmond, Sampson, Wake, Wayne; May, July. SoutH
CAROLINA: Florence; January. TExas: Harrison; May.

The small size is usually sufficient to separate insipiens from other
North American Melanotus. It might be confused with americanus,
but in that species the pronotal punctures are much smaller and
sparser. MM. angustatus and prasinus are also small species of the
americanus group, but they are larger than insipiens and have con-
spicuously different genitalia.

43. Melanotus leonardi (LeConte)

Fiaurss 7k, 12h, 7

Cratonychus Leonardi LeConte, 1853, p. 475.
Melanotus leonardi.—Candéze, 1860, p. 349.—Thomas, 1941, p. 259.—Dietrich,
1945, p. 56.—Fattig, 1951, p. 20.

Mauz.—Head, anterior border of pronotum, elytron, abdomen, and
venter black, antenna and legs dark reddish brown and pronotum
orange colored.

Head: Front usually with marked, anterior depression; margin
scarcely protuberant in front of nasale, rounded or broadly flattened
across center when viewed from above; nasale flat, parantennal fovea
absent or very shallowly indicated. Mandible without pit. Antenna
extends to or beyond tip of hind angle by 1 segment, segment 3 small,
little larger than 2, but much smaller than 4; erect male hairs short
but dense enough to be seen easily.

Pronotum wider than long, ratio of width/length=1.1-1.2; punctures
equal or subequal to those on front, central ones separated by distance
equal to about own diameter; sides of pronotum evenly rounded with
marked concavity at base of hind angles; hind angles divergent; carina
weak, barely extending to hind angle base. Genitalia as figured, apical
blade very small.

Elytron: 6.6 mm (5.9-7.3). 16 spec.

Frmaue.—Antenna extends about to tip of hind angle, segment 3
intermediate in size between 2 and 4; internal genitalia as figured,
bursa with small cluster of spikelike spines, accessory gland short and
thicker than duct, spermathecal duct branches at about basal one-

third.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 67

Elytron: 7.1 mm (6.3-7.8). 20 spec.

Typres.—Lectotype of leonardi: Male, labelled with pink circle ‘‘prob.
Pa.”” (MCZ). Second specimen of type series selected as lectotype by
authors in 1962; second, a male, chosen in preference to first, a female,
because males more accurately identified.

DistriputTion.—Quebec to North Carolina, west to Texas.

Connecticut: Litchfield; June. Inprana: Marion; June. Maine: Hancock,
Oxford, Washington; July. Massacuussrts: Suffolk, Worcester; June. New
HampsuireE: Cheshire, Strafford; no dates. New York: Essex, Jefferson; April,
June. Norra Carouina: Buncombe; June. Trxas: Bexar; June. VERMONT:

No data. Onrarro: Mustota Dist., Orrville; June. QumsBEc: Duparquet, Mt.
St. Hilaire, Perkins Mills; June, July.

44. Melanotus taenicollis (LeConte)
Fiaurss 71, 12]

Cratonychus taenicollis LeConte, 1853, p. 475.
Melanotus taenicollis: Candéze, 1860, p. 348.—Thomas 1941, p. 259.—Dietrich,
1945, p. 56.

Maue.—Black and orange or reddish brown, body largely black
with orange on margins and ventral part of pronotum, on anterior
border of elytra, on legs, and over entire, or on margins of, abdominal
venter, antenna usually dark reddish brown but lobes of flagellar
segments sometimes lighter colored.

Head: Front with anterior depression; margin scarcely protuberant
in front of nasale, coming to point in center when viewed from above;
nasale flat, parantennal fovea absent. Mandible without pit. An-
tenna short of tip of hind angle apex by about 1 segment; segment 3
small, little larger than 2, but much smaller than 4, flagellar segments
broad; erect male hairs may be sparse but easily seen.

Prontotum wider than long, ratio of width/length=1.15-1.26; punc-
tures subequal to those on front, central ones separated by about own
diameter; sides of pronotum evenly rounded with marked concavity
at base of hind angles; hind angles parallel or divergent, carina weak,
extending just to hind angle base. Genitalia as figured; paramere
with apical blade.

Elytron: 6.5+0.3 mm (6.0-6.9). 14 spec.

Frmauy.—Similar to male but less variation in body color; antenna
short of tip of hind angle by about 2 segments, segment 3 inter-
mediate in size between 2 and 4; internal genitalia as figured, bursa
with cluster of spinelike spines, smaller and more numerous than in
leonardi.

Elytron: 6.70.4 mm (6.1-7.5). 16 spec.

Typres.—Lectotype of taenicollis: Male, labelled with pink circle
“Middle States; Pa.” (MOZ 2512). First specimen of type series

68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

selected as lectotype by authors in 1962; second specimen probably
not a type, since only one mentioned in original description.

Distripution.—Kastern U.S.

New Jersey: Essex; no dates. New York: Dutchess; May, June. PENN-
SYLVANIA: Delaware; June.

M. taenicollis and leonardi are the only two North American Melano-
tus with a contrasting, bicolored pattern and are easily distinguished
from other species because of this. The black center of the pronotum
and orange on the abdominal venter of taenicollis, as well as the
genitalic structures, readily separate it from leonard.

The cribricollis Group

Pronotal punctures large and dense; male paramere short, ends far
basad of aedeagus apex.

45. Melanctus cribricollis Candéze

Fiaures 7), 12l,m; PLATE 11
Melanotus cribricollis Candéze, 1860, p. 358.

Matr.—Large, dark reddish species with sparse, short yellowish
vestiture.

Head: Front with marked, anteromedian depression; margin scarcely
protruding in front of nasale, angulate and concave in center when
viewed from above, also sunken or impressed in center; nasale flat,
short, and much wider than high; parantennal fovea shallow (1 spec.
with deep fovea), circular, or semicircular. Mandible with shallow,
elongate pit near lower border. Antenna short, fails to reach tip of
hind angle by 4—5 segments, segment 3 intermediate in size and shape
between 2 and 4; flagellar segments broad and nearly rectangular,
lower angles fulvous; only a few short erect male hairs on base of
lower margin, easily overlooked and difficult to distinguish males
from females.

Pronotum wider than long, punctures very large and dense, margins
nearly contiguous; sides of pronotum straight behind strong anterior
curvature, infrequently with concavity at base of hind angle; hind
angles subparallel, each with strong carina extending to or little
cephalad of hind angle base and sometimes with weak, shorter median
carina about two-thirds length of larger one. Genitalia as figured;
paramere shorter than aedeagus, ending in broad, apical blade.

Elytron: 7.9 mm (6.5-8.5). 12 spec.

FemMALe.—Internal genitalia as figured, accessory gland large and
spherical, bursa with large, dense patch of spikelike spines.

Elytron: 7.9--0.5 mm (7.3-9.0). 18 spec.

Tyrr.—Holotype of cribricollis: Female, “Amer. Bor.” (BMN).
Only specimen in type series.

NO. 3568 MELANOTUS—QUATE AND THOMPSON 69

DistTRIBUTION.—Southwestern U.S., Baja California.

Arizona: Cochise, Maricopa, Pima, Pinal; April, September. Ca.irornia:
San Diego; April. Basa Cauirornia: 10 mi. southwest of Comondu, 25 mi.
west of LaPaz, 8 mi. northwest of Los Angeles Bay, 22 mi. northwest of Pen-
jamo, St. Bartlme Bay, 10 mi. south of San Jose del Cabo; June, September.

M. cribricollis and chiricahuae are readily separable from other North
American Melanotus by the large pronotal punctures, shallow mandib-
ular pit, short antennae, and peculiar male genitalia. Their divergence
from other North American species and distribution in the south-
western U.S. suggests they are of Neotropical origin.

M. cribricollis differs from chiricahuae in its shorter antennae, shape
of the frontal margin, more rectangular shape of the flagellar segments,
better developed parantennal fovea, and different shape of the male
genitalia.

46. Melanotus chiricahuae Knull
Figures 71, 12k

Melanotus chiricahuae Knull, 1962, p. 34.

Ma.e.—Large, dark reddish-brown species with sparse, yellowish
vestiture.

Head: Front with marked anterior depression; margin protruding
in front of nasale by about twice width of rim, rounded when viewed
from above; nasale flat, parantennal fovea very shallow, faintly
crescent shaped. Mandible with shallow, elongate pit near lower
border. Antenna exceeds tip of hind angle by about % seg-
ment, segment 3 intermediate in size between 2 and 4, flagellar
segments triangular; erect male hairs short, usually dense but may
be scarcely visible.

Pronotum wider than long, punctures very large and dense, sep-
arated by less than own diameter; sides of pronotum straight behind
anterior curvature and usually subparallel but may be divergent
posteriorly; hind angles divergent, each with a strong carina extending
cephalad of hind angle base and incurved anteriorly and a weak,
shorter, median carina about ™% length of larger one. Geni-
talia as figured; paramere much shorter than aedeagus, ending
in unusual recurved apex.

Elytron: 8.6+0.5 mm (7.6-9.5). 21 spec.

FrEMALE.—Similar to male; antenna shorter, extends to tip of hind
angle or short of tip by 1 segment; genitalia as figured, bursa with
thick patch of simple spines, accessory gland short, originates well
before apex and origin of spermathecal duct.

Elytron: 9.2+0.5 mm (8.5-10.5). 16 spec.

DistTRIBUTION.—Southwestern U.S.

Arizona: Cochise, Santa Cruz; June-August. New Mexico: Hidalgo; August.

oie Gr 6

70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

The large pronotal punctures of this species are distinctive, although
it might be confused with similis in the same area. The punctures of
chiricahuae are larger than in similis, the antennae are shorter, and
it has a weak, shallow mandibular pit. The male and female genitalia
are easily distinguished from other species.

Species Incertae
Melanotus abdominalis (Erichson)
Cratonychus abdominalis Erichson, 1842, p. 104.—LeConte, 1853, p. 481.
Melanotus abdominalis.—Candéze, 1860, p. 352.

TypE.—Holotype: Female, ‘Amer. Sept.” (ZM, 17036). Only
specimen in type series.

This species has subequal second and third antennal segments, a
jutting frontal margin, fine dense pronotal punctures, and a long,
slender, heavily spied female bursa. It is unlike any species which
we have encountered in North America and is unfamiliar to us. We
suspect that it is not a North American species and the type has been
labelled in error.

Melanotus cinereus (Weber)
Elater cinereus Weber, 1801, p. 77.

This species is usually listed as a synonym of similis (=fissilis) or
communis, but the types apparently are lost, and we are unable to
apply the name to a known species.

Melanotus despectus Candéze
Melanotus despectus Candéze, 1860, p. 343.

We have not seen types of this species, but Mr. Lane (in litt.)
states that he saw specimens at the Deutschen Entomologischen
Institut, Berlin, which might be the types. According to Lane,
these are the same as ignobilis and if those specimens can be shown to
be types, despectus would fall as a synonym of ignobilis.

Melanotus effetus Candéze
Melanotus effetus Candéze, 1860, p. 355.

This may be only a small form of communis according to Lane
(in litt.), who found specimens at the Zoologische Museum der
Humboldt-Universitat that might be the types.

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1939. I Melanotini della fauna italiana: Spheniscocomus Schw. e Melanotus
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1833. Catalogue des Coléoptéres de la collecion . . ., ed. 2, 176 pp.
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1854. Insects of New York. Vol. 5 of pt. 5 (Agriculture) in Natural history
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Ericuson, W. F.

1842. Ueber die Elateren mit Kammférmig gezihnten Krallen. Germar
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1934. Journ. New York Ent. Soc., vol. 42, p. 24.
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1926. Journ. Econ. Ent., vol. 19, p. 502.
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1817. In Schoenherr, Synonymia insectorum, oder: Versuch einer Syno-
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1806. Natursystem aller bekannten in- und auslandischen Insecten,. .
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1874a. Trans. Amer. Phil. Soc., n.s., vol. 10, p. 473.
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1955. Sur les Elatéroides paléaretiques. Bull. Ann. Soc. Roy. Ent. Belgi-
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~3.

Krirsy, W.
1837. Insects. Pt. 4 in Richardson, Fauna Boreali-Americana, xxxix+ 325
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Knut, J. N.

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NO. 3568 MELANOTUS—QUATE AND THOMPSON Ves

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74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Tuomas, C. A.
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1932. Miscellaneous studies in the Elateridae and related families of
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VAN ZWALUWENBURG, R. H.
1921. Melanotus hyslopi, new species (Coleoptera). Proc. Ent. Soe.
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1922. External anatomy of the elaterid genus Melanotus (Coleoptera),
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WEBER, FRIEDRICH
1801. Observationes entomologicae, continentes novorum quae condidit
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WEstTwoop, J. O.
1840. An introduction to the modern classification of insects ..., vol. 2.

No. 3568 MELANOTUS—QUATE AND THOMPSON 75

Ficure 4.—Male genitalia of Melanotus s
castanipes; 7, communis; k, indistinctus;
(Scale lines=0.5 mm.)

pecies: a-d, similis; e, spadix; f-h, decumanus; i,
L, dietrichi; m, verberans; n, miscellus; 0, emissus.

76

PROCEEDINGS OF THE NATIONAL MUSEUM

Ficure 5.—Male genitalia of Melanotus species: a,
clandestinus; g, ignobilis; h, morosus; 1, depressus;
sagittarius; n, corticinus. (Scale lines=0.5 mm.)

VOL. 121

lanei; b, c, pilosus; d, é, opacicollis; f,
j, cribulosus; k, piceatus; 1, hyslopi; m,

77

MELANOTUS—QUATE AND THOMPSON

NO. 3568

d,

1, infaustus; j, americanus; k, cribriventris; 1,

pecies: a, prasinus; b, testaceus; c, trapezoideus;
0.5 mm.)

3h, parallelus;

Ficure 6.—Male genitalia of Melanotus s

difficilis

pertinax; m, beameri. % (Scale lines

tenax; e, f, macer; g,

78

PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

4

1G

te

'

1

'

’

'
1
\

Ficure 7.—Male genitalia of Melanotus species: a, concisus; b, obscuratus; c, lanceatus; d,
hamatus; e, longulus longulus; f, longulus oregonensis; g, gradatus; h, insipiens; i, chiricahuae;

j, cribricollis; k, leonardi; lL, taenicollis. (Scale lines=0.5 mm.)

NO. 3568 MELANOTUS—QUATE AND THOMPSON 79

Ficure 8.—Female genitalia of Melanotus s
labelled; b, similis, “normal”; c, similis, “
(Scale lines= 1.0 mm and 0.1 mm.)

Ae. wan,
o*Vae- 2

o-

Ss less?

WOE
Hy

a LMM RAE
\ Grebe
YEN
somone B.\
pecies: a, hypothetical species with structures
spiny”; d, decumanus; e, spadix; f, castanipes.

80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

miscellus; d,e, dietrichi;
(Scale lines=

Ficure 9.—Female genitalia of Melanotus species: a,b, communis; c,
fig, indistinctus; h, lanei; 1, verberans; j, opacicollis; k, emissus (2); 1, pilosus.
1.0 and 0.5 mm.)

NO. 3568 MELANOTUS—QUATE AND THOMPSON Sl

rn

ne
1}
ty
dn.

Ses

Ficure 10.—Female genitalia of Melanotus species: a, ignobilis; b, depressus; c, sagittarius;
a, cribulosus; e, morosus; f, hyslopi; g,h, prasinus; i, corticinus; J, piceatus; k, pertinax;
I, macer. (Scale lines=1.0 and 0.05 mm.)

121

VoL.

PROCEEDINGS OF THE NATIONAL MUSEUM

82

,d, testaceus;

.
J

a, tenax; b, trapexoideus

infaustus (?); k, gradatus;

Ficure 11.—Female genitalia of Melanotus species

|
|
(
'
|

1, obscuratus. |

1J>

g, parallelus; h, cribriventris;

.
?

americanus

ef,

0.5 and 0.05 mm.)

(Scale lines

NO. 3568 MELANOTUS—QUATE AND THOMPSON 83

Ficure 12.—Female genitalia of Melanotus species: a, concisus (2); b, lanceatus; c,d, longulus
longulus; e,f, longulus oregonensis; g, insipiens; h,i, leonardi; 7, taenicollis; k, chiricahuae;
Im ,cribrjcollis. (Scale lines=0.5 and 0.05 mm.)

U.S. GOVERNMENT PRINTING OFFICE: 1967

PROC. U.S. NAT. MUS. VOL. 121 QUATE AND THOMPSON—PLATE 1

Heads of Melanotus species: a, similis; b, decuman us; C, Castanipes; d, communis; e, verberans;
f, lanet; g, pilosus; h, emissus; i, opacicollis; J, depressus: k, sagittarius; 1, cribricollis; m
tenax; n, parallelus; 0, infaustus; p, gradatus.

’

Proceedings of
the United States

National Museum

SMITHSONIAN INSTITUTION . WASHINGTON, D.C.

Volume 121 1967 Number 3569

SOLDIER FLY LARVAE IN AMERICA NORTH OF MEXICO!
By Max W. McFappen ?

The Stratiomyidae or soldier flies are represented in America north
of Mexico by approximately 237 species distributed through 37
genera. Prior to this study, larvae have been described for only 21
species representing 15 genera. In addition to the lack of adequate
descriptions and keys, classification has seldom been attempted and
a phylogenetic treatment of the larvae has never been presented.

The present study has been undertaken with several goals in mind:
(1) to rear and describe as many species as possible; (2) to redescribe
all previously described larvae of North American species; and (3),
on the basis of larval characters, to attempt to define various taxo-
nomic units and show phylogenetic relationships within the family
and between it and other closely related familes.

Any attempt to establish subfamilial and generic limits must be
regarded as tentative. This is especially true in the present study
since larvae of so many species of Stratiomyidae remain unknown.
Undoubtably, as more species are reared, changes will have to be
made in keys and definitions of taxa.

The keys have been prepared chiefly for identification of last
instar larvae. If earlier instars are known, they either have been

' Modified from a Ph. D. dissertation submitted to the University of Alberta,
Edmonton, Canada.

? Entomology Research Division, U.S. Dept. Agriculture, Tobacco Insects
Investigations, P.O. Box 1011, Oxford, N.C. 27565.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

included in the key or a separate description has been given—or
both. Because pupation takes place within the last larval skin,
the puparium can be identified with the keys to the larvae.

I wish to extend sincere appreciation to Dr. George E. Ball of
the University of Alberta, Edmonton, for his excellent direction of
the study; to Dr. Maurice T. James of Washington State University,
Pullman, for his constant help and advice on many problems; to
Dr. Willis W. Wirth of the U.S. Department of Agriculture, Wash-
ington, D.C., for his advice and for reviewing the manuscript; and
to the University of Alberta and the National Research Council
of Canada for their financial assistance.

I also wish to acknowledge with thanks material loaned from the
following entomologists and institutions (abbreviations that follow
are used throughout this text):

American Museum of Natural History, New York, N.Y. (P. Wygodzinsky) A MNH
California Academy of Sciences, San Francisco, Calif. (P. Arnaud, Jr.)—CAS
Canada Department of Agriculture, Belleville, Ont. (H. G. James)—CDA
Canadian National Museum, Ottawa, Ont. (G. E. Shewell)—CNC

Clemson College, Clemson, 8. C. (R. C. Fox)—CC

Colorado State University, Fort Collins, Colo. (T. O. Thatcher)—CSU

Cornell University, Ithaca, N.Y. (C. O. Berg and H. Dietrich) —CU

Deutches Entomologisches Institut, Berlin, Germany (W. Hennig)—DEIT
Illinois Natural History Survey Collection, Urbana, Ill. (L. K. Gloyd)—INHM
Louisiana State University, University Park, La. (H. V. Daly)—LSU

M. W. McFadden Collection—M WM

North Carolina State College, Raleigh, N.C. (R. L. Rabb)—NCSC

Ohio State University, Columbus, Ohio (J. G. Rozen)—-OSU

Purdue University, Lafayette, Ind. (L. Chandler)—PU

Rockefeller Foundation, Mexico, D.F., Mexico (J. L. Carrillo S.)\—RF
University of California, Berkely, Calif. (P. Ashlock and R. L. Usinger)—-UCB
University of Delaware, Newark, Del. (P. P. Burbutis)—-UD

University of Kansas, Lawrence, Kans. (W. J. Hanson)—UK

University of Maine, Orono, Maine (G. W. Simpson) —UM

University of Massachusetts, Amherst, Mass. (M. E. Smith)—UMASS
University of Michigan, Ann Arbor, Mich. (T. E. Moore)—U MICH

University of Missouri, Columbia, Mo. (W. R. Enns)—UMO

United States National Museum, Washington, D.C. (W. W. Wirth)—USNM
Washington State University, Pullman, Wash. (M. T. James)—WSU

Methods

A suitable technique for removing larvae from various substrates
has been developed (McFadden, 1961). Specimens thus obtained
for this study were divided into two groups: one to be reared, the
other to be preserved.

Larvae were killed and preserved in 70 percent alcohol without
additional treatment. In the case of large specimens, the alcohol
was changed after a month’s lapse.

NO. 3669 SOLDIER FLY LARVAE—McFADDEN 3

Rearing was simplified by collecting in the spring, when larvae
were usually in the last instar, about to pupate, or had already
pupated. Larvae and pupae held for rearing were placed in 5-dram
vials fitted with cotton stoppers. A small amount of the substrate
in which the animal was found was placed in the vial before adding
the specimen. Distilled water was added occasionally to maintain
moist conditions. Larvae and pupae were always reared individually
so that emerging adults or parasites could be associated with the
respective larval or pupal skin. The adults were then identified and
associated with the larvae that had not been reared, and thus the
identity of these larvae was determined. This is by no means a
foolproof system for identification, but in most instances it can be
relied upon because of the gregarious nature of the larvae.

In observing preserved specimens, I frequently found that many
morphological characters that are used in distinguishing the various
taxa were obscured by deposits of calcium carbonate and environ-
mental debris. The larvae were quickly cleaned of most of the
debris by soaking them in 10 percent hydrochloric acid (Lundbeck,
1907, and Johannsen, 1935). Active effervescence always accom-
panied this treatment.

The head of the larva was prepared for study by removing it from
the body and placing it in a test tube containing 2 percent potassium
hydroxide. The tube was placed in a beaker of water and boiled
slowly for 2 hours. The heads then were washed in glacial acetic
acid, dissected, and mounted on a slide. Photomicrographs were
prepared using the technique described by Howden (1955).

Historical Résumé

From 1720 to the present, numerous papers have been written
on various aspects of stratiomyid natural history as well as on mor-
phology and physiology of the larvae. Rather than attempt to
condense a voluminous literature in a few paragraphs here, I shall
discuss these works under their appropriate sections of biology,
morphology, phylogeny, and classification.

Studies on the biology and systematics of larval Stratiomyidae
can be divided into 2 rather distinct periods.

The first period (1720-1868) was devoted to the description of
local faunas. In most instances larvae were associated with adults
through rearing so that correct identifications were obtained; however,
no attempts at classification were made nor were other generalities
presented.

The second period, which was initiated by Brauer (1869), was
characterized by the incorporation of biological features and the

4. PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

characteristics of the immature stages in classification. Using
Schiner’s classification (1868) based on adults, Brauer demonstrated
that the immature stages of Diptera could be used as a check against
an already existing system of classification. Although Brauer did
not organize the Stratiomyidae into its component genera, he did
recognize that the Stratiomyidae, Tabanidae, Rhagionidae (Leptidae),
and Coenomyidae were closely allied and placed them in the tribe
Cyclocera under the Orthorrhapha-Brachycera.

Brauer (1883) once again reviewed the characters of dipterous
larvae and, on the basis of differences in head capsules, established
a new classification in which the Stratiomyidae and Xylophagidae
were considered as separate families but were placed together in the
Notacantha. In the same paper, Brauer divided the Stratiomyidae
into its various genera and listed the larval characters for each genus.

Austen (1899) presented a clear and concise review of the problem
regarding the classification of Solva, subsequent to which the majority
of dipterists have considered the genus to be located in the Xylomyinae
(Solvinae), a subfamily within the Stratiomyidae (for further details,
see page 13).

Malloch (1917) was the first person to arrange systematically
the North American Startiomyidae on the basis of larvae. As might
be expected, numerous genera were omitted since many larvae had as
yet not been described.

Johannsen (1922, 1935) continued the work of Malloch. He added
several new descriptions of larvae and supplied much of the data on
biology that we have today.

The most recent contribution to the classification of stratiomyid
larvae has come from Willi Hennig. In his outstanding work “Die
Larvenformen der Dipteren’”’ (1952) the species are listed for each
genus and their distribution given by zoogeographical region. In
addition, he has included biological notes for nearly all the species
mentioned.

Biology

The following discussion comprises a review of the literature on
the biology of the Stratiomyidae along the lines of a general life
history, with each study of a particular stage included under its
respective heading.

Irwin-Smith (1920, 1921) was the first to achieve some success
in rearing in a laboratory the complete life cycle of a species. Al-
though she was able to obtain a cycle from larva to larva, she was |
unable to determine the duration of the larval stage. More recently, |
Furman, Young, and Catts (1959) reported that newly hatched larvae |
of Hermetia illucens require 2 weeks or longer to attain full growth ©

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 5

under laboratory conditions and that the pupal stage may last from
2 weeks to 5 months. May (1961), also working with HZ. illucens,
gave more precise figures for the duration of the various stages in
the life cycle and, in addition, was able to complete the life cycle
successfully.

Eaes.—Eggs usually are laid in early spring by females that have
overwintered as larvae. When first laid, the eggs are pale yellow but
become progressively darker as the hatching date approaches. The
actual time required for hatching varies from 5 days to 3 weeks.

Hart (1895) mentions that the eggs of Odontomyia are “cigar-
shaped,”’ but this is not the case throughout the family. Wesenberg-
Lund (1943) reports that the eggs may be flattened or highly arched
or in the form of a “regular’’ egg. In addition, he mentions that the
eggs usually are placed in a distinct group or clump (the number
varying with the species) on vegetation overhanging water, in the case
of aquatic forms, or on decaying organic matter, in the case of terres-
trial forms. Wesenberg-Lund (1943) also mentions that the eggs of
Stratiomyidae are difficult to distinguish from those of Tabanidae.

Larvar.—Newly hatched larvae were noted by Irwin-Smith (1921)
to molt as soon as they emerged from the ege. Active feeding com-
menced immediately. As stated previously, the duration of the larval
state is unknown but the larvae of the majority of the northern
Nearctic species appear to be fully mature by fall. These mature
larvae are probably the overwintering stage, as pupation does not
occur until the following spring.

Larvae in each of the subfamilies exhibit a varying degree of simi-
larity with respect to their feeding habits (James, 1960), but there is
considerable variation in habitat selection by species within the
subfamilies. ‘Thus we have papers by Packard (1871), Lucas (1879),
Griffith and Packard (1882), and Brues (1924, 1928, and 1932)
recording the occurrence of larvae in salt water and in hot springs
and lakes. Pearson (1883) reported a larva that was found on an
exposed part of ocean beach, and Florentin (1899) found a great mass
of larvae in excessively saline pools.

The sum of our knowledge on habitats for the larvae of the North
American genera of Stratiomyidae is presented in condensed form
in table 1. Except for genera such as Actina, which are known only
from some other part of the world, the references are primarily to
North American workers. It is hoped that this table will not only
be an aid to collecting larvae but will also be a guide to the needs
for future studies within the family. Additional information on
biology of the larvae is given under the species descriptions.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Pupar.—tThe stratiomyid pupa, surprisingly, is much smaller than
the puparium, often being only one-third the total length of the
latter.

At the onset of pupation the mature larva becomes rigid, the
integument hardens, and the apical segments are usally in a distorted
position. The pupa itself, enclosed in a silky cocoon secreted by
the salivary glands, occupies only the anterior end of the puparium.
Miall (1895) reports that earlier workers often mistook the pupa for
a parasite that had devoured the larva.

Because of the shrinkage of the pupa, an air space is formed within
the puparium, and thus the aquatic pupal stage becomes buoyant.
As a result, the pupa floats on the surface of ponds or other bodies
of water until eventually it is either blown by the wind or carried
by the action of waves to the shore, where it remains with other
debris until the adult emerges.

At the time of emergence the puparium splits not only along the
dorsomedian line to the second abdominal segment, in the manner of
the Nematocera, but also transversely on the second abdominal
segment in the manner of the Cyclorrhapha.

Aputts.—Malloch (1917) and Johannsen (1935) have stated that
adult stratiomyids, of both terrestrial and aquatic forms, may be
seen flying about in areas where species of Umbelliferae and Com-
positae are in flower or resting on vegetation that is close to water.
Although there is relatively little information on feeding habits of
the adults, the regularity with which they are seen on vegetation
correlates with Lundbeck’s observation (1907) that the adults feed
on “plant juices” (nectar?).

The more commonly seen adults that represent species of Stratiomys
and Odontomyia are robust in appearance and are good fliers, but there
are many species in other genera that lack one or both of these at-
tributes. Other species, especially in the genus Sargus, are able to
hover about vegetation in the manner of a syrphid fly.

Adults may be collected by sweeping in grassy meadows, in bog
areas, and in sedges, cattails, and other vegetation along the margins
of streams, lakes, and ponds.

PREDATORS AND PARASITES.—Several investigators have observed
parasites and/or predators attacking larvae of Stratiomyidae. In
table 2, I have presented this information in condensed form and
have added my own observations.

Morphology

Several notable investigations on the morphology of the Stratio-
myidae have been made and have been concerned with the nervous
system, the respiratory system, the head capsule, and the integument.

NO. 3569 SOLDIER FLY LARVAE—McFADDEN a

While the quality of this work is very good, the number of different
species investigated leaves much to be desired. In this section,
an attempt has been made to incorporate previous work with my
own studies in a discussion of the major morphological characteristics
of the larvae, both external and internal.

INTEGUMENT.—The integument of a stratiomyid larva is composed
of many facets, some of which are distincly hexagonal while others
vary in size and shape. This faceted appearance has caused students
of Stratiomyidae to refer to the integument as being ‘“‘shagreened.’’?

Many early investigators (Miall, 1895; Leydig, 1860; Miiller, 1925;
and Kruper, 1930) noticed the presence of these facets and explained
their appearance as representing the upper surface of the “calcareous
nails” that are so evident in a cross section of the integument (fig. 6).
These nails are composed of calcium carbonate, but how they are
formed is unknown. Earlier workers were of the opinion that the
calcium deposits were secreted probably by the malpighian tubules
in an unknown manner. Richards (1951), on the contrary, is of the
opinion that the calcium deposits are laid down on the external sur-
face of the integument as a result of carbon dioxide diffusing outward
through the integument into the calcium-rich medium of water. His
theory, however, will not explain the development of the “nails”
under the acid conditions in which many species of stratiomyid larvae
occur, nor will it explain the development of the ‘‘nails” in terrestrial
species.

The “nails” provide the larva with a strong but flexible armor. To
test its resistence to attack one needs only to try to section it. Flex-
ibility is obtained by the pyramidal shape of the nails, which allows
the integument to bend.

HEAD AND APPENDAGES.—The total length of the head is always
greater than the width. This may seem misleading since the head is
permanently retracted approximately one-half of its length into the
thorax (fig. 3).

Distinctive eye prominences are situated on the lateral margin of
the ocular lobes. Located between the 2 ocular lobes is the clypeus,
which is limited by the frontal sutures. At the distal end of the
clypeus is the labrum, a structure that is not always clearly defined
posteriorly by the clypeolabral suture. The mandibular-maxillary
complex is located in the pocket formed by the labrum and the ex-
tensions of the ocular lobes. The complex itself is formed by the
fusion of the mandibles and the maxillae as the name suggests. It is
so articulated that movement is in a plane perpendicular to the long

3 Derived from the near-eastern word “‘shagreen,”’ a type of uncured leather
prepared in that area. As used here, it doubtless refers to the multifaceted
integument that suggests this type of leather.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

axis of the body. Cook (1949) states that the mandibular-maxillary
complex is used for sweeping food into the oral cavity rather than for
chewing.

The 2-segmented antennae are located near the apex of the ocular
lobes but the exact position varies with the species as does the size
and shape.

On the ventral surface of the head (fig. 4) there is a membranous
area that outlines the labium. The large posterior part of the labium
is the submentum. Directly anterior to the submentum is the pre-
mentum and the palatum. The latter structure, as pointed out by
Cook, is apparently an adaptation for rooting up material. Cook also
reported that labial palpi and a hypopharynx are lacking in stratio-
myid larvae. The arrangement of the previously mentioned struc-
tures is shown in figure 5. The most important feature illustrated
here is the arrangement of 2 sclerotized plates that are located beneath
the pharynx at the posterior end of the head capsule. These struc-
tures function as a mortar and pestle, grinding up the food material
that is swept into the mouth by the manibular-maxillary complex
(Bischoff, 1925; Cook, 1949). The bowl-like mortar is very heavily
sclerotized and convoluted while the pestle, which is formed by the
floor of the pharynx, is modified in the form of a heavy, corrugated
plate and fits into the bowl of the mortar.

If the mortar and pestle do indeed perform the task of grinding the
food, it seems inconceivable that stratiomyid larvae could be pre-
daceous on other larvae as has been reported, especially when the size
of the oral cavity is taken into consideration.

THorax.—The thorax is composed of the first 3 body segments
posterior to the head. The noteworthy characteristics found here are
the prothoracic spiracles, which vary in size and shape, the chaetotaxy,
and the shape of the segments, which seem to become broader toward
the metathoracic segment. Stratiomyid larvae are without legs or
prolegs.

AsBpoMEN.—This tagma consists of the 8 body segments posterior
to the metathoracic segment. They are similar in form to the thoracic
segments but differ in chaetotaxy and in occurrence of special organs
or structures. Several genera can be distinguished by the markings
or color patterns that occur on the thoracic and abdominal segments.

The first and second segments are usually the broadest. The re-
mainder of the segments either taper toward the apical segment
(aquatic species) or several segments are parallel sided and the re-
mainder taper toward the apical segment (terrestrial species). At the
extreme end of the apical segment, which is often much longer than
it is wide (fig. 107), especially in the aquatic species, there is located
a transverse fissure that houses the spiracular chamber. In the genus

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 9

Nemotelus, however, the spiracular chamber is located on the dorsum
of this segment.

A few aquatic genera, notably Euparyphus and Odontomyia, possess
strongly curved sclerotized hooks or spines, as they sometimes are
called (fig. 122). They are usually on the venter of the seventh ab-
dominal segment, but in the genus Hedriodiscus they are on the venter
of the sixth segment as well. A remarkable feature of the spines is
that they curve forward toward the head. This arrangement would
seem to present some difficulty in forward movement, but this has
not been observed. Previous workers (Wesenberg-Lund, 1943; Lenz,
1923) have reported that the hooks serve as anchors to keep the larvae
from being swept away by fast-flowing currents. This statement may
be partially correct, but I have observed this type of larva in shallow
pools that had only convection currents.

DIGESTIVE systEM.—The digestive system has little to offer in the
way of taxonomic characters other than the characteristic shape of
the gut, which is long and highly convoluted. Except for the mortar
and pestle, there are no sclerotized areas present in either the foregut
or the hindgut.

MALPIGHIAN TUBULES.—The malpighian tubules are filled with a
whitish, semiliquid material that is reported to be primarily composed
of calcium carbonate. This material may be associated in some
manner with the calcium carbonate that is secreted in the integument.

RESPIRATORY SYSTEM.—Stratiomyid larvae have frequently been
described as peripneustic, but it is doubtful that they are functionally
so. They appear to be metapneustic or in some cases amphipneustic.
The lateral spiracles almost always are minute and difficult to detect,
especially in the later instars. The only spiracles that function con-
tinually are the terminal or posterior spiracles.

The spiracular chamber contains the ends of the main tracheal
trunks. In the aquatic forms, the lips that form the opening to the
chamber are fringed with long hydrofuge setae, which enable the
larvae to remain suspended from the surface of water for indefinite
periods of time while the spiracles are open to the atmosphere. When
submerged, these setae also aid respiration by enclosing an air bubble
trapped in the process of submerging.

Using the work of Whitten (1959) as a guide, I examined the tracheal
system of 4 genera (2 aquatic and 2 terrestrial) for possible taxonomic
characters. The tracheal system of each larva has the same general
pattern as that given by Whitten for brachycerous larvae and, con-
sequently, cannot be used as a distinguishing character beyond the
suborder.

Nervous system.—Although several studies have been made on
the nervous system of a few species of stratiomyid larvae, the results

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

appear to have little consequence from a taxonomic standpoint
(Viallanes, 1882a, 1882b, 1885; Kiinckel d’Herculais, 1879; Henneguy
and Binet, 1892).

Phylogeny

Handlirsch (1908) postulated that the Stratiomyidae arose in
Jurassic times from a primitive tipuloid stock that had evolved to
the point where it more closely resembled the Xylophagidae than
the Tipulidae. The families most like the Stratiomyidae, the Xylo-
phagidae and the Rhagionidae (Leptidae), are similar to the stra-
tiomyid subfamily Xylomyinae but only in the adult stage (wing
venation, spurs on tibiae, etc.). From the Rhagionidae arose the
Tabanidae and Therevidae and, in turn, this line of descent led di-
rectly to the more advanced Diptera (Handlirsch, 1908; Bischoff,
1925; and Lindner, 1937).

The most primitive subfamily in the Stratiomyidae is the Xylomyi-
nae. Characters occuring in the immature stages of this subfamily
that I consider to be primitive are an integument that is only partly
shagreened, the generalized type of its mouthparts, an incompletely
enclosed pupa, and the type of habitat, which is usually under bark
or in rotten logs. Characters of the adults are equally primitive.

From this primitive subfamily 4 lines appear to have evolved,
one of which represents a continuation of the basic type with slight
modifications in morphology (Xylomyinae, Beridinae, and Pachy-
gastrinae). The other lines represent a divergence toward different
ecological zones in the larval stage: one to an aquatic or semiaquatic
mode of life (Stratiomyinae and Nemotelinae); another to life in a
rich, soft food source such as animal excrement and decaying organic
matter (Sarginae and Clitellariinae); and another line, represented
by Altermetoponia rubriceps (Macquart), which lives in sod
(Chiromyzinae).

As previously indicated, the environment in which the larvae live,
their feeding habits, and the type of mouthparts are closely allied
and provide an insight into the evolution of the family. Table 3
illustrates the distribution of these characters through the subfamilies.
In an attempt to show degree of relationship, I have analyzed 11
characters and given them an arbitrary numerical value (for details
of this technique, see James, 1953). A list of the characters and their
numerical values is as follows:

A. Habitat of larva
1. aquatic or semiaquatic
2. terrestrial
3. terrestrial-arboreal (close association with trees)

NO. 3569 SOLDIER FLY LARVAE—McFADDEN i

B. Food of larva
1. aquatic microorganisms
2. plant roots
3. decaying organic matter (includes microorganisms)
C. Labium of larva
1. well developed
2. not well developed
D. Distribution of maxillary setae in larva
1. setae absent
2. setae present, located in notch
3. setae present, other arrangement
KE. Types of maxillary setae in larva
1. cylindrical brushes
2. cylindrical brushes absent or not cylindrical
F. Transverse series of dorsal setae with accompanying setae in larva
1. extra setae present
2. extra setae absent
G. Posterior spiracles of larva
1. with coronet of setae
2. without coronet of setae
H. Anal armament of larva
1. large teeth anterior to anal opening
2. large teeth absent
I. Body conformation of larva
1. attenuate
2. parallel sided
J. Mandibles of larva
1. well developed
2. not well developed
K. Posterior spiracles of larva
1. located at apex of last abdominal segment
2. located on dorsum of last abdominal segment

These results are presented in graph form in figure 1. The phylo-
genetic tree in figure 2 represents the sum of the similarities and dif-
ferences among the subfamilies and indicates the relationships of the
extant species.

The Beridinae and succeeding subfamilies of the Stratiomyidae
probably arose from a group of the subfamily Xylomyinae that had
developed a completely shagreened integument. This separation
was no doubt strengthened subsequently by 2 changes in the
ecological requirements of beridine larvae: a change in habitat
(larvae of Beridinae are not found in the same habitat as larvae of
Xylomyinae) and a change in food preference (as evidenced by larvae
of Actina incisuralis).

The subfamily Pachygastrinae represents an advanced group on
the basis of adult characters, but the mouthparts of the larvae are

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

similar to those found in the larvae of Xylomyinae and Beridinae.
In addition, pachygastrine larvae are found under the bark of trees
as are larvae of Xylomyinae. During the course of time, however,
the chaetotaxy and size of the larvae have evolved.

On the basis of similarity of mouthparts, the Clitellariinae and
the Pachygastrinae appear to have arisen from a beridine stock.
The former represents a radiation into a new ecological zone, and
the latter represents evolutionary improvement for living in an
already occupied niche. Clitellariine larvae have mouthparts much
the same as beridine larvae except for the labium, which has become
more sclerotized and spatulate in the former. This change in the
labium is probably an adaptation for feeding on the pulpy flesh of
succulents.

The Sarginae appear to have arisen from a clitellariine stock,
probably closely related to the tribe Hermetiimi. The mouthparts of
the latter are very similar to those of the Sarginae and both groups
live in the same type of enviroment—decaying organic material.
Both groups are highly evolved: the mouthparts have degenerated
so that the mandibular-maxillary complex is almost indistinguishable.

The subfamily Nemotelinae, although similar in general appear-
ance to the Stratiomyinae, apparently has evolved from a clitel-
lariine stock. This cleavage, however, must have occurred shortly
after the Clittellariinae had broken away from the pachygastrine
stem since all three taxa are closely related (fig. 2). Both adult and
larval characters of the Nemotelinae appear to substantiate the con-
clusion that the Stratiomyinae and Nemotelinae are not closely re-
lated and probably never have been.

The Stratiomyinae have few characters in common with any of
the extant subfamilies. Larval characters indicate that the sub-
family probably evolved from a beridine stock at a very early period.

Because the Chiromyzinae are represented in North America by
only Altermetoponia rubriceps—an introduced species—no further
consideration need be given to the phylogeny of this subfamily.

Classification

The classification of the Stratiomyidae presented here is based
upon opinions and conclusions drawn from the relationships of the
subfamilies presented in the previous section. This system differs
somewhat from the classification of James (pers. comm.), which is
based upon adult characters. A comparison of the 2 systems is
as follows (asterisks indicate genera not seen in immature stages and
thus not in my classification):

NO. 3569 SOLDIER FLY LARVAE—McFADDEN LS

James McFadden

1 1. Xylomyinae—Xylomya, Solva

2. Chiromyzinae—Altermetoponia 2. Chiromyzinae—as in James

3. Beridinae—Allognosta, Actina, 3. Beridinae—as in James
Beris, Exodontha

4. Sarginae—Sargus, Ptecticus, 4. Sarginae—as in James
Microchrysa, Chloromyia, *
Merosargus

5. Cyphomyiinae—Cyphomyia, 5. Cyphomyiinae—see under
Dicyphoma Clitellariinae

6. Hermetiinae—Hermetia 6. Hermetiinae—see under

Clitellariinae

7. Clitellariinae—Adozomyia, 7. Clitellariinae—Dieuryneura
Dieuryneura, Brachycara,* Cyphomyiini—Cyphomyia,
Euryneura,* Oxycera,* Adozomyia, Dicyphoma
Euparyphus, Caloparyphus, Hermetiini— Hermetia
Akronia,* Nemotelus

8. Stratiomyinae—Stratiomys, 8. Stratiomyinae—Stratiomys,
Hoplitomyia,* Labostigmina,* Hedriodiscus, Odontomyza,
Anoplodonta,* Hedriodiscus, Myxosargus, Euparyphus,
Odontomyia, Nothomyia,* Caloparyphus, Aochletus
Myzxosargus

9. 9. Nemotelinae—Nemotelus

10. Pachygastrinae—Neopachygaster, 10. Pachygastrinae—as in James

Eupachygaster, Pachygaster,
Zabrachia, Berkshiria

There are 2 major points of difference between the present
arrangement and that of James. First, the latter includes the genera
Xylomya and Solva in a separate family, the Xylomyidae. AlI-
though there is little information in the literature regarding the
larva of Xylomya, many workers have recorded their thoughts on the
larva of Solva and its importance in classification; thus, we have
papers by Westwood (1840), Osten Sacken (1882), Brauer (1883),
Lindner (1937), and Hennig (1952) pointing out that Solva should be
placed in the Stratiomyidae. In 1899 Austen presented a summation
of previous work concerning the classification of Solva and came to
the conclusion “that Solva represents a primitive ancestral form of
Stratiomyidae, given off from the common stem after the evolution
of the characteristic type of larva and mode of pupation, but before
the assumption on the part of the imago of the equally characteristic
features (venation, spurless tibiae, etc.) exhibited by the more special-
ized types of the family.”” Inasmuch as the larvae of Xylomya and
Solva are almost indistinguishable morphologically, I have extended
Austen’s discussion to include Xylomya and have reorganized the
subfamily Xylomyinae.

A second area of disagreement between the 2 systems is in the
classification of the Clitellariinae. A study of the mouthparts re-
vealed that there were 3 distinct types present in this 1 subfamily.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

One type was found in the genus Adozomyia. 'The mouthparts of
the species in this genus are very similar to those found in the larva
of Cyphomyia. In addition, the larvae of both genera are known
only from wounded or decaying succulents in North America; con-
sequently, Cyphomyia and Dicyphoma have been transferred to the
Clitellariini (Cyphomyinae of James) along with Adozomyia.

A second tribe, the Hermetiini (Hermetiinae of James), has been
transferred to this subfamily on the basis of similarities in morpholog-
ical and ecological characters of both adults and larvae of Hermetia
to the clitellariine genera Adoromyia, Cyphomyia, and Dicyphoma.

The genus Dieuryneura has been placed in this subfamily also but
not in either tribe. A comparison of larval mouthparts shows it to
be more closely related to the Hermetiini than to the Clitellariini,
but a more definite attempt at classification is impossible at this time.

The larvae of the remaining genera in the Clitellariinae of James
are either aquatic or semiaquatic and the mouthparts are similar
to the genera originally placed in the Stratiomyinae. These 2
facts by themselves give sufficient cause to unite the genera in a
single subfamily; in addition, all of the genera except Nemotelus
that were removed from the Clitellariinae possess ventral hooks.
These curious structures may represent a clinging device for living
in fast-flowing water. Stratiomys larvae lack these structures but
they are present on larvae of Odontomyia. It is possible that the
hooks have evolved twice but it seems more likely that the genera are
more closely related than they were previously thought to be.

Larvae of Nemotelus pose a special problem. In addition to lack-
ing the ventral hooks, the orifice of the spiracular chamber is on the
dorsal surface of the last abdominal segment and possesses a cornet
of hydrofuge setae as contrasted with the terrestrial larvae that lack
this feature and as contrasted with other aquatic larvae that also
possess a coronet of hydrofuge setae but have the spiracular chamber
at the apex of the last segment. This combination of characters is
rather unique and provides the basis for elevating this genus to
subfamilial status.

Key to Suborders, Divisions, and Principal Families of Brachycera-
Orthorrhapha Based on Larval Characters

1. Head complete, or the posterior portion with deep longitudinal incisions;
mandibles capable of horizontal movement; body consisting of 13 segments
in addition to head; with 9 pairs of spiracles . . Suborder NEMATOCERA

Head incomplete, frontal region not strongly arched; mandibles capable of
vertical movement; body comprising fewer than 13 segments and only ex-
ceptionally with as many as 9 pairs of spiracles.

Suborder BRACHYCERA 2

a

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 15

2. Free cephaloskeleton present; head very poorly developed, completely un-
sclerotized dorsally; antennae poorly developed or absent, when present
situated on a membranous surface; mandibles short and hooklike, usually

capable of protrusion much beyond the poorly developed maxillae; maxillary
palpi rarely visible. . ... ye ee Division CYCLORRHAPHA
No free cephaloskeleton within fhe! head capsule, the exoskeleton of the head
at least dorsally indicated; antennae well developed, situated on the upper
surface of the lateral lobe or on the slightly arched, sclerotized frontal plate;
mandibles normally sickle shaped, not protruding much beyond apices of
the well-developed maxillae, often much shorter, maxillary palpi distinct.
Division ORTHORRHAPHA 3

3. Posterior spiracles situated within a terminal or subterminal cleft or chamber,
usually concealed, or with a terminal breathing tube; pupa free or enclosed

AAS tueaveule skal eg) seul a Te Ae ae eA
Posterior spiracles visible, atuedie on po analimaatore or Se penne
segment, body not shagreened or visibly striated; pupa free. . ... .5

4. Body 12-segmented, cylindrical, not eieeeeceel usually longitudinally
striated, abdomen with a girdle of pseudopods on each segment; head
retractile; spiracular fissure vertical. . . dials pa LABDANDDAE

Body 11-segmented, bristly, surface finely peso without pseudopods;
head not retractile; spiracular fissure transverse, sometimes rather small;
peripneustic or amphipneustic ..........STRATIOMYIDAE

Posterior spiracles situated on penultimate or antepenultimate segment.

THEREVIDAE, SCENOPINIDAE, MYDAIDAE, ASILIDAE,
BOMBYLIIDAE

Posterior spiracles situated on apical segment .... . sshd [tee “ol
6. Last abdominal segment obliquely truncate and with Sraicntiee processes;
projecting portion of head and the flattened apical plate of last abdominal
segment heavily sclerotized, the former cone shaped. . XYLOPHAGIDAE
(including Rhachiceridae and Coenomyidae)

Last abdominal segment lacking projecting processes; projecting portion of
head not pointed cone shaped; last abdominal segment without a heavily
sclerotized flattened terminal plate . . . . . NEMESTRINIDAE,
EMPIDIDAE, DOLICHOPODIDAE, RHAGIONIDAE, ACROCERIDAE.

on

Family Stratiomyidae

In addition to the characters given in the key to families, stratiomyid
larvae may be readily distinguished from other dipterous larvae (ex-
cept for psychodid larvae) by the presence of calcium carbonate
deposits on the integument (see p. 7).

Famity piaGNosis.—Head elongate and narrow except for ocular
prominences; not retractile; antennae usually located anterolaterally
but varying among species; setae on dorsum of head variable; body
11-segmented; prothoracic segment with functional spiracles, that
segment wider than the head; segment posterior to prothorax either
parallel with the lateral margins of the prothorax or tapering in a
posterior direction from the second abdominal segment as in Stratiomys
(fig. 105); posterior spiracles located in a transverse cleft on last
abdominal segment; with or without a fringe of hydrofuge setae;

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

pseudopods lacking, but strong curved hooks may be present on venter
of penultimate and/or antepenultimate segment.

Key to Larvae of Subfamilies of Stratiomyidae of America North of
Mexico

1. Last abdominal segment with a coronet of plumose or pinnate setae at the

DOK yee cers Jo ee 2 SERA ELOMY TNAB

Last abdominal peement ei caeenee of plumose or Be setae absent
FLOM APEXce oe oy HN canieees

2. Last abdominal eens race a peoranee Gs Sree or iam setae on
GOTSUIR. Pa... ... . .NEMOTELINAE
Last abdominal er Pelays a cor Sones of oTamene or pinnate setae . .3

3. Pro- and mesothoracic segments with a smooth field on dorsum; anus bordered
anteriorly by a transverse row of strong, posteriorly directed teeth.

XYLOMYINAE
Pro- and mesothoracie segments with normal shagreened pattern on dorsum;
anus not bordered anteriorly by teeth ... . .4

4. Mouthparts highly sclerotized and foreshortened, snag tisiles eet Bereiaped
no setae or bristles present on mouthparts (fig. 15); recorded from San
Francisco area only .... soe 0 6 SAOIENOROIM ICANN:

Mouthparts with a different eaepinaioe of characters than those listed
above; occurring in San Francisco area or elsewhere .. . AC KAT necO.

5. Dorsal row of transverse setae surrounded by smaller setae; Poe as
in figure 16; uniformly colored larvae .. . ee S oe oe DER EDINAR

Dorsal row of transverse setae not surrounded By smaller setae; mouthparts
different than above; larvae uniformly colored or witha pattern ... .5

6. Larva with a well-developed labium (fig. 19); primarily restricted to south-

western United States, where found in Opuntia or Agave species.
CLITELLARIINAE, Tribe CLITELLARIINI
Larva lacking a well-developed labium; occurring in southwestern United
States and elsewhere; not recorded from hosts given above ......7

7. Small larva, under 10 mm in length; mouthparts similar to those of Beridinae
(fig. 18); found under bark of trees ...... .PACHYGASTRINAE

Larva over 10 mm in length; mouthparts lacking sclerotized areas . . . .8

8. Mouthparts simple, setae restricted to a linear patch on inner margin of

mandibular-maxillary complex (fig. 21); large, robust, color uniform.
CLITELLARIINAE, Tribe HERMETIINI
Mouthparts more elaborate, setae present on mandibular-maxillary complex,
in addition to setae restricted to notch on inner margin of complex (fig. 17) ;
smaller, usually with a vittate pattern ........ =. .SARGINAE

Subfamily Xylomyinae

This subfamily is represented in North America by the 2 genera
Xylomya Rondani and Solva Walker. It is considered the most prim-
itive group of the Stratiomyidae because it shares at least 1 charac-
ter of the adult stage with the corresponding stage of the generally
more primitive Xylophagidae.

SUBFAMILIAL CHARACTERS.—Mouthparts as in figure 14. Other
characters as given in key to subfamilies.

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 17

HABITAT OF LARVAE.—The few records available indicate that the
larvae live under the bark of trees and in rotting logs.

Key to Genera of Xylomyinae

Prothoracic segment of larva with a tubercle anterior to the spiracle giving a cleft
appearance to margin (fig. 10); apical abdominal eee with a transverse row

of tubercles on dorsum .. . . . .. .Solva Walker
Prothoracic segment of larva nears alent appearance ot margin; no tubercles pres-
ent on dorsum of apical abdominal segment. . . . . . .Xylomya Rondani

Genus Xylomya Rondani, 1861

Of the 7 species listed by James (pers. comm.) for this genus, the
larvae of only 2 species have been collected and 1 lacks a specific
determination.

GENERIC CHARACTERS.—As given in key to genera.

Xylomya americana (Wiedemann), 1821
Ficures 149, 150

DescripTion.—Puparium: length (head capsule lacking) 15.2 mm;
width 4.1mm. Other characters as given in key to genera.

Biotocy.—Larvae of this species were removed from a hole in a
sycamore tree.

MATERIAL EXAMINED.—Virginia: Alexandria, Oct. 14, 1951, W. W.
Wirth, 1 puparium (USNM).

Xylomya species
Ficurss 7, 11, 25

Description.—Mature larva: length 13 mm; width 3.2mm. Other
characters as given in key to genera.

Brotoay.—No data available for this species.

MATERIAL EXAMINED.—Massachusetts: Amherst, 1945, M.E.S., 1
larva, MSC.

Genus Solva Walker, 1860

James (pers. comm.) lists 3 species for this genus but the larva of
only 1 species, Solva pallipes (Loew), has been collected.

Lindner (1938) and Hennig (1952) include the species of Xylomya
in this genus, but Steyskal (1947) has demonstrated that the adults
of these 2 genera can be distinguished. This shows that the immature
stages can also be differentiated.

GENERIC CHARACTERS.—As given in key to genera.

Solva pallipes (Loew), 1863
Fiaures 8-10, 12-14
Description.—Mature larva: length 7-8 mm, mean 7.8 mm; width
1.75-2.0 mm, mean 1.85 mm. Other characters as given in key to
species.
221-533 67-2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Pupa (after Greene, 1926): Very thin, shining, transparent, with
a yellowish tinge. Antennal capsules large, poimting outward,
faintly annulated to the tip; just posterior to the base of these are 3
faint ocellar punctures. Thorax slightly longer than wide, smooth;
wing pads smooth, reaching to the apex of the third abdominal
segment; thoracic spiracles situated on a small elevation; the spiracular
entrance is golden yellow, sinuous, and contains a great number of
short radiating slits. Abdomen cylindrical, composed of 8 segments;
first and last segment without any spines; segments 2-7 with a
transverse, dorsal row of reddish-yellow spines just posterior to the
middle of the segment; segments 1-7 have a spiracle on the antero-
lateral surface; spiracle small, golden yellow, slightly elevated; last
segment rounded at apex and entirely smooth. Length 6.0 mm;
diameter of thorax 1.85 mm; diameter of abdomen 1.75 mm.

BroLtocy.—Larvae have been collected from the following trees:
Ulmus species, Populus species, Robinia species, and Liriodendron
tulipifera. The larvae seem to prefer trees on which the bark has
become loosened. Malloch (1917) intimates that he found the larva
of Solva to be predaceous on the larvae of Euzesta, Lonchaea, and
Heterominga. Greene (1926) states that the larvae may be predaceous
but are usually scavengers.

As the larva of Solva matures, it forms a pupa within the last
larval skin as is the case with all other stratiomyid larvae; however,
before emergence, the puparium splits dorsally down the median line
and the pupa forces itself about two-thirds of the way out of the
puparium and remains there until the adult emerges. ‘The emergence
of the pupa causes the head of the puparium to become loose and
often completely separated as in the higher flies. Adults of Solva
can be found on tree trunks in June, July, and August.

MATERIAL EXAMINED.—Illinois: Rock Island, Apr. 5, 1932, C. O.
Mohr, 2 larvae, INHM. Pennsylvania: Blain, Aug. 6, 1955, F.
Craighead, lot no. 55 11825, 6 larvae, 7 puparia from oak, USNM.
Wisconsin: Madison, July 1938, Dodge, 23 larvae, 6 puparia
from under elm bark, OSU.

Subfamily Chiromyzinae

Although the genus Altermetoponia Miller has been placed in this
subfamily, there is some doubt as to whether it should be left here or
placed in a subfamily of its own as Hennig (1952) has done. Because
of a scarcity of specimens in related genera within the subfamily,
it has been impossible to make a comparison of characters. Raff
(1931) has provided a vague description of the larva of Chiromyza

ee Le

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 19

australis Macquart, the only published information available on the
larvae of this subfamily.

The subfamily is represented in North America only by Altermeto-
ponia rubriceps (Macquart).

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

HABITAT OF LARVAE.—The only species of Chiromyzinae for which
habitat data are available is A. rubriceps.

Genus Altermetoponia Miller, 1945
Fieures 15, 25, 33, 34

DescripTion.—Mature larva: length 8.8-9.8 mm, mean 9.27
mm; width 1.8-2.0 mm, mean 1.87 mm. Body segments yellow;
head brown, darkest at apex.

Brotoay.—It is quite probable that this species will assume eco-
nomic importance in the area around San Francisco, just as it has
come to be considered a pest in Australia, where it causes extensive
damage to lawns. Other than evidence it lives in sod as a larva,
nothing is known about its life history in North America.

MATERIAL EXAMINED.—California: San Francisco, Mission Park,
Nov. 20, 1959, T. R. Haig, 35 larvae, 3 puparia, CAS.

Subfamily Beridinae

Of the 5 genera that represent this subfamily in North America,
the larvae of only 2 are known: Allognosta and Exodontha; however,
larvae of 2 of the remaining genera are known from other geographical
areas and descriptions of these species have been included.

SUBFAMILIAL CHARACTERS.—As given in the key to subfamilies.

HABITAT OF LARVAE.—The larvae in this subfamily are terrestrial
but may be associated with semiaquatic environments; e.g., Fuller
(1934) found larvae of Actina in and on the carcass of a dead sheep,
Johannsen (1922) reported larvae of Allognosta from decaying organic
material, and the larvae of Beris have been found in decaying leaves.
In addition to bemg the center of the larvae’s food source, the semi-
aquatic medium facilitates their movement and ingestion.

Key to Genera of Beridinae

1. Larva robust, greaterthanl10 mminlength. .. . . . Exedontha Rondani
Larva smaller, less than 10 mm in length. ......... Ree eRe cot

2. Abdominal segments with lateral margin bilobed, the anterior smaller than
he; POsterlOrONG cicen 5. Ss hs =) 5 3 st . . . Allognosta Osten Sacken
Abdominal segments with lateral margin smooth .... . nt cient

3. Last abdominal segment broadly indented at distal end and ernged with fine
BELA IT RMT hy Senaa tees Lee roma is Cag a) Bae pert R aL BR CS fi . Beris Latreille

Last abdominal segment not indented at distal end; fringe lacking
Actina Meigen

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Genus Allognosta Osten Sacken, 1883

There are 4 species recognized in this genus, but only the larva of
Allognosta fuscitarsis (Say) has been reported (Johannsen, 1922).
GENERIC CHARACTERS.—As given in the key to genera.

Allognosta fuscitarsis (Say), 1823
Figures 31, 36-38

DescripTion.—Mature larva: length 9.5 mm; width 3.5 mm.
Prothoracic spiracle located in a distinct marginal notch. Other
characters as given in key to genera.

Brotogy.—Little is known of the biology and life history of this
species. Malloch’s (1917) specimen was obtained from woods near
Urbana, Ill., while Johannsen’s report (1922) stated that his speci-
mens were found in decaying organic matter.

MATERIAL EXAMINED.—Rhode Island: Kingston, Aug. 20, 1942,
H. Knutson, 1 early instar larva from glass of milk, MSC. New York:
Ithaca, C. Hamilton, 2 larvae, 1 puparium, CU; Ithaca, 2 early instar
larvae collected with larvae of Fannia species, CU. North Carolina:
Great Smoky Mountain National Park, Oct. 4, 1951, J. S. Ayars,
acc. no. 49768, 11 larvae, NCS.

Genus Actina Meigen, 1804

Only 1 species of this genus is known from North America, the
immature stages of which have never been reported.

The characteristics that are given below are taken from a paper by
Fuller (1934) in which she described the immature stages of an
Australian species, Actina incisuralis Macquart.

GENERIC CHARACTERS.—As given in key to genera.

Actina incisuralis Macquart, 1847
Figures 40, 41, 45

Description.—Mature larva (?): length 9 mm. Color varying
between greyish brown and brown.

Head elongated, narrow and pointed. Body strongly constricted between the
segments giving the larva a scalloped appearance along the sides. Dorso-laterally
the integument between each abdominal segment, and between the third thoracic
and first abdominal, is produced into a small papilla projecting outwards. The
thoracic segments become broader passing from the head, the abdominal segments
are of uniform width and length, with the exception of the eighth, which is nar-
rower and longer than the others, more flattened and produced into two blunt
projections at the posterior corners (Fuller, 1934, p. 191).

Setal arrangement typical for family and subfamily.
Brotoay.—Fuller (1934) reported that larvae of this species ‘‘were
found in abundance on the undersurface of a sheep carcass”? and ‘‘on

NO. 3569 SOLDIER FLY LARVAE—McFADDEN Dit

the earth under masses of rotting grass” (p. 190). Larvae were also
present in soil among the roots and stems of growing grasses. Pupae
were obtained 7-8 months after the larvae were collected.

MATERIAL EXAMINED.—No larvae of this species were examined.

Genus Beris Latreille, 1802

Three species are recorded for this genus in North America but
the larvae or pupae have never been reported. The larval stages of
a few European species of Beris have been described (Lenz, 1923).
The larva described below was loaned to me by W. Hennig.

GENERIC CHARACTERS.—As given in key to genera.

Beris vallata Forster, 1771
FIGURES 23, 26, 28

Description.—Mature larva (?): length 7-8 mm; width 2 mm.
Head and body segments yellow; body of larva extremely flattened
dorsoventrally; prothoracic spiracle elevated but relatively incon-
spicuous; body segments from second thoracic to seventh abdominal
with a distinct papilla at intersegmental fold.

Brotogy.—Nothing is known about the biology of the North
American species. Lenz (1923) reports that larvae of Beris vallata
and 2 other species of this genus were found under the bark of a
fallen tree, which was lying in a marshy area between fallen leaves
(presumably wet) and at the margins of springs.

MATERIAL EXAMINED.—Two larvae borrowed from the Deutsches
Entomologisches Institut; no collecting data.

Genus Exodontha Rondani, 1856

According to James (pers. comm.), 2 species occur in North America,
but larvae and pupae of only Erodontha luteipes (Williston) have
been found.

GENERIC CHARACTERS.—As given in key to genera.

Exodontha luteipes (Williston), 1885
FiaureEs 29, 30, 35

Descrietion.—Mature larva: length 11.7-15+ mm, mean
13.23 mm; width 3.8-4.5 mm, mean 4.77 mm. Head and body seg-
ments pale yellow.

Brotoay.—Larvae of this species were found in moist rotting wood
under large boulders on a mountainside at an elevation of approxi-
mately 6000 feet. Adults emerged on June 27, 1960, inthelaboratory.

MATERIAL EXAMINED.—Alberta: Banff National Park, Mt. Ishbel,
June 13, 1960, Ball, Madge and McFadden, 1 larva, 4 puparia MWM.

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Subfamily Sarginae

Of the 6 genera listed for this subfamily by James (pers. comm.),
larvae have been described for only the following 4 genera: Sargus,
Ptecticus, Microchrysa, and Merosargus.

The keys to the genera of this subfamily will work for mature
larvae only, but, where earlier instars are known, descriptions of
these forms are included.

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

HABITAT OF LARVAE.—Larvae of the Sarginae are terrestrial
scavengers. They have been collected from various types of manures,
rotting vegetation—especially garden refuse—and from other types
of decaying organic matter.

Key to Genera of Sarginae

1. Venter of sixth abdominal segment with sternal patch extending almost the
length of that segment; venter of fifth abdominal segment with a smaller

sternal patch (fig. 50) . Sates s ... . . Ptecticus Loew
Venter of sixth abdominal ase ite a eae sternal patch; sternal patch
lacking on venter of fifth abdominal segment. ... . Swollen

2. Dorsolateral margin of segments 1-10 with a series of eae dark plaques
arranged in a circular pattern; segments 1-3 with a transverse band of
these plaques connecting the 2 lateral circles; 5 vittae on dorsum, the mesal
vitta much broader than the others. . . . Sargus (Pedicellina) James

Dorsolateral margin of segments 1-10 lacking markings described above . . 3

3. Lateral margin of head lacking a protruding tubercle; no constriction of head

posterior to eye prominence . . . Sargus Fabricius (sensu stricto, in part)
Lateral margin of head with a protruding tubercle; head constricted posterior
tom tuberclemsman.. |e Ae ADS, oe! ced “eye ge!

4. Eye prominence anterior 2 Sonia Theat fabeecles . Merosargus Loew

Eye prominence on protruding lateral tubercle .... Thus ee aoe OD

5. Width of head at tubercle not more than 0.5 mm... UVilerachevan Loew

Width of head at tubercle 0.8-1.0 mm.
Sargus Fabricius (sensu stricto, in part)

Genus Sargus Fabricius, 1798

Larvae of this genus are almost impossible to identify to species
unless they are fully mature specimens. Earlier instars lack the char-
acteristic patterns of the species and the head capsule apparently
varies from instar to instar.

GENERIC CHARACTERS.—As given in key to genera.

Subgenus Pedicellina James, 1952
Sargus (Pedicellina) lucens Loew, 1866
Figures 49, 56, 61

Description.—Mature larva: length 9.2-11 mm, mean 10.1 mm;
width 2.5-3.3 mm, mean 2.85 mm. Other characters as given in key
to genera.

a

f
;
:

NO. 3569 SOLDIER FLY LARVAE—McFADDEN ey

Broutoay.—Larvae of this species have been collected from leaf axils
of cattails only where the axils were above the waterline.

MATERIAL EXAMINED.—New York: Ithaca, Apr. 8, 1921, P. W.
Claasen and O. A. Johannsen, 5 larvae, 15 puparia collected from leaf
axils of cattails, CU; Ithaca, July (?), 2 larvae from leaf axils of
cattails, CU.

Subgenus Sargus Fabricius, 1798

Key to Species of Subgenus Sargus

1. Body segments lacking vittae; prothoracic spiracle on stalk like structure.
elegans Loew
Body segments with alternating dark and light vittae; prothoracic spiracle

RCHEUI C Soi mene ee ch ens Wie Bw Ao tee cas ol as noun a Yc ve Se eth rete ee
2. Dorsal surface of body segments with 3 dark vittae. .. . decorus Say
Dorsal surface of body segments with more than 3 dark vittae ..... 3

3. Median light band between inner 2 dark vittae parallel sided.
bipunctatus Scopoli
Median light band between inner 2 vittae geniculate; each of inner pair of
setae located in a dark patch in median light band. . . . . viridis Say

Sargus (Sargus) elegans Loew, 1866
Ficures 52, 53, 61

Description.—Mature larva: length 7.8 mm, width 2.4 mm; head
not restricted posterior to eye prominence, ratio of width of head at
eye prominence to width at neck 1.0. Other characters as given in
keys to genera and subgenera.

Instar no. ?: length 4.5-5.2 mm, mean 4.96 mm; width 1.4-2.0 mm,
mean 1.68 mm; differ from mature larvae since they lack the stalked
prothoracic spiracle.

Biotocy.—Larvae of this species have been collected only from
under horse manure.

MATERIAL EXAMINED.—Ontario: Marmora, June 19, 1952, J. R.
Vockeroth, 2 puparia, larvae collected from under horse dung in
woods, CNC. Virginia: Alexandria, June 24, 1951, W. W. Wirth,
15 larvae (early instar), from under horse dung, USNM.

Sargus (Sargus) decorus Say, 1824
Ficures 55, 58, 59

Description.—Mature larva: length 4.0-4.2 mm, mean 4.1 mm;
width 3.0-3.3 mm, mean 3.13 mm; width of head at eye prominence
0.81—0.94 mm, mean 0.87; width of head at neck 0.48-0.59 mm, mean
0.52 mm, mean ratio of width of head at eye prominence to width at
neck 1.67. Other characters as given in key to species.

BioLoay.—Larvae of this species have been collected from rotting
leaves and cow manure.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

MatTERIAL EXAMINED.—Saskatchewan: Saskatoon, Aug. 1, 1949,
A. R. Brooks, 14 puparia from cow manure, CNC. Washington:
Seattle, Sept. 15, 1942, lot no. 42-11571, det. by C. T. Greene, 4
larvae, 5 puparia from rotting leaves, USNM.

Sargus (Sargus) bipunctatus Scopoli, 1763
Ficures 48, 54, 57

Description.—Mature larvae: length 9 mm; width 3mm. Other
characters as given in key to species.

Instar no. ?: length 6.5 mm; width 2.4mm. Differs from mature
larva since it lacks the vittae.

Brotocy.—Larvae of this species have been collected from cow
manure. Malloch (1917) reported the larvae as being very sluggish.

MATERIAL EXAMINED.—Maryland: Braddock Heights, Oct. 1,
1921, A. N. Caudell Collection, 1 larva (early instar), USNM.

Discussion.—All data regarding description of the mature larva
and notes on its biology have been taken from Malloch.

Sargus (Sargus) cuprarius (Linnaeus), 1758
Ficures 17, 62-64

DescripTION.—Instar no. ?: length 6.5-7.4 mm, mean 7.04 mm;
width 1.7-2.0 mm, mean 1.84mm. Similar to S. decorus in this stage
but much smaller.

Brotogy.—Larvae of this species have been collected from under
cow manure.

MATERIAL EXAMINED.— Nebraska: Dunbar, Sept. 13, 1950, W. W.
Wirth, 39 larvae from cow manure (early instars), USNM.

Genus Ptecticus Loew, 1855

Of the two species listed by James (1960) for this genus, only the
larvae of P. trivittatus Say have been collected north of Mexico.
GENERIC CHARACTERS.—As given in key to genera.

Ptecticus trivittatus (Say), 1829
Figures 438, 47, 50, 51

Description.—Mature larva: length 11.2-13.2 mm, mean 12.68
mm; width 3.5-3.9 mm, mean 3.76 mm; apical segment with 2 setae
at each corner of spiracular opening; no anal spines present; other
characters as given in key to species.

Antepenultimate instar: length 12.2-15.1 mm, mean 13.67 mm;
width 3.0-4.5 mm, mean 3.67 mm; larvae testaceous, no visible pat-
tern; body setae short, partially hidden by fine pubescence that covers
body; anal spines present.

No. 3569 SOLDIER FLY LARVAE—McFADDEN Zo

Biotoay.—Larvae of this species have been collected from a corn
compost pile, from decaying tomatoes, decaying garbage, rotting
paper, decaying fruit, and from watermelon rinds. Larvae have also
been collected from a fungus (Laetiporus speciosus).

MATERIAL EXAMINED.—Georgia: Silver Lake, Bradley, 7 larvae
from decaying garbage and rotting paper, CU. Illinois: Urbana,
Aug. 3, 1942, H. H. Ross, 14 larvae, INHS. Maryland: Cabin John
Bridge, July 31, 1913, Barber and Shannon, 11 larvae from fungus
(Laetiporus speciosus), USNM. North Carolina: Faison, Aug. 25,
1950, P. O. Richter, 5 larvae from watermelon, NCS; Faison, Oct. 14,
1952, Dogger and Howden, 1 larva from decaying vegetables, NCS;
McCulley’s, Jan. 20, 1951, Weisman, 3 larvae from watermelon rinds,
NCS. Ohio: O.A.E.S., dump, Aug. 22, 1943, 17 larvae from decaying

tomatoes, OSU.
Ptecticus sackeniti Williston, 1885

Description.—Mature larva: length 11 mm; width 4.5 mm.

BroLtocy.—No data available.

MATERIAL EXAMINED.— Mexico: No other data, 2 larvae collected
by A. Herrana, AMNH.

Discusston.—According to W. W. Wirth of the U.S. Department
of Agriculture, this species does not occur in Mexico. The 2 larvae
examined by the author were indistinguishable from the antepenulti-
mate instar of P. trivittatus. The identification is made less creditable
by the fact that both larvae were neither reared nor associated with
adult flies in the collection.

Genus Microchrysa Loew, 1855

James (pers. comm.) lists 2 species for this genus but only the
larvae of Microchrysa polita (Linnaeus) have been collected to date.
GENERIC CHARACTERS.—AS given in key to genera.

Microchrysa polita (Linnaeus), 1758
Ficures 42, 44, 46

Description.—Mature larva: length 5.2-7.1 mm, mean 6.68 mm;
width 1.6-2.0 mm, mean 1.84 mm; width of head at eye prominence
0.45-0.50 mm, mean 0.47 mm; width of head at neck 0.32-0.42 mm,
mean 0.36 mm; mean ratio of width of head at eye prominence to
width at neck 1.32; venter of sixth abdominal segment with the setae
on each side of the sternal patch reduced; anal spines lacking.

Brotoay.—Larvae of this species have been collected from cow
manure and from decaying vegetation.

Material EXAMINED.—Colorado: Woodland Park, Station 4, Aug.
4, 1943, J.A.R. and H. H. Ross, 3 larvae from cattle droppings, UC.
Massachusetts: Amherst, July 10, 1942, M.E. Smith, 15 larvae, 2

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

puparia from decaying vegetation, MSC. New York: Ithaca, July 8,
1915, 4 larvae from cow manure, CU; Ithaca, July 1917, 5 larvae,
3 puparia from decaying rhubarb, CU; Ithaca, July 1919, 9 larvae,
CU. North Carolina: Rowan County, Dec. 14, 1955, C. E. Jernigan,
1 larva from woods trash, NCS.

Genus Merosargus Loew, 1855

Of the 2 species listed by James (pers. comm.) for this genus, only
the larvae of Merosargus caerulifrons (Johnson) have been collected
to date.

GENERIC CHARACTERS.—AS given in key to genera.

Merosargus caerulifrons (Johnson), 1900
Figure 151

DEscrIPTION.—Puparium: length 5.6 mm (head capsule and pro-
thorax lacking) ; width 2.3 mm; head capsule similar to that of Sargus
lucens but differing in key characters.

BroLtocy.—A puparium of this species was collected from debris at
the base of a squirrel’s nest.

MATERIAL EXAMINED.— Maryland: Linnieville, Jan. 1, 1914, R. C.
Shannon, 1 puparium with head capsule detached, USNM.

Subfamily Clitellariiae

A study of both adults and larvae of the subfamilies Cyphomyiinae
and Hermetiinae (James, pers. comm.) has shown that these two taxa
are closely related to the Clitellariinae as defined by me (p. 13). For
this reason, I have combined the Cyphomyiinae with the genus
Adoxomyia to form the tribe Clitellariini and have reduced the sub-
family Hermetiinae to tribal status.

The genus Dieuryneura has not been placed in either tribe due to
a lack of characters, but it appears to be more closely allied to the
Hermetiini than to the Clitellariini.

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

HABITAT OF LARVAE.—All larvae of this subfamily are terrestrial
scavengers and, as such, have been reported from a great variety of
habitats, the most common of which is decaying organic matter.

Tribe Clitellariini

This tribe includes 3 genera: Adoxomyia, Cyphomyia, and Di-
cyphoma. Their range extends from South America through the
southwestern region of the United States. It is interesting to note
that 2 species of Adoxomyia, lata and rustica, have extended their

NO. 3569 SOLDIER FLY LARVAE—McFADDEN Qe

range as far north as Oregon and Washington on the west coast and a
third species, swbulata, is found throughout many of the Eastern
States. Fourteen species have been collected from the southwest as
adults, but the genus Adozomyia, which contains 11 of the 14 species,
is known only from 3 puparia that lack the head capsule.

HABITAT OF LARVAE.—AIl specimens collected to date have been
taken from decaying or wounded cactus. Larvae of an unknown
species of Cyphomyia have been reported as occurring under the bark
of cedar logs in Honduras.

Key to Genera of Clitellariini

1. Head without a distinct notch posterior to eye prominence; dorsal seta on
median line of first abdominal segment shorter than outer 2 setae (fig. 77) ;
all 3 setae set on a line perpendicular to the body axis. Adoxomyia Kertesz

Head with a distinct notch posterior to eye prominence. . ....... 2

2. Median seta on dorsum of first abdominal segment longer than setae on either

side of it (fig. 84); all 3 setae set on a line diagonal to the body axis.
Cyphomyia Wiedemann
Median seta no longer than setae on either side of it (fig. 78); all 3 setae set
on a line diagonal to the body axis. . . ... .. . Dicyphoma James

Genus Cyphomyia Wiedemann, 1819

James (pers. comm.) recognizes 2 species in this genus: Cyphomyia
bicarinata Williston and C. marginata Loew. Larvae have been col-
lected for both species, but because the specimens of CO. bicarinata are
an early instar, pin mounted, and in poor condition, I have substituted
the description of a neotropical species, C. pilosissima Gerstaecker.

Cyphomyia marginata Loew, 1866
Ficures 152-154

Description.—Puparium: length 20.1 mm; width 2.9 mm; other
characters as given in key to genera.

Biotocy.—Larvae of this species have been collected from papaya
in Florida.

MaTERIAL EXAMINED.—Florida: Miami, July 11, 1917, T. E.
Snyder, 1 puparium, USNM.

Cyphomyia pilosissima Gerstaecker, 1857
Figures 83-85

Description.—Mature larva: length 21 mm; width 5 mm; dorsally
the anterior and posterior margins of the thoracic and of the first 7
abdominal segments somewhat elevated with low transverse ridges oc-
cupying approximately the anterior and posterior fourth or fifth of the
segment; ventrally these areas are only slightly elevated.

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Brotocy.—Larvae of C. bicarinata as well as larvae of C. pilosissima
have been taken from wounds in cacti, C. bicarinata from prickly pear
(Opuntia species), and C. pilosissuma from maguey (Agave species).
No other data are available.

Genus Dicyphoma James, 1937

Only 1 species, Dicyphoma schaefferr (Coquillett), occurs in the
Nearctic area. It is restricted to the southwestern states, where it
has been taken from wounded and decaying cacti.

GENERIC CHARACTERS.—As given in key to genera.

Dicy phoma schaefferi (Coquillett), 1904
Fiaures 78, 81, 82

Description.—Mature larva: length 14.0-15.8 mm, mean 15.2
mm; width 3.6—4.1 mm, mean 3.87 mm.

Brotoay.—Larvae have been collected from Opuntia (Platyopuntia)
species and Myrtillocactus geometrizans in Mexico.

MATERIAL EXAMINED.—Mexico: 54 miles north of San Luis Potosi,
S.L.P., Aug. 4, 1960, Ryckman, Ryckman and Christianson, ADM
61, 2 larvae, WSU; 8 miles south of Aguascalientes, June 19, 1960,
ADM 46, Ryckman, Ryckman and Christianson, 1 larva WSU.
Texas: El Paso, Aug. 12, 1908, F. C. Pratt, 1 headless puparium,
USNM.

Discussion.—Except for the key characters, there appears to be
little difference between the larva of D. schaefferi and that of C.
pilosissima.

Genus Adoxomyia Bezzi, 1908

James (pers. comm.) lists 11 species for this genus, all of which
(except A. subulata, A. lata, and A. rustica) are confined to the south-
western and western states. Unfortunately, this group of species is
represented by only 3 puparia, all of which lack the head capsule;
however, in an attempt to present a complete analysis of the generic
characters, | have included the description of an Indian species,
Adoxomyia heminopla (Wiedemann).

GENERIC CHARACTERS.—As given in key to genera.

Adoxomyia rustica (Osten Sacken), 1877
Figures 155, 156
DescripTion.—Puparium: length 19.6 mm (head capsule missing) ;
width 2.8mm. Other characters as given in key to genera.

Brotocy.—No data available for A. rustica but larvae of A. clari-
pennis have been collected from decaying cacti.

ee

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 29

MarTerIAL EXAMINED.—Washington: Tampico, Mar. 1, 1931, A. RB.
Rolfs, 1 puparium, USNM.

Discussion.—In addition to the specimens described above, the
author has examined 2 puparia of A. claripennis that were collected
in Pima County, Ariz., Dec. 27, 1951, by Ryckman, Ames, and Ara-
kawa USNM. Both specimens lacked the head capsule but were in
agreement with other key characters of A. rustica.

Adoxomyia heminopla (Wiedemann), 1819
Fiaures 74, 77, 80

DescripTion.—Mature larva: the characters that appear to be of
generic significance are given in the key to genera. Specific characters
are unimportant since the species does not occur within the geographic
range covered in this paper.

Brotoey.—Larvae of this species were collected from Kaempferia
species, a tuberous-rooted plant. It has also been reported from
papaya by Brunetti (1923).

MATERIAL EXAMINED.—India: Calcutta, May 26, 1955, HO-19477,
lot no. 55-11257, 10 larvae, 5 puparia, USNM.

Tribe Hermetiini

Composed of the single genus Hermetia, this tribe is represented
in North America by 8 species. Seven of them appear to be re-
stricted to the southwestern United States. One species, Hermetia
ilucens (Linnaeus), has been recorded from as far north and east as
Maryland and Delaware.

TRIBAL CHARACTERS.—As given in key to subfamilies.

Hasirat or LARVAE.—Hermetiine larvae are terrestrial scavengers.
They have been reported from excrement (mammal and avian),
decaying vegetable matter, wax in beehives, and a cadaver (Dunn,
1916). H. illucens has been involved in the human disease en-
teric myiasis (James, 1947).

Key to Species of Hermetia Latrielle

1. Lateral and transverse setae (dorsal and ventral) very prominent; body
segments with dense yellow pubescence especially on apical segment and
Onvlaternl Mienrm users. gs eek st ee eee tine illucens (Linnaeus)

Lateral and transverse setae not prominent; dense yellow pubescence lacking
or, if present on dorsum of thorax, then divided into 2 patches by a bare

ae diane lien iy. vel RAN ee Bier ies. Slee ete ek ed bei Lt 2
2. Posteroventral region of head with coarse yellow pubescence; 2 setae in
prothoracic leg group. . . ; . . . species

Bib on venter of head lacking (fig. 108); more than 2 setae in prothoracic
FO VeTOUD ae meena Ms TNE, MERRY Sy ME eS tig

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

3. Labrum barely extending beyond ocular lobes; point not drawn out, blunt

APWEATATECS Se 6 ek ce eye. os) othe te) or coe ae nae concinna Williston
Labrum extending beyond ocular lobes, end portion drawn out into a taper-
AT PUPOUN EIS Se sea ih ere 6! Te oo! er weeny nae oe aurata Bellardi

Genus Hermetia Latreille, 1804

I have seen larvae or puparia of 4 of the 8 species listed by James
(pers. comm.) for this genus. One species, however, remains unnamed.

Hermetia illucens (Linnaeus), 1758
Fiaures 21, 65, 75, 79

Descriptions.—Mature larva: length 14.5-21.5 mm, mean 18.6
mm; width 4.8-6.5 mm, mean 5.8 mm; head and body segments
reddish brown; spiracles present on lateral margin of segments 3-10,
accompanied by a small papilla on segments 2-5; anus not armed
with spines; no transverse series of short spines.

Penultimate instar: length 19.0-25.5 mm, mean 18.6 mm; width
4.0-5.5 mm, mean 4.8 mm; body segments white to creamy yellow,
head and prothoracic spiracles almost black; anus armed with spines;
all abdominal segments with a transverse series of darkly pigmented
spines on venter near anterior margin.

Brotoay.—Larvae of H. illucens are terrestrial scavengers. They
have been reported from beehives (Copello, 1926), decaying crabs
(Buxton and Hopkins, 1927), decaying fruits and vegetables, catsup,
animal cadavers, waste material in beehives, and from privies (James,
1947). May (1961) reported that large numbers of H. tllucens,
living and dead larvae, pupae, and some dead adults, were found
in covered crocks containing tuna remains preserved in a solution of
10 percent formaldehyde. Furman, Young, and Catts (1959) have
shown that large numbers of larvae of H. illucens will prevent develop-
ment of larvae of the housefly Musca domestica (Linnaeus) in poultry
manure.

A detailed life history has been presented by May (1961) in which
she reports on the time required for the egg, larval and pupal stadia,
number of instars, and the morphological changes associated with
each of the 6 larval instars.

In North America this species appears to be limited to one genera-
tion per year with the adults emerging in the spring after a winter of
dormancy. Copello (1926) mentioned that mating takes place during
flight and later the females oviposit at the edge of decaying organic
material.

MATERIAL EXAMINED.—Arkansas: Fayetteville, Sept. 27, 1927,
W. J. Baerg, 3 larvae from kraut. California: 10 miles east of

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 31

Rosario Cirio, June 29, 1938, Ross and Michelbacher, 12 larvae.
Delaware: Magnolia, October 1958, P.P.B., 10 larvae from polluted
water, UD; Primehood Neck, M.S.C., 8 larvae from lima bean cull
pile, UD; Thompsonville, Oct. 24, 1957, M.S.C., 1 larva from lima
bean refuse pile, UD. Florida: Collection 1884 from USNM, 5
larvae, 1 pupa, USNM. Georgia: Bissel, Sept. 15, 1938, 1 larva
from rotten potatoes; Camp Stewart, Sept. 21, 1944, E. R. Willis,
8 larvae taken from pit latrine; Camp Stewart, Dec. 6, 1944,
E. R. Willis, 31 larvae taken from pit latrine. Louisiana: Baton
Rouge, Sept. 30, 1959, H. V. Daley, 13 larvae from chicken manure,
LSU. Maryland: Beltsville, Aug. 9, 1957, J. C. Hwang, 4 larvae
from turkey manure, USNM. Missouri: St. Louis, July 20, 1960,
C. W. Robinette, 4 larvae, UMO. New Mexico: Roswell, Aug. 30,
1957, 7 larvae taken from worm bed. North Carolina: Clay County,
Oct. 15, 1954, extension service, 5 larvae from silo, NCSC; Clayton
County, July 5, 1959, W. Brooks, 6 larvae from corn pile, NCSC;
Faison, Aug. 14, 1952, Dogger and Howden, 9 larvae from rotting
pumpkin, NCSC; Fuguay, Sept. 12, 1950, 5 larvae from manure,
NCSC; Long Beach, Sept. 5, 1951, 1 larva from refuse, NCSC;
Onslow County, Sept. 18, 1956, H. E. Scott, 2 larvae from poultry
litter, NCSC; Raleigh, Aug. 4, 1941, 8 larvae from garbage, NCSC;
Raleigh, July 1, 1953, D. M. Weisman, 3 larvae from ensilage, NCSC;
Raleigh, Sept. 4, 1954, W. Spink, 5 larvae, NCSC; Rockingham,
Oct. 25, 1955, M. Farrier, 4 larvae from under caged hens, NCSC.
South Carolina: Clemson, Nov. 18, 1959, Schroeder and Skelton,
2 larvae from lab culture (dung), CC; Clemson, Nov. 19, 1959, 1
larva from manure, CC; Clemson, 30 larvae (no data), CC; Wild
Cat Creek, Six Mile, Sept. 29, 1959, D. H. Peterson, 1 larva, CC.
Tennessee: Knoxville, Harwood, 1 pupal case. Washington: Ev-
erett, June 22, 1957, F. Johansen, 10 larvae from soy beans in freight
car, origin unknown, WSU.

Hermetia concinna Williston, 1900
Ficures 66, 69, 72

Description.—Puparium: length 17.3 mm; width 4.8 mm; head
and body segments testaceous; no spines in transverse series on
any segments.

Brotogy.—The puparium studied was taken from decayed sotol
(Dasylirion species). No other data are available.

MATERIAL EXAMINED.—Arizona: Sierita Mountains, 30 miles
southwest of Tucson, Nov. 27, 1913, E. A. Schwarz, 1 larva, 1
puparium, USNM.

oo PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Hermetia species

Ficures 68, 70, 76

Description.—Mature larva: length 17.5 mm; width 5.5 mm;
head and body segments orange yellow, head more darkly pigmented
at distal end; eyespots not distinct; transverse row of small spines
present on segments 1-7 near anterior margin.

BroLtogy.—Same as noted for H. concinna.

MaTERIAL EXAMINED.—Three larvae with same data as for H.
concinna.

Discusston.—There may be some doubt as to the validity of
this species because the specimens used to describe it were found
with those of H. concinna.

Hermetia aurata Bellardi, 1859
Ficures 67, 71, 73

DescripTion.—Puparium: length 17.8-22.3 mm, mean 20.05 mm;
width 5.1-6.0 mm, mean 5.55 mm. Other characters as given in
key to species.

Brotogy.—Larvae have been collected from prickly pear (Opuntia
occidentalis).

MATERIAL EXAMINED.—California: San Dimas Canyon, Pomona,
Los Angeles County, Dec. 4, 1960, A.D.M. 72, Ryckman and Olsen,
two puparia, WSU.

Genus Dieuryneura James, 1937

Dieuryneura obscura (Coquillett), 1902
Figures 16, 24, 27, 32

Description.—Mature larva: length 14.9 mm; width 4.0-4.3 mm,
mean 4.1 mm; head and body segments dark brown.

Biotoay.—Larvae of this species were collected from a decaying
sotol plant stem (Dasylirion species).

MATERIAL EXAMINED.—Texas: 12 miles north of Presidio, Apr. 18,
1952, 52-6319—Presidio-3119-L, 1 larva, 11 puparia, USNM.

Subfamily Stratiomyinae

This subfamily is composed of the species of Stratiomyidae, whose
larvae are aquatic or semiaquatic. ‘The larvae vary in size from a
few millimeters to 50 millimeters. One commonly finds larvae of
this subfamily as representatives of this family in general collections
of immature stages. This can be explained best by the fact that the
aquatic species are more numerous and, therefore, easier to locate
than the terrestrial species.

SS .aaaaaaSaQQSaQrF wee _

NO. 3669 SOLDIER FLY LARVAE—McFADDEN 33

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

HaBITAT OF LARVAE.—Stratiomyine larvae have been collected
from almost every conceivable type of aquatic environment, ranging
from hot springs and fast-flowimg brooks to ocean shores. Although
larvae in the genus Sératiomys are at home in water depths of several
feet, larvae in most of the remaining genera prefer to remain hidden
in naturally occurring vegetation along the shore or margin of the
water. Quite frequently these larvae are found in and under shore
debris such as boards, logs, cans, cardboard boxes, weeds, and other
types of decaying organic matter.

Key to Genera of Stratiomyinae

1. Seventh abdominal segment of larva with curved sclerotized hooks on venter

(ge 22 OS eas s Pa
Seventh abdominal Scoyaesaly sation ee pcieenred hooker on anes 2) 2
2. Antenna located at apex of ocular lobe (fig. 126) ... . REP elec eee

Antenna not located at apex of ocular lobe . Galoparynhns iene (in part)

3. Integument of larva covered with minute, peltate scales (fig. 91).
Odontomyia Meigen, subgenus Catasina
Integument lacking peltate scales. ..... . . .Stratiomys Geoffroy
4. Sclerotized hooks present on venter of each of the first 7 abdominal segments,
those on the seventh larger than the others. . . . Aochletus Osten Sacken
Sclerotized hooks absent from each of the first 6 segments; those on the
SEVEDUNOPLESCMU wwe tes we Sepae in cic ce Si) cy esl ueh Sr bah es keds. oo va cistasne: okt SD
5. Hydrofuge setae on last abdominal segment attaching to 2-lobed structures
on lower lip of spiracular cleft (fig. 88); prothoracic spriacles located at
anterior corner of that segment... . . Myxosargus Brauer
Hydrofuge setae attaching to straight dae of fore lip, lobed structures
absent; prothoracic spiracle not located in anterior corner of prothorax . . 6

6. Antenna dorsal, not at apex of ocular lobe; prothoracic spiracles elevated or

stalked sd. << ts PIMOROL Se
Antenna at apex of eee one: Gareiianae ppiearies wounee elevated nor
SUMO et eh tee ote CES OER Sa Vesh o) (ase) ceveR A ate tee ATs ule aki et Bt oa ands

7. Apical segment with lateral margin straight; distance from antenna to eye
prominence greater than twice length of antenna.
Caloparyphus James (in part)
Lateral margin of apical segment tapering basally toward median line; distance
from antenna to eye prominence approximately equal to length of an-
tenMAN Eee hen nish eh os feiouore wits oreo Uparypnus)» Gerstaccker
8. Venter of sixth and seventh abdominal segments with sclerotized hooks . .9
Venter of only seventh abdominal segment with sclerotized hooks.
Odontomyia Meigen, subgenus Odontomyiina
9. Body segments with broad dorsal vittae or body segments with fine white
pubescence or multiple hooks.
Odontomyia Meigen, subgenus Odontomyia
Without the above combination of characters . . . Hedriodiscus Enderlein

221-533—67——3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Genus Stratiomys Geoffroy, 1762

This is one of the largest genera of soldier flies that occurs in
North America. Of the 21 species listed by James (pers. comm.), I
have seen associated larvae or puparia of 7 species.

GENERIC CHARACTERS.—As given in key to species

Key to Species of Stratiomys Geoffroy

(Although associated puparia of 7 species were available, I was able to separate
only 1 species (S. discaloides Curran) from the remaining 6).

Length of last abdominal segment approximately 2 times the basal width of

Phat rSeeTseN bir ous! Gion muse hia9 <o 6 Uieet eatenn rete om earn ies discaloides Curran
Length of last abdominal segment greater than 2 times the basal width of
thabvseemembi. @ <) ogee: res ee eee adelpha Steyskal, badia Walker,

barbata Loew, meigenii Wiedemann, norma Wiedemann, normula Loew.

Stratiomys discaloides Curran, 1922
Ficure 119

Duscription.—Puparium: length 31 mm, width 6.8 mm; head and
body segments dark brown; faint dark markings at basal margin of
body segments. Other characters as given in key to species.

Brotocy.—No data are available for this species.

MATERIAL EXAMINED.—Montana: Harlan Gulch, Rav. County,
Jan. 9, 1932, C. B. Philip, 1 broken puparium, WSU.

Stratiomys species
Figures 20, 105-109

Description.—Puparium: length 31.3-41.0 mm; width 4.9-6.0 mm;
color varying from grey brown to dark brown; markings same as for
S. discaloides. Other characters as given in key to species.

Brotoay.—See biology section in this paper.

MATERIAL EXAMINED.—Stratiomys adelpha Steyskal: New York:
Ithaca, Decker Pond, Mar. 27, 1950, H. H. Schwardt, 1 puparium,
WSU. Saskatchewan: Watson, May 15, 1956, M. E. Taylor, 5
puparia, MWM._ Stratiomys badia Walker: Alberta: 4 miles north
of Devon, May 16, 1961, G. Pritchard, 1 puparium, MWM; Banff
National Park, Mt. Ishbel, June 14, 1960, Ball, Madge and McFadden,
1 larva, 1 puparium from under moss, elevation approximately 6000
ft., MWM. _ Stratiomys barbata Loew: Manitoba: Churchill, July 16,
1949, 2 puparia, CNC. Alberta: Banff National Park, Mt. Ishbel,
June 14, 1960, Ball, Madge and McFadden, 1 puparium, elevation
approximately 6000 ft., MWM. Stratiomys meigeni Wiedemann:
Kansas: Manhatten, Feb. 8, 1936, H. H. Schwardt, 2 puparia, WSU.
Stratiomys norma Wiedemann: Wisconsin: T9N Knapps Creek,
Richland County, R2W, June 10, 1954, R. H. Jones, 4 puparia, WSU.

No. 3569 SOLDIER FLY LARVAE—McFADDEN 35

Stratiomys normula Loew: Louisiana: Baton Rouge (?), 2 puparia,
LSU.

Genus Myxosargus Brauer, 1882

Four species of Myzosargus occur in North America but only the
puparium of Myxosargus nigricornis Green is known.
GENERIC CHARACTERS.—As given in key to genera.

Myxosargus nigricornis Green, 1918
FiaureEs 88, 92, 93

Descriprion.—Puparium: length 6.7-7.0 mm (less head and pro-
thorax), mean 6.85 mm; width 2.0-2.1 mm, mean 2.05 mm.

Brotogy.—No data are available for this species.

MarTeRIaL EXAMINED.—Missouri: 5 miles north of Vichy, June 15,
1955, P. J. Spangler, 2 puparia, WSU.

Genus Euparyphus Gerstaecker, 1857

James (pers. comm.) divided this genus into 2 subgenera as follows:
Euparyphus (11 species) and Aochletus (3 species). Of the 11 species
listed for Huparyphus, only the larvae of E. limbrocutris Adams have
been collected. Aochletus has been elevated to full generic status.

GmNERIC CHARACTERS.—As given in key to genera.

Euparyphus limbrocutris Adams, 1903
Ficures 94, 95, 99

Description.—Mature larva: length 10.6-11.2 mm, mean 10.9 mm;
width 2.3-2.5 mm, mean 2.4 mm; prothoracic spiracles located on a
medial-lateral projection and extending at least halfway to the
dorsomedian line of that segment.

Penultimate instar: length 9.2-10.6 mm, mean 9.9 mm; width
2.0-2.1 mm, mean 2.05 mm; differs from mature larva since pro-
thoracic spiracles are elevated as high as their basal diameter.

Biotogy.—No data are available for this species.

MATERIAL EXAMINED.—Washington: O’Sullivan Dam, Grant
County, May 7, 1955, J. A. Quist, 2 mature and 2 penultimate instar
larvae, WSU.

Genus Aochletus Osten Sacken, 1886

On the basis of adult characters, James (pers. comm.) considered
Aochletus to be a subgenus of Euparyphus; however, the difference or
degree of difference between the larvae of Aochletus and EHuparyphus
is so great that I feel full generic status should be given to Aochletus.

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Of the 3 species listed by James (pers. comm.) for this taxon, larvae
have been collected for the following 2: A. cinctus Osten Sacken
and A. brevicornis Loew.

GENERIC CHARACTERS.—As given in key to genera.

Key to Species of Aochletus Osten Sacken

Prothoracic spiracles distinctly stalked. . . - +--+: cinctus Osten Sacken
Prothoracic spiracles almost flush with integument. is cue brevicornis Loew

Aochletus cinctus Osten Sacken, 1866

Description.—Puparium: length 9.5-10.5 mm, mean 10.0 mm;
width 2.7-2.8 mm, mean 2.73 mm. Other characters as given in
key to species.

Bro.ocy.—No data are available for this species.

MATERIAL EXAMINED.—California: Topanga Canyon, Los Angeles
County, May 10, 1953, 3 puparia, WSU; Sespe Creek, Ventura
County, June 15, 1948, W. W. Wirth, 1 puparium, USNM.

Aochletus brevicornis Loew, 1866
Froeurss 96, 97, 101

Description.—Instar no. ?: length 4.5-5.3 mm, mean 4.93 mm;
width 1.3-1.7 mm, mean 1.50 mm; other characters as given in key to
species.

Biotocy.—Larvae have been collected from springs in Yellowstone
National Park. No mention is given regarding the type of spring
other than the location.

MATERIAL EXAMINED.—Wyoming: Mammoth, Yellowstone Na-
tional Park, Jan. 25, 1956, J. R. Murphy, 3 early instar larave from
a cavern spring, WSU; Mammoth, Yellowstone National Park,
June 20, 1956, J. R. Murphy, 6 larvae from hillside springs, WSU.

Genus Caloparyphus James, 1939

On the basis of male genitalia, both James (pers. comm.) and Quist
(Thesis) consider Caloparyphus to merit generic status. The larvae,
however, do not seem to corroborate this. In fact, it is rather diffi-
cult to distinguish between the larvae of Caloparyphus and those of
Euparyphus (see key to genera, p. 33). This seems to suggest that
perhaps Caloparyphus should be returned to its former status as a
subgenus of Huparyphus.

James (pers. comm.) lists 11 species for this genus in North America.
Associated larvae or puparia have been collected for 5 species, in-
cluding 1 unidentified species.

GrNnpRIC CHARACTERS.—As given im key to genera.

ee EEE ______eeeee

i

NO. 3569 SOLDIER FLY LARVAE——McFADDEN 37

Key to Species of Caloparyphus James

1. Venter of seventh abdominal segment without strong sclerotized hooks.
species
Venter of seventh abdominal segment with strong sclerotized hooks. . . 2
2. Sclerotized hooks half the length of seventh abdominal segment.
amplus (Coquillett)
Sclerotized hooks less than half the length of the seventh abdominal

Sea) Soro ot Bale WoL ce Lado Lol tea oe aoe oe ane
3. Prothoracic spiracle oval shaped, diameter at widest point twice that of the
elevation; 4 setae in mesothoracic leg group... .. major (Hine)

Prothoracic spiracle varying in shape but elevation approximately equal to
basal diameter; number of setae in mesothoracic leg group other than4 . 4

4. Three setae in mesothoracic leg group... .. . . tetraspilus (Loew)
Five setae in mesothoracic leg group... . . . crotchi (Osten Sacken)

Caloparyphus species
Ficures 112, 115, 118

Descrietion.—Puparium: length 12.4-13.4 mm, mean 12.88 mm;
width 2.5-3.2 mm, mean 2.90 mm; other characters as given in keys
to genera and species.

Briotoay.—Larvae of this species were collected from a sphagnum
bog located part way up a mountain (elevation approximately
6000 ft.).

MATERIAL EXAMINED.—Alberta: Banff National Park, Mt. Ishbel,
Apr. 14, 1960, Ball, Madge and McFadden, 5 puparia, MWM.

Caloparyphus amplus (Coquillett), 1902
Ficgurss 122, 127, 128, 131

Description.—Mature larva: length 7 mm or under; width 1.0-
1.4 mm, mean 1.2 mm; dorsum of body segments with short blunt
setae; other characters as given in key to species.

Brotoay.—No data are available for this species.

MATERIAL EXAMINED.—California: Sespe Creek, Ventura County,
June 15, 1948, W. W. Wirth, 20 larvae, USNM.

Discusstion.—Accurate measurements of length were unobtainable
due to improper preservation of the specimens.

Caloparyphus major (Hine), 1901
Ficures 111, 114, 117

Description.—Mature larva: length 9.6—12.1 mm, mean 10.94 mm;
width 2.7-2.9 mm, mean 2.8 mm; other characters as given in key to
species.

Brotocy.—Larvae of this species were collected from moss on a
large floating board in a cold, spring-fed roadside pool.

MATERIAL EXAMINED.—British Columbia: Pole no. 187/18, Route 3,
June 9, 1960, Ball, Madge, and McFadden, 5 larvae, 3 puparia, MWM.

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Caloparyphus tetraspilus (Loew), 1866
Fiaeures 110, 113, 116

DescripTion.—Mature larva: length 9.5-11.0 mm, mean 10.25 mm;
width 1.8-2.3 mm, mean 2.05 mm; other characters as given in key
to species

BrioLtocy.—Larvae of this species were collected on a sandy lake
beach under decaying vegetation and other debris at the water’s edge.

MATERIAL EXAMINED.—Alberta: Dilberry Lake, 54°34’30’’ N lat.,
110°60’45’’ W long., June 5, 1960, Ball, Madge and McFadden, 2 lar-
vae, MWM.

Caloparyphus crotchi (Osten Sacken), 1877
Fiaurss 125, 129, 132

Description.—Puparium: length (specimen broken, impossible to
measure accurately but close to 15 mm); width 3.4 mm; other charac-
ters as given in key to species.

Brotocy.—No data are available for this species.

MATERIAL EXAMINED.—Colorado: Fort Collins, Aug. 4, 1910, 1
puparium, WSU.

Genus Hedriodiscus Enderlein, 1914

James (pers. comm.) lists 7 species for this genus in North America.
Only the larva of H. vertebratus (Say) is known.

Hedriodiscus vertebratus (Say), 1824
FIGURES 87, 91

DescripTion.—Puparium: length 19 mm, width 2.7 mm; other
characters as given in key to genera

Brotogy.—James (pers. comm.) reported that larvae of this species
live among floating vegetation in smallstreams. They feed on micro-
organisms, algae, and the soft parts of plants.

MATERIAL EXAMINED.—Locality (?); July 1, 1938, larva from weedy
lake, 1 puparium, WSU.

Genus Odontomyia Meigen, 1803

James (pers. comm.) has divided this taxon into 3 subgenera as fol-
lows: Catasina (9 species), Odontomyiina (7 species) and Odontomyia
(14 species). The character combinations of the immature stages
seem to support this classification.

NO. 2569 SOLDIER FLY LARVAE—McFADDEN 39
Subgenus Catasina Enderlein, 1914

Odontomyia (Catasina) pubescens (Day), 1882
Fiaures 86, 89, 90

Description.—Puparium: length 13.8-17.0 mm, mean 15.80 mm;
width 2.7-3.6 mm, mean 3.1 mm; other characters as given in key to
genera.

Biotoay.—Larvae of this species have been collected from moss-
covered logs (in boggy areas), from under stones at water’s edge, and
from the margins of marshy areas.

MATERIAL EXAMINED.—Alberta: Beaverhills Lake near Tofield,
May 21, 1960, G. E. Ball, 1 puparium, MWM;; Flatbush, May 12,
1960, M. W. McFadden, 1 larva, 1 puparium, MWM;; Dilberry Lake,
54°34’30” N lat., 110°60’45’’ W long., June 5, 1960, Ball, Madge, and
McFadden, 1 puparium, MWM. Saskatchewan: Saskatoon, May
15, 1949, A. R. Brooks, 1 puparium, CNC.

Subgenus Odontomyiina Enderlein, 1930

Odontomyia (Odontomyiina) virgo (Wiedemann), 1830
Fiaures 98, 102

Description.—Puparium: length 15 mm (less head and prothorax) ;
width 3.50 mm; 4 white vittae on dorsal surface of body segments,
inner 2 narrow; penultimate segment half as long as apical segment;
other characters as given in key to genera.

Brotoey.—Larvae of this species were collected from shore debris at
the edge of a small pond.

MATERIAL EXAMINED.—Alberta: 4 miles north of Devon, May
10, 1961, M. W. McFadden, 1 puparium, 1 early instar larva, MWM.

Subgenus Odontomyia Meigen, 1803

Key to Species of Odontomyia Meigen

1. Venter of sixth and seventh abdominal segments with multiple hooks.
occidentalis James
Venter of sixth and seventh abdominal segments with a single pair of hooks . 2
2. Dorsum with 2 broad vittae extending length of body but dividing into 4
Victae on apicalises mente (ikea nh ee PANS ER cincta Olivier
Vittae in different pattern; body segments with fine white pubescence.
communis James

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Odontomyia (Odontomyia) occidentalis James, 1936
FiaurREs 135, 136, 138

Descriprion.—Puparium: length 20 mm; width 3.5 mm; other
characters as given in key to species.

Biotocgy.—Larvae of this species have been collected from hot
springs.

MATERIAL EXAMINED.—Sleeping Child Hot Springs, reared June
28, 1930, 2 puparia, WSU.

Discussion.—The mensural data given above were taken directly
from the puparia. These specimens had the apical segments turned
up in the typical manner making accurate measurements an
impossibility.

Odontomyia (Odontomyia) cincta Olivier, 1811
FiaureEs 126, 130, 133

DescripTion.—Mature larva: length 18.5-20.0 mm, mean 19.25
mm; width 3.5-3.9 mm, mean 3.7 mm; other characters as given
in key to species.

BroLtogy.—Larvae of this species have been collected from a peat
bog and from a pool of unidentified type.

MATERIAL EXAMINED.—Ohio: Delaware County, July 29, 1940,
1 larva, OSU. Wisconsin: Dane County, University of Wisconsin
Arboretum, May 22, 1954, R. H. Jones, 1 puparium, WSU.

Odontomyia (Odontomyia) communis James, 1939

Ficures 140, 142

DescripTion.—Mature larva: length 19-21 mm, mean 20 mm;
width 2.9-3.6 mm, mean 3.25 mm; other characters as given in key
to species.

Biotocy.—Larvae of this species have been collected from 5
percent saline water in Death Valley.

MATERIAL EXAMINED.—California: Bad Water, Death Valley,
May 1, 1958, D. P. Furman, 2 larvae, WSU.

Subfamily Nemotelinae

James (pers. comm.) placed the genus Nemotelus in the Stratio-
myinae solely on the basis of adult characters; however, when both
adult and larval characters are taken together and compared with
similar characters of other members of that subfamily, the differences
observed are great enough to warrant separation. For this reason
I have erected the new subfamily Nemotelinae.

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 41

Genus Nemotelus Geoffroy, 1762

In North America this genus contains 33 species that are divided
into 2 subgenera: Nemotelus (12 species) and Camptopelta (21 species).
I have seen puparia of 2 species of Camptopelta and larvae of a single
species of Nemotelus.

GENERIC CHARACTERS.—As given in key to genera.

Subgenus Nemotelus Geoffroy, 1762

The specimens representing this taxon consisted of 3 headless
puparia in poor condition.

Nemotelus (Nemotelus) kansensis Adams, 1903
Figures 120, 123

Description.—Puparium: length 7.2 mm (less head and prothor-
ax); width 2.6 mm

Biotocy.—Larvae of this species were collected from a salt spring
(probably from the margin)

MatTERIAL EXAMINED.—Missouri: Petersburg, June 1, 1955 aloud.
Spangler, 3 puparia, WSU.

Discussion.—The pattern on the dorsum of the body segments and
the arrangement and placement of setae both agree very closely with
that of Nemotelus canadensis Loew

Subgenus Camptopelta Williston, 1917
Key to Species of Camptopelta Williston

Larva less than 4 mm in length; dorsum of body segments lacking vittae and

PACS Re Met Le AeA. TE Seed aids vate ee eee cn) ls wes Ey centralis Hanson
Larva 5 mm or more in length; dorsum of body segments with vittae and plaques
Bei 2 ee ie ae Ae hak Fao RAE be canadensis Loew

Nemotelus (Camptopelta) centralis Hanson, 1958
Ficures 100, 103, 104

Description.—Mature larva: length 3.5-3.7 mm, mean 3.6 mm;
width 0.8-1.0 mm, mean 0.9 mm; head light brown, body white;
prothoracic spiracles dark.

Biotogy.—No data are available for this species.

MatTeRIAL EXAMINED.—Michigan: Cheboygan County, Aug. 17,
1957, W. J. Hanson, 3 larvae, lab reared, KU

Nemotelus (Camptopelta) canadensis Loew, 1863
Ficures 121, 124, 134
Description.—Puparium: length 7.7-8.3 mm, mean 8.0 mm;
width 1.8-2.0 mm, mean 1.9 mm; other characters as given in key
to species.
221-533—66——_4

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Bioutoay.—Larvae of this species have been collected from under
cow manure and rotting vegetation at the edge of highly alkaline
lakes (pH 8.6).

MaTERIAL EXAMINED.—Alberta: Chappice Lake, 18 miles north of
Medicine Hat, June 7, 1960, Ball, Madge and McFadden, 3 puparia,
MWM;; Gooseberry Lake Provincial Park, 9 miles north of Consort,
June 5, 1960, Ball, Madge and McFadden, 8 puparia, MWM.

Subfamily Pachygastrinae

In their recent revision of this subfamily, Kraft and Cook (1961)
have presented an up-to-date resumé of the biology and taxonomy
of both adults and larvae. Keys were presented for distinguishing be-
tween larvae in each of the 5 genera and within the 2 genera Zabrachia
and Hupachygaster.

I have found variation in the measurements given by Kraft and
Cook as compared with those taken by myself. In one species, |
Eupachygaster henshawi Malloch, this variation is one millimeter and
represents an error of 25 percent. In spite of this, I do not believe
that these mensural variations represent specific differences but, —
rather, indicate a small sample that may have been taken from a
limited geographic area. I have placed the measurements given by
Kraft and Cook in brackets and have presented my own measure-
ments including range and mean.

SUBFAMILIAL CHARACTERS.—As given in key to subfamilies.

HasiratT oF LARVAE.—Pachygastrine larvae have been found under |
the bark of both deciduous and coniferous trees. Oviposition usually |
occurs on wounded or dead trees but in either case there must be
enough moisture present in the host for the larvae to be able to obtain |
nourishment. Host trees mentioned by Kraft and Cook are: apple,
quaking aspen, dwarf elm, American elm, cottonwood, hickory, white |
pine, Douglas fir, Engelmann spruce, oa shore pine.

The larvae are gregarious (I have collected approximately 100
larvae beneath the bark of a single log), extremely slow moving, and |
feed on the sap or microorganisms that occur in the moist areas
beneath the bark. Malloch (1917) has suggested that pachygastrine
larvae are predatory on other insect larvae but this has not been
observed.

Key to Genera of Subfamily Pachygastrinae
(modified after Kraft and Cook, 1961)

1. Teeth along anal opening prominent; setae on margin of last segment short, |
no longer than one-fourth width of last segment (fig. 141)....... 2

No prominent teeth along anal opening; setae on margin of last segment long,

at least one-third as long as width of last segment (fig. 140). ...... 3 |

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 43

2. Midventral line of abdominal segment 6 with a round sternal patch, located
anterior to transverse row of setae; 8 or more pairs of conspicuous plaques
along dorsal midline of last segment. .... . Berkshiria Johnson

Sternal patch on midventral line of abdominal segment 6 oval, located between
setae of transverse row; no more than 3 or 4 pairs of conspicuous plaques
along dorsal midline of last segment. ...... Neopachygaster Austen

3. Each thoracic leg group with 2 setae (fig. 140) . . . Eupachygaster Kertesz

Each thoracic leg group with 3 setae (fig. 142). . . SOMO iil aan 4

4. Abdominal segments 1-7 each with 18 setae (fig. 148). . Pachygaster Meigen

Abdominal segments 1-7 each with 20 setae (fig. 142). Zabrachia Coquillett

Genus Berkshiria Johnson, 1914

This genus contains the single species Berkshiria albistylum, the
larva of which has been collected from beneath the bark of deciduous
trees only. As Kraft and Cook (1961) have pointed out, it resembles
Neopachygaster but can be readily distinguished on the basis of the
form of the sternal patch.

GENERIC CHARACTERS.—As given in key to genera.

Berkshiria albistylum Johnson, 1914
Fiaure 142

Description.—Mature larva: length (5.0-7.2 mm) 5.0-5.9 mm,
mean 5.3 mm; width (1.6-2.0 mm) 2.0-2.4 mm, mean 2.2 mm.

Brotogy—Larvae of B. albistylum have been collected from under
the bark of poplar (Populus deltoides) and elm (Ulmus pumila).
Cook (1953) reported that the larvae have at least 4 instars that
apparently do not form distinct size groups. He also mentioned
that the pupal period lasted from 8 to 10 days and that the adults
did not live for more than 5 days in the laboratory.

MATERIAL EXAMINED.—Light larvae from the Ohio State University
collection with the following data: Dec. 5, 1942, under bark of dead
poplar.

Genus Zabrachia Coquillett, 1901

Of the 11 species listed for this genus by Kraft and Cook (1961),
the larvae of only 2 species have been found. Both species were
taken from beneath the bark of coniferous trees.

GENERIC CHARACTERS.—As given in key to genera.

Key to Species of Zabrachia Coquillett

(after Kraft and Cook, 1961)

Ventral surface of abdominal segment 6 with 18 large plaques . politum Coquillett
Ventral surface of abdominal segment 6 with 16 large plaques.
plicatum Kraft and Cook

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Zabrachia politum Coquillett, 1901
Fiaure 145

DescriptTion.—Mature larva: length 4.3 mm; width 1.0 mm; other
characters as given in key to species.

Brotocy—No data are available for this species.

MATERIAL EXAMINED.—No larvae of this species were examined.

Zabrachia plicatum Kraft and Cook, 1961
Fiaure 142

Description.—Mature larva: length 4.0-5.0 mm, mean 4.7 mm;
width 0.7-1.0 mm, mean 0.5 mm; other characters given in key to
species.

Brotocy.—Larvae have been collected from beneath the bark of
Pinus contorta, P. ponderosa, Picea engelmanni, and Pseudotsuga
mucronata.

MATERIAL EXAMINED.—North Carolina: Raleigh, 1941-1942, 9
larvae from fallen pine, NCSC.

Discusston.—In keying out this species, I found it necessary to
slide-mount the integument in order to be sure of the number of
plaques on segment 6.

Genus Neopachygaster Austen, 1901

Kraft and Cook (1961) recognize 4 species in this genus. The
larvae have been collected from both coniferous and deciduous trees.
GENERIC CHARACTERS. —As given in key to genera.

Key to Species of Neopachygaster Austen

Kraft and Cook (1961) were unable to differentiate between the
larvae of the species in this genus. Because of a lack of material,
I am also unable to contribute toward the identification of these
species.

Neopachygaster occidentalis Kraft and Cook, 1961

Figure 147

Duscriprion.—Larva: length 5.58 mm; width 1.69 mm; tufts of }
setae of thoracic leg group with 2 inner seate of equal length, outer —
seta much shorter; otherwise indistinguishable from other larvae of
this genus. |

Brotocy.—Larvae were taken from under bark of Pinus ponderosa.

MATERIAL EXAMINED.—No larvae of this species were examined.

Discuss1on.—All data presented for this species are taken directly |
from Kraft and Cook (1961).

:

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 45

Neopachygaster maculicornis (Hine), 1902
Ficures 18, 146

Description.—Mature larva: length (5.50-6.00 mm) 6.0-6.5 mm,
mean 6.17 mm; width (1.40—-1.50 mm) 1.1-1.50 mm, mean 1.27 mm;
tufts of setae in thoracic leg group same as in WN. occidentalis.

Biotocgy.—Larvae have been collected from beneath the bark of
a fallen poplar (Populus species).

MATERIAL EXAMINED.—Alberta: Medicine Hat, South Saskatche-
wan River, June 7, 1960, Ball, Madge, and McFadden, approximately
100 specimens of larvae and puparia, MWM.

Discusston.—Adults of this species were reared from larvae and
the measurements outside of the parentheses are based on these
specimens.

Neopachygaster vitrea Hull, 1930

Description.—According to Kraft and Cook (1961), the larva of
this species is essentially the same as the larva of N. maculicornis.

BtoLtocy.—No data are available for this species.

MATERIAL EXAMINED.—No larvae of this species were examined.

Neopachygaster reniformis Hull, 1942

Description.—Larva: length 4.93 mm; width 1.45 mm; very simi-
lar to larvae of N. maculicornis; tufts of thoracic leg setae with the
middle setae longest, inner seta next longest.

Biotocy.—No data are available for this species.

MATERIAL EXAMINED.—No larvae of this species were examined.

Discusston.—All data presented for this species were taken directly
from Kraft and Cook (1961).

Genus Euphachygaster Kertesz, 1911

Kraft and Cook (1961) recognize 3 species in this genus. The
larvae have been collected from beneath the bark of both deciduous
and coniferous trees.

GENERIC CHARACTERS.—As given in key to genera.

Key to Species of Eupachygaster Kertesz
(modified after Kraft and Cook, 1961)

1. Abdominal terga 1-7 of approximately equal length with setae in transverse
TOW Sime rey Use en a Sel eR Behe ee de punctifer Malloch
Setae in transverse rows with the outermost setae much shorter than the
OURErsiminptneGrrOWer eerie ote hoc Ss eas ne co Ley eee ene GREE eg 2

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

2. Transverse row of 6 setae on abdominal sterna 1-7 with all setae of approxi-
mately equalilengthcw. curl toue) hte eiuetie) ene bes henshawi Malloch

Setae in transverse row with outermost setae much longer than others.
fusca Kraft and Cook

Eupachygaster punctifer Malloch, 1915
Figure 143

DescripTion.—Mature larva: length (6.2 mm) 5.0-6.5 mm,
mean 5.8 mm; width (1.7 mm) 1.0-1.3 mm, mean 1.2 mm; other
characters as given in key to species.

Brotogy.—Larvae have been collected from under the bark of
Carya, Populus, and an unknown species of Pinus.

MatTERIAL EXAMINED.—Delaware: Sussex County, 1952, W. A.
Connell, 6 larvae from under bark, UD. North Carolina: Auburn,
Nov. 2, 1956, Bowden and Wright, 3 larvae from under bark of oak
log, NCSC; Herring, August 23, 1956, C. G. Wright, 1 larva from
under bark of oak log, NCSC; West End, July 14, 1941, 12 larvae
from under bark of pine log, NCSC.

Eupachygaster fusca Kraft and Cook, 1961
Fiaure 140

Description.—Mature larva: length (5.85 mm) 4.2-7.0 mm,
mean 6.0 mm; width (0.169 mm) 0.9-1.7 mm, mean 1.5 mm; other
characters as given in key to species

Brotoagy.—The only recorded host for this species is the willow
(Salix species).

MATERIAL EXAMINED. —North Carolina: Rocky Mountain, Mar.
25, 1954, D. M. Weisman, 20 larvae from under bark of log, NCSC.
Ohio: Wooster, Apr. 22, 1940, H. R. Dodge, 5 larvae from under
willow bark, NCSC.

Eupachygaster henshawi Malloch, 1917
Figure 144

Description.—Mature Larva: length (4.95 mm) 5.7-6.9 mm,
mean 6.5 mm; width (1.44 mm) 1.2-1.3 mm, mean 1.23 mm; other |
characters as given in key to species

Brotoagy.—Larvae have been collected from under the bark of
apple, elm, and oak trees. |

MATERIAL EXAMINED.—North Carolina: Auburn, Jan. 8, 1956, |
C. G. Wright, 3 larvae from under bark of oak log, NCSC; Auburn,
Aug. 25, 1956, C. G. Wright, 3 larvae from under bark of oak tree, |
NCSC.

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 47

Genus Pachygaster Meigen, 1803

Kraft and Cook (1961) list 3 species for this genus but larvae of
only Pachygaster pulchra have been found.
GENERIC CHARACTERS. —As given in key to genera.

Pachygaster pulchra Loew, 1863
Fieure 148

DescripTion.—Larva: length 4.82 mm; width 0.94 mm; narrow,
dirty white, dorsal setae knobbed at tips; only 9 pairs of setae on ab-
dominal segments 1-7.

Biotogy.—Larvae of this species have been collected from tree
crotch debris, treeholes, and hollow trees.

MATERIAL EXAMINED.—No larvae of this species were examined.

Discussion.—All data presented for this species were taken
directly from Kraft and Cook (1961).

Literature Cited

AustTEn, E. E.

1899. On the preliminary stages and mode of escape of the imago of the
dipterous genus Xylomyia Rondani (Subula Meigen et auct.) with
special reference to Xylomyia maculata F., and on the systematic
position of the genus. Ann. Mag. Nat. Hist., vol. 3, no. 7, pp.

181-190.
Baker, C. F.
1895. Biological notes on some Colorado Diptera. Ent. News, vol. 6, no.
6, p. 6.
BEING, T.

1883. Beitrag zur Metamorphose zweifliigeliger Insekten aus den Familien
Tabanidae, Leptidae, Asilidae, Empidae, Dolichopidae, und
Syphidae. Arch. Nat., vol. 48, pp. 186-240.

BEARD, L.

1861. Saggio di Ditterologia Messicana. Mem. Reale Accad. Sci. Torino,

vol. 19, pp. 201-278.
Bere, C. O.

1952. Biology and metamorphosis of some Solomon Islands Diptera, 2:
Solva bergi James (Erinnidae), with a comparison of related species.
Pan-Pacific Ent., vol. 28, no. 4, pp. 203-215.

BERTRAND, H.

1948. Note sur deux larves du genre Hermione Meigen. Bull. Soc. Ent.

France, vol. 53, pp. 55-58.
Biscuorr, W.

1925. Ueber die Kopfbildung der Dipterenlarven, 3: Die Kopfe der
Orthorrhapha-Brachycera-Larven. Arch. Naturg., vol. 90, pp. 1-
105.

BorGMEIER, T.

1930. Ueber das Vorkommen der Larven von Hermetia illucens L. in den

Nestern von Meliponiden. Zool. Anz., vol. 90, pp. 225-235.
Boucuf, P. F.

1834. Naturgeschichte der Insekten besonders in Hinsicht ihrer ersten

Zustande als Larven und Puppen.
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221-533—66——5

58

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VOL. 121

TABLE 1.—Habitats of stratiomyid larvae occurring in America north of Mexico

Genus

Xylomya
Solva

Altermetoponia
Allognosta

Actina

Beris

Exodontha

Sargus

Ptecticus

Microchrysa

Merosargus

Chloromyia

Cyphomyia
Dicyphoma
Adoxomyia

Hermetia

Dieuryneura

Oxycera

Habitat

Terrestrial; in crotch of tree
Terrestrial; under bark of
trees; in rotted logs

Terrestrial; in sod

Terrestrial; in decaying
organic material

Terrestrial; in decaying
plant and animal material

Terrestrial; in decaying
leaves; under bark of
fallen trees; in moss?

Terrestrial; in rotten wood
under large rocks

Terrestrial; in decaying
plant and animal material;
in excrement

Terrestrial; in decaying
organic material

Terrestrial; in decaying
organic material; in
garden soil; in excrement

Terrestrial; in debris at
base of squirrel’s nest

Terrestrial; in decaying
organic material

Terrestrial; in decaying
plant material
Terrestrial; in decaying
plant material
Terrestrial; in decaying
plant material
Terrestrial; in decaying
organic material; in
excrement

Terrestrial; in decaying
plant material

Aquatic; on margins of
lakes, ponds, and streams

(genera arranged by subfamily as given on p. 13)

Reference

New record

Townsend, 1893; Malloch,
1917; Johannsen, 1922;
Greene, 1926; Peterson,
1951; Hennig, 1952

Irwin-Smith, 1920

Malloch, 1917; Johannsen,
1922

Fuller, 1934

Williston, 1908; de Meijere,
1916; Lenz, 1923

New record

Westwood, 1840; Lundbeck,
1907; Williston, 1908;
Malloch, 1917; Johannsen,
1922; Peterson, 1951;
Hennig, 1952

Lindner, 1928; Hennig,
1952

Lundbeck, 1907; Malloch,
1917; Johannsen, 1922;
Seguy, 1926; Hennig,
1952

New record

Brauer, 1883; Cornelius,
1860; Lundbeck, 1907;
Seguy, 1926

James, 1957

James, 1962
New record

Williston, 1908; Malloch,
1917; Johannsen, 1922;
Copello, 1926; James,
1935; James, 1497;
James, 1957

New record

Heeger, 1856; Lundbeck,
1907; Johannsen, 1922;
Lenz, 1923; Johannsen,
1935; Wesenberg-Lund, 1943

NO. 3569

Genus

Euparyphus

Caloparyphus

Stratiomys

Hedriodiscus

Odontomyia

Myzxosargus

Nemotelus

Berkshiria

Zabrachia

Neopachygaster
Eupachygaster

Pachygaster

SOLDIER FLY LARVAE—McFADDEN 59

TaBLE 1.—Continued

Habitat

Aquatic; on margins of
aquatic environments;
usually associated with
mossy conditions

Aquatic; in bog or swamp
areas; usually associated
with mossy conditions

Aquatic; on margins of
aquatic environments;
usually associated with
Typha spp.; occasionally in
hot springs or saline
habitats

Aquatic; on margins of lakes
and ponds

Aquatic; in much the same
habitat as given for
Stratiomys

Aquatic; no specific data
available

Aquatic; under debris at
margins of lakes and
ponds; frequently in
saline habitats

Terrestrial; under bark of
trees

Terrestrial; under bark of
coniferous trees

Terrestrial; under bark of
trees

Terrestrial; under bark of
trees

Terrestrial; in tree holes; in
crotch debris and in hollow
trees

Reference

Johannsen, 1922;
Johannsen, 1935;
Wesenberg-Lund, 1943;
Peterson, 1951; James,
1960

Quist (thesis) ; James,
1960

Hart, 1895; Johnson,
1895; Miall, 1895;
Lundbeck, 1907;
Williston, 1908;
Malloch, 1917;
Johannsen, 1922, 1935;
Peterson, 1951;

James, 1960b

James, 1960b

Hart, 1895; Johson, 1895;
Lunbeck, 1907; Williston,
1908; Malloch, 1917;
Johannsen, 1922, 1935;
Peterson, 1951; James,
1960b

New record

Haliday, 1857a; Lunbeck,
1907; Malloch, 1917;
Johannsen, 1922; Lenz,
1923; Johannsen, 1935;
Wesenberg-Lund, 1943;
Hanson, 1958; James,
1960

Kraft and Cook, 1961;
Cook, 1953

Malloch, 1915; Malloch,
1917; Kraft and Cook,
1961

Malloch, 1917; Cook, 1953;
Kraft and Cook, 1961

Malloch, 1917; Kraft and
Cook, 1961
Kraft and Cook, 1961

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

TABLE 2.—Predators and parasites of stratiomyid larvae in America north of Mexico

Parasite or predator Host Reference

Coleoptera
Coccinellidae
Megilla maculata Egg mass of Odontomyia

species

Hart, 1895

Hymenoptera
Pteromalidae
Rhicnocoelia, new species!
Genus species

Odontomyia species New record
Stratiomys species

Chloromyia formosa Lundbeck, 1907

Eulophidae
Tetrastichus species Microchrysa polita Lundbeck, 1907
Chalcidae
Chalcis barbara Stratiomys norma Hart, 1895
Chalcis microgaster Odontomyia cincta; Hart, 1895
Odontomyia species;
Odontomyia vertebrata; egg
mass of Odontomyia species
Ichneumonidae
New genus, new species?| Stratiomys species New record

Lundbeck, 1907
Lundbeck, 1907

Nemotelus species
Sargus species

Genus, new species
Genus, species

1 Taxonomic status as given by B. D. Burks, U.S. National Museum.
2 Taxonomic status as given by L. H. Walkely, U.S. National Museum.

TaBLE 3.—Ecological and morphological specializations in stratiomyid larvae

Subfamily Environment Nutrition Mouthparts
Xylomyinae terrestrial- | micropantophagous cylindrical brushes
arboreal present
Chiromy zinae terrestrial phytophagous brushes and setae
absent
Beridinae terrestrial micropantophagous cylindrical brushes
present
Sarginae terrestrial copraphagous, degenerate, cylindrical
sapronecrophy- brushes reduced
tophagous
Clitellariinae terrestrial copraphagous, variable, degenerate to
sapronecrophy- well developed
tophagous
Stratiomyinae aquatic micropantophagous cylindrical brushes
absent
Nemotelinae terrestrial micropantophagous | cylindrical brushes
absent
Pachygastrinae terrestrial- | micropantophagous | cylindrical brushes
arboreal present

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 61

Xylomyinae
Chiromyzinde

Clitellariinae
Stratiomyinae

Xylomyinae * : A
comme |

}O| >| a>
ea >i>ia

Beridinae

se
esters
[seternnee

Renetae |_|

Pachygastrinae

[> [0 [0[0] > [> [0 [rscuostin

Ficure 1.—Subfamily relationships of the Stratiomyidae. Each symbol represents a range
of numerical value, which is the sum of the differences in 11 characters (25 variates)

between a pair of subfamilies. Maximum values indicate maximum differences and
distant relationships; minimum values indicate minimum differences and close relationships.

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

62

SYNIAWOTAX

avNianaa

AVNIONVS

AVNIV SLND

AYNILSVSDAHDVd

AVNITALOWIN

AVNIAWOILVELS

AVNIZAWOUIHD

INasidd

isd

Ficure 2.—Proposed phylogeny for the eight stratiomyid subfamilies.

HY NX

Te (Ce

JY

iT 12

Ficures 3-13.—3, Odontomyia species, head, dorsal view (after Cook, 1949; A=antenna,
B= mandibular-maxillary complex, C=labrum, D=eye, E=clypeus, F=ocular lobe, G=
frontal suture); 4, Odontomyia species, head, ventral view (after Cook, 1949; H= maxillary
palp, I=palatum, J=prementum, K=submentum, L=pharynx); 5, Odontomyia species,
head, lateral view (after Cook, 1949; M=pestle, N=labrum, O= mandibular articulation,
P=mandibular-maxillary complex, Q=labium); 6, Odontomyia species, integument,
cross-section (CN=calcareous nail, MC=mesocuticle, EC=endocuticle); 7, Xylomya
species, head, dorsal view; 8, Solva pallipes, first abdominal segment, dorsal view; 9,
S. pallipes, apical segment, dorsal view; 10, S. pallipes, outline of prothoracic segment,
dorsal view; 11, Xylomya species, apical segment, dorsal view; 12, S. pallipes, head,
dorsal view; 13, S. pallipes, apical segment, ventral view.

64. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 14-24.—Dorsal view: 14, Solva pallipes, mandibular-maxillary complex; 15, Alter-
metoponia rubriceps, mouthparts; 16, Dieuryneura obscura, mouthparts; 17, Sargus cuprarius
mouthparts; 18, Neopachygaster maculicornis, mandibular-maxillary complex; 19, Cypho-
myia bicarinata, labium and mandibular-maxillary complex; 20, Stratiomys norma,
mandibular-maxillary complex (after Malloch, 1917); 21, Hermetia illucens, mouthparts
(labrum removed); 22, Xylomya species, first abdominal segment; 23, Beris vallata, apical
segment; 24, D. obscura, apical segment.

No. 3569 SOLDIER FLY LARVAE—McFADDEN 65

Ficures 25-41.—Dorsal view: 25, Altermetoponia rubriceps, head; 26, Beris vallata, head;
27, Dieuryneura obscura, head; 28, B. vallata, first abdominal segment; 29, Exodontha
luteipes, head; 30, E. luteipes, apical segment; 31, Allognosta fuscitarsis, head; 32, D.
obscura, first abdominal segment; 33, Altermetoponia rubriceps, apical segments; 34,
A. rubriceps, first abdominal segment; 35, E. luteipes, first abdominal segment; 36, 4.
fuscitarsis, first abdominal segment; 37, 4. fuscitarsis, apical segment; 38, 4. fuscitarsis,
outline of prothoracic segment; 39, D. obscura, apical segment; 40, Actina incisuralis,
head; 41, 4. incisuralis, first abdominal segment.

Mn —_

Ficures 42-64.—Dorsal view: 42, Microchrysa polita, head; 43, Ptecticus trivittatus, head;
44, M. polita, first abdominal segment; 45, Actina incisuralis, apical segment; 46, M.
polita, apical segment; 47, P. trivittatus, first abdominal segment; 48, Sargus bipunctatus,
head; 49, S. lucens, head; 50, P. trivittatus, sixth and seventh abdominal segments (ventral
view); 51, P. trivittatus, apical segment; 52, S. elegans, head; 53, S. elegans, first abdominal
segment; 54, S. bipunctatus, first abdominal segment; 55, S. decorus, head 56, S. lucens,
apical segment; 57, S. bipunctatus, apical segment; 58, S. decorus, first abdominal segment;
59, S. decorus, apical segment; 60, S. elegans, apical segment; 61, S. lucens, first abdominal
segment; 62, S. cuprarius, apical segment; 63, S. cuprarius, head; 64 S. cuprarius, first
abdominal segment.

71

we A

“NH Xt

Ficures 65-82.—Dorsal view: 65, Hermetia illucens, head; 66, H. concinna, head; 67, H.
aurata, head; 68, Hermetia species, apical segment; 69, H. concinna, first abdominal seg-
ment; 70, Hermetia species, head; 71, H. aurata, apical segment; 72, H. concinna, apical
segment; 73, H. aurata, first abdominal segment; 74, Adoxomyia heminopla, apical seg-
ment; 75, H. illucens, first abdominal segment; 76, Hermetia species, first abdominal
segment; 77, A. heminopla, first abdominal segment; 78, Dicyphoma schaefferi, first
abdominal segment; 79, H. illucens, apical segment; 80, 4. heminopla, head; 81, D.
schaefferi, apical segment; 82, D. schaefferi, head.

97 100

104

Ficures 83-104.—Dorsal view: 83, Cyphomyia pilosissima, head; 84, C. pilosissima, first
abdominal segment; 85, C. pilosissima, apical segment; 86, Odontomyia pubescens, head;
87, Hedriodiscus vertebratus, head; 88, Myxosargus nigricornis, apical segment; 89, O.
pubescens, first abdominal segment; 90, O. pubescens, apical segment; 91, H. vertebratus,
apical segment; 92, M. nigricornis, first abdominal segment; 93, M. nigricornis, head;
94, Euparyphus limbrocutris, head; 95, E. limbrocutris, apical segment; 96, Aochletus
brevicornis, first abdominal segment; 97, A. brevicornis, head; 98, O. virgo, first abdominal
segment; 99, £. limbrocutris, first abdominal segment; 100, Nemotelus centralis, head;
101, 4. brevicornis, apical segment; 102, O. virgo, apical segment; 103, N. centralis, first
abdominal segment; 104, N. centralis, apical segment.

NO. 3569 SOLDIER FLY LARVAE—McFADDEN 69

105
er iow rs

AN

Ficures 105-119.—Dorsal view: 105, Stratiomys norma; 106, S. norma, head; 107, S.
norma, apical segment; 108, S. norma, head (ventral view; Abib); 109, S. norma, apical
segment (ventral view); 110, Caloparyphus tetraspilus, head; 111, C. major, head; 112,
Caloparyphus species, head; 113, C. tetraspilus, first abdominal segment; 114, C. major
first abdominal segment; 115, Caloparyphus species, first abdominal segment; 116, C.
tetraspilus, apical segment; 118, C. major, apical segment; 118, Caloparyphus species,
apical segment; 119, S. discaloides, apical segment.

118 119
AA

ae TNS
LE OWN \ WS
C7 dX SSS

¢

12

129

128

135

134 138

jee

Ficures 120-139.—Dorsal view: 120, Nemotelus kansensis, first abdominal segment;
121, N. canadensis, first abdominal segment; 122, Caloparyphus amplus, ventral hooks
(lateral view); 123, N. kansensis, apical segment; 124, N. canadensis, apical segment;
125, C. crotchi, head; 126, Odontomyia cincta, head; 127, C. amplus, head; 128, C. amplus,
first abdominal segment; 129, C. crotchi, first abdominal segment; 130, O. cincta, first
abdominal segment; 131, C. amplus, apical segment; 132, C. crotchi, apical segment;
133, O. cincta, apical segment; 134, N. canadensis, head; 135, O. occidentalis, head; 136
O. occidentalis, apical segment; 137, O. communis, apical segment; 138, O. occidentalis,
first abdominal segment; 139, O. communis, first abdominal segment.

NO. 3669 SOLDIER FLY LARVAE—McFADDEN 71

Ficures 140-148.—140, Eupachygaster fusca, dorsal view on right; 141, Berkshiria albistylum,
dorsal view on right; 142, Zabrachia plicatum, dorsal view on right; 143, E. punctifer,
abdominal terga; 144, E. henshawi, abdominal sterna; 145, Z. politum, terminal segments;
146, Neopachygaster maculicornis, dorsal view on right; 147, N. occidentalis, dorsal view
on right; 148, Pachygaster pulchra, dorsal view on right. (After Kraft and Cook, 1961.)

te PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

155

151
ii
152
AE
154
156

Ficures 149-156.—149, Xylomya americana, first abdominal segment, dorsal view; 150,
X. americana, apical segment, ventral view; 151, Merosargus caerulifrons, apical segment,
ventral view; 152, Cyphomyia marginata, head, dorsal view; 153, C. marginata, first ab-
dominal segment, dorsal view; 154, C. marginata, apical segment, ventral view; 155,
Adoxomyia rustica, first abdominal segment, dorsal view; 156, 4. rustica, apical segment,
ventral view.

U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3570

REVISION OF THE FAMILY PANDARIDAE
(COPEPODA: CALIGOIDA)!

By Rocer CRrEssEY

Associate Curator, Division of Crustacea

In 1907, C. B. Wilson published a revision of the subfamily
Pandarinae as part of a series of papers dealing with caligoid copepods.
We now recognize that much of this work was superficial, containing
descriptions of species often incomplete and inadequately figured;
nevertheless, it served to focus attention on a group of parasites,
caligoid copepods, which were then and are still today poorly known
in most cases.

Between 1960 and 1965 I collected and solicited material of the
family Pandaridae from as many different areas as possible. As a
result of this accumulation of material and data, I feel that a revision
of the family is in order. Ecological relationships are now more
evident than before.

Because of inadequate species descriptions that exist for most
members of this family, positive identification of material is often
difficult. This results in the publication of records that obscure our
understanding of existing host-parasite relationships. I believe that,
in most cases, I have examined enough samples of members of this

1 Modified from a Ph. D. dissertation submitted to Boston University, Boston,
Mass.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

group to be able to draw a clearer picture. Also, as a result of these
collections, I have been able better to define important taxonomic
characters and to discount others on which new species descriptions
have often been based. It is with the foregoing in mind that I have
made the following family revision. The Pandaridae as defined here
is composed of 12 genera and 33 species.

The material examined was preserved in 10 percent formalin or 70
percent ethyl alcohol. For detailed examination of the appendages,
the copepods were dissected in lactic acid and mounted in Hoyer’s
mounting medium. Whole specimens were often treated with 5
percent potassium hydroxide to render them more transparent. No
distortion was noticed by this method.

All drawings were made with the aid of a camera lucida. The
letter following the explanation of the figure refers to the scale at
which it was drawn. In all text tabulations, Roman numerals refer
to spines, Arabic numerals to setae.

All specimens are deposited in the U.S. National Museum unless
otherwise designated.

I wish to acknowledge the constant encouragement and helpful
advice offered by Dr. Arthur Humes, Boston University, during the
course of this study.

I also wish to acknowledge the following persons who generously
donated or loaned to me material for study: Dr. Thomas E. Bowman
and Dr. Robert Gibbs, U.S. National Museum; Dr. Eugenie Clark,
Cape Haze Marine Laboratory, Fla.; Dr. Richard Gooding, University
of Singapore; Mr. Ju Shey Ho, Boston University; Miss Leonie
Joubert, Oceanographic Research Institute, South Africa; Mr. Susumu
Kato, California Fish and Wildlife Service; and Dr. Jan Stock,
Zoological Museum, Amsterdam.

A portion of this work was supported by the National Science
Foundation as a part of the U.S. Program in Biology, International
Indian Ocean Expedition.

Family Pandaridae Milne-Edwards, 1840

FEMALE.—Body caligiform, usually with dorsal plates. First
thoracic segment fused with cephalon. Thoracic segments 2-4 free.
Genital segment conspicuous. Abdomen of 1 or 2 segments with or
without dorsal plates. Oral area with or without adhesion pads.
First antenna 2-segmented. Mandible in form of stylet with 10-12
apical teeth. Mandible inserted within mouth tube. Maxilliped
with terminal claw. Legs 1-4 biramose. Leg 5reduced. Egg strings
consisting of long strings of eggs arranged in linear series.

Maue.—Body caligiform, without dorsal plates. First thoracic

NO. 3570 PANDARIDAE—CRESSEY 3

segment fused with cephalon. Thoracic segments 2—4 free. Oral
area generally asin female. Legs 1—4 biramose, rami always with long
plumose setae. Legs 5 and 6 present. Abdomen 1- or 2-segmented.
Caudal rami large.

Discusston.—Members of the family Pandaridae are generally con-
sidered to be parasites of elasmobranch fishes. Occasionally investi-
gators report these copepods from teleost fish, but occurrence on such
hosts is undoubtedly accidental and does not indicate the true host.
The copepods are found on the body surface, cloacal aperture, gills and
gill arches, mouth, and nasal passages. Those which are found on the
body surface are often heavily pigmented; those in more protected
areas are devoid of pigment.

These parasites are well adapted for attachment to the host, the
female more so than the male. The principal attachment structure
is the maxilliped. In the case of Perissopus the maxilliped becomes
cemented to the host. Adhesion pads scattered over the ventral sur-
face of the parasite seem to help resist its sliding over the surface of
the host. The pads do not actually attach to the host but rather have
surface striations for increased friction.

Locomotion is usually restricted to the males. The swimming legs
of the females are often lamelliform (bearing reduced setae) and prob-
ably afford increased area for respiration with a reduction in the
swimming function. Males are not so modified and undoubtedly
move much more freely than the female.

The life history of pandarid copepods is still unknown. Wilson
(1907) has outlined a composite life history of this group based on
fragments of information from various representative caligoids. In
general, the life history includes the following stages: egg, nauplius,
copepodid, chalimus, and adult. The number of molts between stages
is completely unknown for this group. The nauplius is of the usual
copepod form with 3 pairs of anterior appendages. This stage is
apparently free in the plankton and undoubtedly of short duration.
The copepod attaches to the host as a copepodid (=“‘metanauplius,”
Wilson 1907). The second antenna of this stage is greatly enlarged
and projects anteriorly. The body of the copepodid is divided into
a cephalon, 2 thoracic segments, and a fused genital segment-abdomen
with rami attached distally. It is not clear whether this stage lasts for
more than 1 molt. The chalimus assumes a body form similar to
the adult. Wilson (1907) describes the chalimus stage as seen in
Perissopus dentatus. The chalimus stage of caligoid copepods is
characterized by the presence of a frontal filament. This is produced
by glands in the anterior portion of the cephalon and serves to attach
the copepod to the host. Wilson’s description includes this structure.
I have collected occasional chalimus stages in the genera Pandarus

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

and Echthrogaleus and have not observed this filament. More ma-
terial will eventually show the true nature of this structure in pandarid
copepods. The chalimus apparently undergoes a series of molts, in-
creasing the segmentation until the adult form is reached. Lewis
(1963) has described the life history stages of Lepeoptheirus dissimu-
latus Wilson. This copepod includes the following stages in its life
history: nauplius (first and second), copepodid (1 stage), chalimus
(6 stages), and adult. The life history of pandarids is probably quite
similar. Heegaard (1947) described 2 copepodid stages for Caligus
curtus (Muller).

The family Pandaridae is composed of 2 well-defined groups on the
basis of external characters of both sexes. These characters are in-
cluded in the key below. Group I includes the following genera:
Pandarus, Pseudopandarus, Phyllothyreus, Gangliopus, Perissopus, and
Pannosus. Group II includes: Dinemoura, Demoleus, Pagina, Ech-
throgaleus, Nesippus, and Paranesippus. Keys to the genera of
females of the 2 groups also are provided below (not enough informa-
tion is available to construct a key to the males).

Key to Groups and Genera of Pandaridae

FEMALES

Dorsal thoracic plates present on segments 2-4; penultimate segment of second
maxilla with 2 prominent distal spines .. . 5 56 oo GIROWIP I
Dorsal thoracic plates, if present, on segment 4 pales penultimate segment of
second maxilla with 1 spine and a patch of spinules or setules . GROUP II

MALES
Outer distal corner of last endopod segment of leg 3 smooth . . . .GROUP I
Outer distal corner of last endopod segment of leg 3 modified with roughened
areasvand Short, SpineS, =-)\s « serps) 24 28 esd eerie oO ee

GENERA OF GROUP I FEMALES

1. Dorsal thoracic plates of segment 3 extending beyond plates of segment 3 . 3
Dorsal thoracic plates of segment 3 not extending beyond plates of segment

2. Abdomen and caudal rami hidden dorsally. . .... . . . .Perissopus
Abdomen and caudal rami visible dorsally... . . . . « « Pandarus
3. Plates of segment 2 large, overlapping those of peomeat Bi4iels hoe tales
Plates of segment 2 reduced and lateral to those of segment3 ..... 5
4. Maxilliped with spatulate'tip §- 5 <5... ~ he.) « fem - .eannosus
Maxilliped with pointed tip... . . . . . .Phyllothyreus

5. Maxilliped with pointed tip, genital poement ese on \% body length.
Gangliopus

Maxilliped with spatulate tip, genital segment at least 44 body length.
Pseudopandarus

f
|

NO. 3570 PANDARIDAE—CRESSEY 5

GENERA OF GROUP II FEMALES

ip Odomen 2-sepmenved,— liv PIO ie ail) Besse EE lt Rad OT 2
BGOMmen Se Qimen bed zt NUON. 91 retell. aNd, mal oee cals fe fan esta. 3
Beebourvoles lamellifornn:. 1... .).2, 3/4) 3.)- Verne een oy sue) \y: st Dinemoura
Rounthudegsnmot-lamelliforms <0. iid & oe gS te kebew olg oh ee
pebourthwlesiamelliform ; p20. 406 he Ses de ces 8 ee Chthrogaleus
Fourth leg not lamelliform . SNe Chay ge Hea Setar N SO ct ad ore Uriah PNT ay tage ta oe
4. Abdomen with large dorsal plate. ....... =... .. .. .Demoleus
Abdomen. withouv dorsaliplate: 3. =) suct eis ss em cl ick ey ek cee O
5. Exopods of legs 1-3 3-segmented ......... .. . . . Paranesippus
Exopods of legs 1-8 2-segmented ............. . .Nesippus

Genus Pandarus Leach, 1816

Pandarus Leach, 1816, p. 405. [Type-species: P. bicolor.]
Caligus—Lamarck, 1818, p. 137. [Refers to C. bicolor only.]
Nogagus Leach, 1819, p. 536.

Frema.e.—Frontal plate distinctly separate. First thoracic seg-
ment fused with cephalon. Thoracic segments 2-4 free, bearing dorsal
plates. Dorsal plates of segment 3 never extending beyond plate
of segment 2. Plates of segment 4 fused basally. Abdomen 1-
segmented with dorsal plate. Abdomen (or its plate) visible dorsally
and attached to distal end of genital segment. Caudal rami lateral
to abdomen. First antenna 2-segmented. Adhesion pads present on
cephalon in association with first and second antennae and maxilliped ;
pads also located on ventral surface of posterior corners of cephalon.
Claw of maxilliped spatulate. Legs 1-4 biramose, rami of legs 1-3
2-segmented. Rami of leg 4 1I-segmented. Leg 5 present. Egg
strings long and straight.

Matr.—No dorsal plates present. Cephalic appendages in general
like those of female. Legs 1-4 biramose, all rami 2-segmented and
bearing plumose setae. Fifth and sixth legs present. Leg 3 endopod
unmodified.

Discusston.—Since 1816, 29 species have been described and as-
signed to this genus. Only 10 of these remain valid today. The
synonymies are discussed with the species descriptions. I propose to
designate each of the following 4 species described by Hesse in 1883
as a nomen dubium on the basis that the descriptions and figures are
so poor that it is impossible to assign these to any known taxon:
Pandarus mustelilaevis, Pandarus unicolor, Pandarus spinaciiachantias,
and Pandarus carchiiglaucus.

Members of the genus Pandarus are parasites on the body surface of
the host. They have been reported from both pelagic and inshore
species and are the most frequently encountered pandarid copepod.
The females of some species are heavily pigmented and those of others

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

show some signs of pigmentation. They frequently occur in clusters
of more than 100 individuals on the fins of the shark.

Members of this genus can easily be separated from other genera
on the basis of the arrangement of the dorsal thoracic plates and the
nature of the caudal rami.

Key to Adult Females of Pandarus

1. Dorsal plate of thoracic segment 2 extending only as far as the posterior edge
ofthe jplate of segment 3°. « =; « «.:..... sbiCOLOR (GROUPS 2
Dorsal plate of thoracic segment 2 extending well beyond posterior edge of
plate of segment 3... .. i ae CORANCHEE GROUPS

2. Cephalon only % of total body ean caudal rami small, scarcely visible
Gorsallivaemsene . . . bicolor
Cephalon about 4 of ae Body ee ae rami Manse Sere dorsally . 3

3. Caudal rami long, about 3 times as long as dorsal abdominal plate and extend-
ing well beyond it; Eee edge of Pail of segment 4 without conspicuous
Sinise he - pet niger
Caudal rami only acu 1% meas as iene as genni Cpaominnl plate and
extending only ore beyond it; posterior edge of plate of segment 4 with

deep sinus. .. . . . . Ccarcharhini

4. Caudal rami with Ae inner been half ied i. Becta lobe . . smithii

Caudal rami without such a lobe... . 5

5. Caudal rami not extending more than % icaern af Boral Bhdawaeel nigral

satyrus

Caudal rami extending at least to tip of dorsal abdominal plate or well

DEYOnGEIE yan arn sel Pk RO

6. Dorsal plates of poeta eeeenis 3 pad 4 paced aa ae peonal posterior

SHOUIS ene ses eae tad

Dorsal plates oF hora! eeements 3 aaa 4 Hivided oF deep sinus into 2

nearly separate lobes... . i SEEeS

7. Dorsal abdominal plate subtriangular eye oe Separated in fully pigmented

FORMS: saeco: s/t)... Ckanehn

Dorsal podomnse aI areeaca ‘body ae or ee once be darkly pig-
mented .

8. Most of body deride: Perel pe eye anoes Pomme by eae posterior

liayey |) La) . . Sinuatus

Body only elichale piemeated amet Piece edi pontned to anterior

portion of cephalon and plate of fourth thoracic segment . . . fioridanus

9. Thoracic plate of segment 2 extending to posterior tip of thoracic plate of

SCCmen tie eee nee ecle ezyeaenae

Thoracic plate of eign 2 eedine to iadille a homer plate of seg-

MHEG CAA ERE TS RO Ted Tae LE Ane 2 nO

Pandarus satyrus Dana, 1852

Figures 1-26
Pandarus satyrus Dana, 1852-3, p. 13868.—Brady, 1883, p. 134.—Bassett-Smith,
1899, p. 467.—Wilson, 1907, p. 415; 1914, p. 71—Yamaguti, 1936, p. 5.—
Bere, 1936, p. 595.—Shiino, 1957, p. 364; 1959a, p. 315; 1959b, p. 352;
1960b, p. 493.—Ho, 1963, p. 90.

NO. 3570 PANDARIDAE—CRESSEY |

SPECIMENS sTuDIED.—AII collections from Prionace glauca (Lin-
naeus). Twenty-six collections made in the western North Atlantic
Ocean between latitudes 30°N to 49°N and longitudes 60°W to 72°W.
Six collections in the Indian Ocean (0°58’N, 55°E; 0°14’S, 55°04’E;
6°37’S, 55°00’E; 34°32’S, 74°48’E; 33°11’S, 54°58’E; 02°06’S, 75°10’E.
Three collection in Pacific Ocean (3°18’N, 101°54’W; 1°00’S,
101°40’W; 9°56’/N, 135°16’W).

FEMALE.—Body form as in figure 1. Length 8.2 mm and width
(measured at widest point) 4.4 mm based on average of 10 specimens.

First thoracic segment fused with head. Dorsal thoracic plates
present on segments 2-4. Plates on segment 2 separate, extending
laterally beyond tip of plate of segment 3. Plates of segment 3
fused at base, divided by a broad median distal sinus in middle of
distal margin. Plates of segment 4 extending over the genital seg-
ment, fused, and with distal median sinus. Genital segment 1.8 mm
long and 2.7 mm wide. Abdomen 1-segmented and joined broadly to
genital segment ventrally. Abdomen covered by dorsal plate
(1.8 by 1.5 mm), longer than wide, extending beyond tips of rami.
Caudal rami (fig. 2) long, slender, widest at base, tapering distally
(990u by 228u), bearing 4 short spines. Rami extend only to about
middle of abdominal plate.

Oral area (fig. 3). Adhesion pads present at bases of first antennae,
second antennae, and maxillipeds. Pads also present at posterior
corners of cephalon. First antenna (fig. 4) 2-segmented. First
segment bearing 26 setae, 4 setae small and plumose, remainder stout
and armed as in figure. Second segment bearing 12 naked setae.
Second antenna (fig. 5) 3-segmented. Terminal segment bearing
large terminal spine and 2 setae. Mouth tube (figs. 6, 7) of usual
caligoid type. Labrum with pari of subterminal processes within tube
and fringed at its tip. Labium somewhat expanded at tip and with
fringe as in figure 7. Mandible (fig. 8) composed of basal podomere
bearing long slender shaft with serrate tip. Mandibular process ex-
tending within tube as in figure 6. First maxilla (fig. 6) 2-segmented.
Basal segment bearing 3 short setae. Terminal segment with small
seta and large terminal spine. Second maxilla (fig. 9) with 3 segments.
Basal segment unarmed. Second segment with 2 distal spines, longer
one fringed, shorter plumose. Terminal segment bearing large claw
with rows of spinules and apical patch of spinules. Maxilliped (fig.
10) 2-segmented. Basal segment stout, bearing an adhesion pad.
Terminal segment in shape of a claw, with 1 seta. Tip of claw bilobed,
with an adhesion area that, when closed, is in contact with adhesion
pad of basal segment. Legs 1—4 biramose with spine and setal formula
as follows:

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

leg 1 leg 2 leg 8 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 120 0:0 1:0 0:0 VI 0
seg. 2 VI Iil xX IV VI II - -

Leg 1 (fig. 11) with both rami 2-segmented. Leg 2 (fig. 12) with both
rami 2-segmented. Inner 7 spines on last segment of exopodite not
articulated. Leg 3 (fig. 13) with both rami 2-segmented. Two inner
spines on last segment of exopodite not articulated. Leg 4 (fig. 14)
with both rami 1-segmented. Endopodite without spines. Legs 1-4
with patches of spinules and adhesion pad as in figures. Leg 5 (fig.
15) consisting of outer plumose seta and inner lobe with single terminal
spine.

Egg strings long and slender, extending several times length of body,
not recurved. Eggs disc shaped.

Color of adult female dark brown to black with eye spots separated
as In figure 1.

Ma.sr.—Body form as in figure 16. Length (not including setae
on caudal rami) 10.3 mm and width (measured at widest point) 5.6
mm based on an average of 10 specimens. Cephalon rounded when
viewed dorsally with head and first thoracic segment fused. Thoracic
segments 2-4 free, without dorsal plates except for lateral winglike
plates on segment 2. Genital segments 2.3 mm by 2.3 mm, with
posterior corners attenuated, terminating in inwardly directed tip.
Spermatophores (fig. 17) often visible through genital segment.
Abdomen 2-segmented. First segment 432u long. Second segment
576u long. Caudal ramus (fig. 18) 1050u by 665y, bearing 4 long and
2 short setae. Outermost long seta 1365u long, innermost 910 long,
and 2 median setae 1500u long. Oral area as in female except as
follows: second antenna (fig. 19) with last segment in form of large
claw, not subdivided as in female; maxilliped (fig. 20) terminating in
pointed claw whereas female claw terminating as spatulate tip.

Legs 1-4 biramose, each ramus of 2 segments, with spine and setal
formula as follows:

leg 1 leg 2 leg 8 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1-6 O:0% Hed O21 Gobet Orie) oho 0:1
seg. 2 Vie 3 IV:6 8 LVig5 6 IVigdia0 5
All setae densely plumose. Leg 1 as in figure 21. First segment of
endopod with prominent inner adhesion pad. Leg 2 as in figure 22.
Leg 3 (fig. 23) with bilobed outer edge on first segment of exopodite.
No modified setae on last endopodite segment. Leg 4 (fig. 24) also ©
possessing bilobed outer edge on exopodite first segment. Leg 5 |
(fig. 25) borne on genital segment as lateral projection with 4 setae and
1 stout terminal spine (see also fig. 16). Leg 6 (fig. 26) consisting of |

NO. 3570 PANDARIDAE—CRESSEY 9

single plumose seta and stout spine borne on genital segment near
origin of abdomen (see also fig. 16).

Color in life whitish yellow and generally devoid of darker pigment.

Discusston.—Pandarus satyrus is a widely distributed copepod
nearly always found associated with Prionace glauca. Reports in
the literature of this copepod from other hosts are probably the result
of misidentification of the copepod or even of the host shark. It is
closely related to Pandarus cranchii Leach and was placed in synonymy
with it by Shiino (1954). Ihave examined a great number of specimens
of both of these species and conclude that both are valid species. The
principal differences between them are discussed under the description
of Pandarus cranchii.

This copepod is parastic on the body surface of the host and is often
found in large clusters near the posterior edge of the fins. Generally
the number of females far exceeds the number of males.

Pandarus cranchii Leach, 1819
FIGuREs 27-33

Pandarus cranchii Leach 1819, p. 535.—Burmeister, 1833, p. 331.—Kr¢gyer, 1837,
p. 202.—Steenstrup and Liitken, 1861, p. 390.—Rathbun, 1884, p. 488;
1886, p. 317.—Beneden, 1892, p. 221.— Wilson, 1907, p. 403; 1908, p. 453;
1932, p. 485; 1936, p. 333.—Brian, 1908, p. 4; 1912, p. 14.—Leigh-Sharpe,
1934a, p. 27.—Pesta, 1934, p. 30.—Oorde and Schuurmans Stekhoven, 1936,
p. 141.—Heegaard, 1943b, p. 27.— Barnard, 1948, p. 249; 1955, p. 258.—
Capart, 1953, p. 660; 1959, p. 98 — Markevitch, 1956, p. 151.

Nogagus latreilli Leach, 1819, p. 536.

Pandarus carchariae Leach, 1819, p. 535.

Pandarus concinnatus Dana, 1852, p. 59.

Pandarus pallidus Milne-Edwards, 1840, p. 468.

Pandarus vulgaris Milne-Edwards, 1840, p. 468.

Pandarus dentatus Milne-Edwards, 1840, p. 469.—Heller, 1868, p. 206.—Thom-
son, 1889, p. 363.—Bassett-Smith, 1899, p. 466.

Pandarus armatus Heller, 1868, p. 202.—Thomson, 1889, p. 363.—Bassett-Smith,
1899, p. 467.—Wilson, 1907, p. 448.—Stebbing, 1910, p. 558.—Capart, 1953,
p. 659.—Barnard, 1955, p. 258.

SPECIMENS stupIED.—Kighteen collections from Pterolamiops longi-
manus (Poey) between latitudes 30°N to 49°N and longitudes 60°W
to 72°W. ‘Two collections from Eulamia falciformis (Miller and
Henle) (30°49’N, 64°02’W; 57°44’N, 65°42’W). Two collections
from Hulamia obscura (Lesueur) (34°45’N, 73°41’W; 36°42’N,
70°00’W). Ten collections from Eulamia floridanus (Bigelow and
Schroeder) between latitudes 30°N to 49°N and longitudes 60°W to
72°W. All above collections in the North Atlantic Ocean. Indian
Ocean: 8 collections from P. longimanus (07°17'N, 55°00’E; 02°25/N,
55°04’E; 00°14’S, 55°04’E; 06°37’S, 55°00’E; 11°08’S, 55°04’E;

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

12°38’S, 54°40’E; 04°09’S, 74°58’E; 03°54/N, 74°59’E). Two collec-
tions from E. floridanus (07°17’N, 55°00’E; 00°14’S, 55°04’E).
Pacific Ocean: from Carcharinus malpeloensis (Fowler) (9°42’N,
85°46’W), Carcharinus galapagensis (14°3’N, 92°8’W) and Sphyrna
zygaena (Linnaeus) off Formosa. Also 1 collection from P. longimanus
from Durban, South Africa, and 4 collections from this host in the
Gulf of Mexico. Two collections from Galeocerdo cuvier (Lesueur), 1
from Formosa and the other from Pointe Noire, West Africa.

FrMALE.—Body form as in figure 27. Length 8.4 mm and width
(measured at the widest part) 4.6 mm. Based on an average of 10
specimens.

In general, dorsal view and arrangement of thoracic plates of
female is same as P. satyrus with the following exception: caudal ramus
(fig. 28) of P. cranchii always extends to distal tip of abdominal
plate and often even beyond while ramus of P. satyrus is much shorter
in relation to abdominal plate (see fig. 2). Ramus of P. cranchii
measures 1.4 mm in length.

Oral area as in P. satyrus except that maxilliped (fig. 29) of P.
cranchii possesses a larger basal segment. Leg 1-4 with spine and
setal formula as P. satyrus. Endopodite of leg 2 of P. cranchit
(fig. 30) with patch of more rugose spines along outer edge than that
of P. satyrus. Leg 5 (fig. 31) asin P. satyrus. Eggs and egg strings as
in P. satyrus.

Color as in P. satyrus.

Maur.—Body form as in P. satyrus. Total length 10.4 mm.
Greatest width 5.2 mm, based on an average of 10 specimens. Male of
P. cranchii differs from P. satyrus as follows: second antenna of P.
eranchii (fig. 32) with smaller claw; posterior corners of genital
segment of P. cranchii attenuated but not curved inwardly as in
satyrus (see fig. 33).

Discussion.—This copepod appears to be closely related to P.
satyrus but the two species are easily separated on the basis of the
caudalrami. The rami of P. cranchii extend at least to the tip of the
abdominal plate (often beyond) whereas the rami of P. satyrus extends
only about half the length of the abdominal plate. The legs of P.
cranchii have the same spine and setal formula but the patches of
spinules are much heavier in P. cranchit.

Pandarus cranchii seems to be confined primarily to carcharinid
sharks. It is also common on the tiger shark, Galeocerdo cuvier. The
parasite is found on the body surface of the host and like other mem-
bers of the genus is sometimes found in large clusters on the fins.
It is worldwide in distribution.

Shiino (1954) has described and figured both sexes of this species
under the name of P. satyrus.

NO. 3570 PANDARIDAE—CRESSEY iy

Pandarus smithii Rathbun, 1886
Fiaures 34-35

Pandarus smithii Rathbun, 1886, p. 315.—Wilson, 1907, p. 410; 1932, p. 158.—
Leigh-Sharpe, 1934, p. 27.—Bere, 1936, p. 595.—Carvalho, 1940, p. 281; 1945,
p. 110; 1951, p. 1389.—Brian, 1944, p. 202.—Barnard, 1948, p. 249; 1955,
p. 259.—Causey, 1955, p. 6.—Shiino, 1959b, p. 353.

Pandarus lugubris Heller, 1868, p. 205.—Bassett-Smith, 1899, p. 467.—Brian,
1902, p. 8; 1906, p. 56.— Wilson, 1907, p. 395.—Rose and Vaissiere, 1953, p. 86.

Pandarus marcusi Carvalho, 1940, p. 284; 1951, p. 140.

SPECIMENS sTUDIED.—Two collections from Jsurus oxyrhynchus
Rafinesque (42°18’N, 64°02’W; 35°00’N, 70°00’W). Two collections
from Eulamia obseurus (Lesueur) (36°42’N, 70°00’W; 34°45’N,
73°41’W). Three collections from Eulamia floridanus Bigelow and
Schroeder (36°07’N, 73°25’W; 38°00’N, 68°00’W; 40°25’/N, 62°35’W).
All above in North Atlantic Ocean. From the Indian Ocean 2 collec-
tions from Alopias vulpinus (Bonneterre) (9°24’N, 54°58’E; 07°17’N,
55°00’E). Single collections from Jsurus oxyrhynchus (16°13’N,
63°29’E) and Eulamia floridanus (07°17’N, 55°00’E). From the
Pacific Ocean 2 collections from Carcharinus azureus (9°52’N, 85°29’W ;
10°09’N, 86°04’W). Single collections from Carcharinus galapagensis
(9°45’N, 85°34’W), Carcharinus malpeloensis (Fowler) (Trés Madres
Island), Carcharinus limbatus Miiller and Henle, off San Pedro, Calif.
Also single collections from Hypoprion signatus Poey in the Gulf of
Mexico, Carcharodon carcharias (Linnaeus) off Cape Cod, Mass., and
Sphyrna zygaena (Linnaeus) off the coast of Brazil.

FremaLe.—Body form as in figure 34. Length 8.2 mm and width
4.6 mm based on an average of 10 specimens. Dorsal thoracic plates
similar to those of P. satyrus and P. cranchii except that plates of
segment 2 extend only to about middle of plate of segment 4. Genital
segment 2.2 mm long and 3.0 mm wide. Abdomen 1-segmented as in
P. satyrus. Dorsal plate of abdomen nearly round (see fig. 35).
Caudal rami (figs. 35, 36) 1.78 mm long and with an inner lobe extend-
ing ventrally below abdomen. In some specimens lobes extending
toward midline almost touching and thus forming shelflike structure
bridging the rami.

Oral area similar to that of P. satyrus. Second antenna (fig. 37)
and maxilliped (fig. 38) are illustrated to show minor differences in
shapes of segments and lengths of spines. Legs 1-4 modified as in P.
satyrus and with the following spine and setal formula:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 10 0:0 TOM O30 Var ord’
seg. 2 VI Iil xX IV Vile or ul

Formula like that of P. satyrus and P. cranchii except in variation of
numbers of spines on last podemere of expodite of legs 3 and 4 (fig. 39).

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Leg 5 (fig. 40) with inner lobe in form of large claw. Egg strings as in
other Pandarus species. Color dark brown to black in pigmented
areas (see fig. 34). Eye spots joined at midline and not separated
as in P. satyrus.

Maur.—Body form as in figure 41. Length 9.4 mm (not including
setae on caudal rami) and width (measured at widest point) 4.4 mm
based on an average of 10 specimens. General configuration similar
to other Pandarus species. Genital segment (fig. 42) 2.0 mm by
1.8 mm with posterior corners not produced as in P. satyrus and P.
cranchit. Spermatophore visible through genital segment. Oral area
as in P. satyrus. Legs 1.4 biramose with spine and setal formula as
in P. satyrus. Leg 1 (fig. 43) showing minor differences in adhesion
areas and length of setae when compared with P. satyrus. Leg 2
(fig. 44) with papillose area on outer corner of coxopodite rather than
patch of fine setules as in P. satyrus. Leg 5 (fig. 45) with 3 plumose
setae and 1 small spine and borne laterally on genital segment as in
P. satyrus. Leg 6 consisting of 1 short spine and a single seta located
at junction of genital segment and abdomen.

Color in life whitish yellow and generally devoid of darker pigment.

Discusston.—Pandarus smithii is a widely distributed copepod
found on a number of hosts and often occurring with Pandarus cranchii
on carcharinid sharks. P. smithit can be easily distinguished from
the other species of the cranchit group on the basis of the caudal
ramus and eye spots in pigmented females. Males can be separated
from cranchii and satyrus on the basis of the shape of the genital
segment and nature of leg 5.

In 1940 Carvalho described a new species, Pandarus marcusi. I
have examined material identified by Carvalho as this new species
from Brazil and have concluded that it is in fact a synonym of P.
smithvi. The sole basis for separating marcusi from smithii was the
shape of the dorsal plate of the abdomen. P. marcusi had a plate
with a deep median sinus. This irregularity in the shape of plates is
not uncommon in the genus. I have observed these irregularities in
collections of Pandarus that show an epiphyte (algal or fungal)
growing on the surface of the copepod. The “roots” of the epiphyte
seem to have a corrosive action on the integument and often leave
the area malformed.

Pandarus floridanus, new species
Figures 46-66
SPECIMENS sTupIED.—Thirty-two females and 1 male collected
from Carcharodon carcharias (Linnaeus) caught at Dennis, Mass.
Holotype female, allotype male, and 10 paratype females in alcohol
deposited in the U.S. National Museum, 10 paratype females in

NO. 3570 PANDARIDAE—CRESSEY 13

alcohol deposited in the British Museum (Natural History) and the
remaining paratypes in the author’s collection. Other specimens
studied: 3 collections from Carcharodon carcharias (USNM 107299,
Miami, Fla.) (USNM 101876, off Cape Lookout, N.C.) (USNM
104859, St. Augustine, Fla.); a single collection (USNM 32765) from
Lamna nasus (Bonneterre) Woods Hole, Mass.

Frma.Le.—Body form as in figure 46. Total length (based on an
average of 5 specimens) 7.5mm. Greatest width (measured at widest
part of cephalon) 3.8mm. Cephalon widest posteriorly narrowed
anteriorly, measuring 3.5 by 3.8mm. First thoracic segment fused
to head. Dorsal thoracic plates present on segments 2-4. Plates of
segment 2 extending well beyond those of segment 3 to about middle
of lateral edge of plate of segment 4. Plates of segments 3 and 4
fused at bases. Genital segment nearly square measuring 2.8 mm
long and 2.6 mm wide. Dorsal posterior border of genital segment
has broad sinus to accommodate dorsal abdominal plate. Abdomen
1-segmented with a dorsal plate (see fig. 47) slightly wider than long
(7.9 by 7.2mm). Caudal ramus of usual generic type, bearing 4
setae (see fig. 47) and 0.6 mm long.

Oral area as in figure 48. First antenna 2-segmented. First
segment 4904 long bearing 25 spines and setae. Spines and setae
armed as in the figure. Second segment 168u long bearing 11 naked
setae. Second antenna (fig. 48) 3-segmented. Terminal segment in
form of claw and bearing 2 spines. Mouth tube with associated
mandible and first maxilla as in other members of genus. Second
maxilla with usual fringed clawlike tip. Short plumose seta (fig. 49)
near base of terminal segment. Maxilliped of the usual type (see
fig. 48).

Head appendages, except maxilliped, as in female. Maxilliped
with distinct claw at tip as in P. satyrus (see fig. 20).

Legs 1-4 not modified as in female and with spine and setal formula
as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 Lek O:1 lesan O:1 TA 0:1
seg. 2 Wes IV:6 8 LVS: 1 AG JAY Ray

Legs 1-4 biramose and each ramus 2-segmented. All setae plumose.
Leg 1 (fig. 61) with adhesion pads as indicated in figure. Leg 2
(fig. 62) exopod with first segment bearing long hairs along outer
edge and outer distal corner. Leg 3 (fig. 63) with no modifications
for holding on last endopod segment. Leg 4 as in figure 64. Leg 5
(fig. 65) consisting of 3 short plumose setae and one short, stout
spine located on midlateral margin of genital segment. Leg 6 (fig. 66)
located at junction of genital seement and abdomen, consisting of a
spinelike process with stout plumose seta near base.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Color in life cream and generally devoid of pigmentation.
Legs 1-4 biramose, with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
end. exp. end. exp. end. exp. end.
seg. 1 T:0 0:0 1:0 0:0 1:0 0:0 Vip d
seg. 2 VI Ill IX V VI II

Leg 1 (fig. 52) with both rami 2-segmented. First exopod segment
with outer spine. Last exopod segment with 6 spines, outer 3 naked,
inner 3 finely plumose. First endopod segment with adhesion pad.
Last endopod segment with 3 inner spines. Basipod having an
inner and outer spine with 1 adhesion pad. Leg 2 (fig. 53) with both
rami 2-segmented. First exopod segment with 1 outer spine and
patch of fine spinules on outer distal half. Last segment with 9
spines, outer 4 spinose and inner 5 naked. First endopod segment
unarmed. Second endopod segment with 5 short spines at tip.
Leg 3 (fig. 54) with both rami 2-segmented. First exopod segment
with spine on outer distal corner. Second segment with 6 short
spines. First endopod segment unarmed. Second segment with 2
short terminal spines. Leg 4 (fig. 55) with both rami 1-segmented.
Exopod segment with 6 spines. One spine placed at midpoint of
outer edge suggesting a 2-segmented ramus. Endopod unarmed.
Leg 5 (fig. 56) consisting of naked spine and separate plumose seta.
Leg 5 located near point of attachment of spermatophores (see fig. 56).
Area of spermatophore attachment located at junction of abdomen
to genital segment and composed of a heavily sclerotized area with
shelf to receive neck of spermatophores, attaching in these areas
with necks crossing to opening of seminal receptacle on opposite side.
Ege strings long and of usual generic type.

Color in life creamy yellow with light brown pigmentation as indi-
cated in figure 46.

Maue.—Body form as in figure 57. Total length based on 1 speci-
men, 9.5 mm, including caudal ramus but not setae. Greatest width
4.5 mm, measured at widest part of cephalon. Cephalon rounded,
slightly longer than wide (4.9 by 4.4 mm).

First thoracic segment fused with head. Segments 2-4 free without
dorsal plates. Genital segment longer than wide, 2.2 by 1.7 mm with
spermatophore visible within. Abdomen of 2 segments. First
segment 3604 long. Second segment 6124 long. Caudal ramus
(fig. 58) bearing 6 setae, inner 4 long and plumose, inner margin of
caudal ramus bearing row of hairs.

Oral area similar to that of female. In adult male adhesion pad
associated with second antenna only about one-half length of pad of
first antenna (264:576u) (see fig. 59). Collected with this mature
male was also a young male (7.1 mm) with spermatophores developing

SSS eee en

NO. 3570 PANDARIDAE—CRESSEY 1

within genital segment (by itself, this might have been mistaken for
fully developed male). Examination of its appendages showed it to
be identical with its larger counterpart, except pad of second antenna
is about as long as pad of first antenna (408:420z) (see fig. 60). Other
species of this genus with same reduction of adhesion pad of second
antenna in mature males.

Discussion.—Pandarus floridanus is closely related to P. sinuatus
but the females can be readily separated on the basis of the following
differences. ‘The dorsal abdominal plate of P. floridanus is consider-
ably longer than the dorsally exposed caudal ramus while in P. sinuatus
they are about equal in length (see table 1). In P. florrndanus the

TasBLE 1.—Comparison of the length of the dorsal abdominal plates with the
length of dorsally exposed caudal ramus in some specimens of Pandarus jflori-
danus with P. sinuatus (CHML=Cape Haze Marine Laboratory, Fla.)

Specimens Plate Ramus

P. floridanus
ex Carcharodon carcharias (avg. of 4) Dennis, Mass. 702u 450yu
(avg. of 2) USNM 107299 810u 576
ee : (1 spec.) USNM 101876 7220p 540u
ex Lamna cornubica (1 spec.) USNM 32765 720u 360u

P. sinuatus

ex Negaprion brevirostris (avg. of 2) CHML 495p 468u
ex Carcharinus leucas (avg. of 2) CHML 621yu 630u

dorsal thoracic plate of segment 2 extends beyond the middle of the
plate of segment 4 while in P. sinuatus the plate of segment 2 does
not extend to the middle of the plate of segment 4. P. floridanus is
not so heavily pigmented as P. sinuatus. Young females of P. sinuatus
first show pigmentation in the lateral areas of the cephalon (see fig. 66),
whereas P. floridanus is not pigmented in this area at all (its young
females showing first areas of pigmentation to be the anterior margin
of the cephalon).

From Latin, the word floridanus refers to the fact that the species
was seen first on a shark from Florida.

Pandarus floridanus can be separated from all other species of the
cranchwi group on the basis of the spine and setal formula. The male
of P. floridanus is similar to that of P. sinuatus. Since only 1 mature
male of the new species was available, a more detailed comparison
could not be made.

This species seems to be a well-established parasite on the body
surface of Carcharodon carcharias since it was recovered from that host

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

on four different occasions in the Western Atlantic. It should also be
noted as having been collected from Lamna nasus as well.

Pandarus sinuatus Say, 1817
Ficures 67, 68

Pandarus sinuatus Say, 1817, p. 486.—Milne-Edwards, 1840, p. 470.—Smith,
1874, p. 283.—Rathbun, 1886, p. 310.—McClendon, 1906, p. 44; 1907,
p. 114; 1910, p. 229.—Wilson, 1907, p. 417; 1932, p. 437.—Bere, 1936, p.
595.—Carvalho, 1940, p. 283; 1945, p. 111; 1951, p. 139.—Pearse, 1952a,
p. 27; 1952b, p. 218.—Causey, 1953, p. 12; 1955, p. 6.

Pandarus affinis Beneden, 1892a, p. 224.—Bassett-Smith, 1899, p. 467.— Wilson,
1907, p. 394.—Capart, 1953, p. 660.

SPECIMENS stTuDIED.—Two collections from Carcharinus leucas
Miiller and Henle from Sarasota, Fla. From the same locality a single
collection from Negaprion brevirostris (Poey). In addition to these, I
examined 88 collections from the U.S. National Museum collected at
various localities along the eastern coast of the United States. These
collections are from a wide variety of shark hosts generally restricted
to inshore species.

FrmaLe.—Body form as in figure 67. Total length (based on an
average of 5 specimens) 6.5mm. Greatest width (measured at widest
part of cephalon) 3.2 mm. Appendages of this species inseparable
from those of P. floridanus except last segment of exopod of leg 2 bears
10 spines instead of 9 as in P. floridanus. Since a description of ap-
pendages of this species would only duplicate that of P. floridanus,
none will be given here. Only those features that separate it from
P. floridanus will be emphasized.

Dorsal plate of segment 2 extending only to anterior third of plate
of segment 4. Abdominal plate small and rounded, only about as long
as exposed caudal ramus (see table 1). Color dark brown to black
with eye spots fused and continuous with median unpigmented line
from posterior edge of eye spots to posterior margin of cephalon.
Young females (fig. 68 )with pigment first developing in lateral areas
of cephalon.

Maur.—As in P. floridanus.

Discusston.—P. sinatus is apparently found only on sharks inhab-
iting coastal waters of the western North Atlantic Ocean. It occurs
only on the body surface of the host usually in clusters on the fins as
do other members of the genus. It is closely related to P. floridanus
but is easily separated from it on the basis of the above characteristics.
Superficially, this copepod also may be confused with P. bicolor because
of the pigmentation, but it can be easily distinguished from this species
on the basis of the thoracic plates. The dorsal plates of segment 2 of
P. bicolor do not extend beyond the plate of segment 3.

NO. 3570 PANDARIDAE—CRESSEY 17

Pandarus katoi, new species
Fiaures 69-96

SPECIMENS STUDIED.—Thirty females and 10 males from Carcharinus
malpeloensis (Fowler) from the Pacific (09°55’N, 85°51’W). Holotype
female, allotype male, and 14 paratypes (102 2 and 40’) in alcohol
deposited in the U.S. National Museum. Fourteen paratypes (109 9
and 407”) deposited in the British Museum (Natural History) and
the remaining paratypes in the author’s collection. Additional speci-
mens studied: 4 collections from C. malpeloensis from the Pacific
Ocean (09°42’N, 85°51’W; 21°20’N, 106°50’W; 10°03’N, 85°53’W;
08°25’N, 83°45’W), 2 collections from C. azureus from the Pacific
(9°52’N, 85°29’W; 10°09’N, 86°04’W), and a single collection from
the Cocos Islands from C. platyrhynchus.

Frma.Le.—Body form as in figure 69. Total length (based on an
average of 5 specimens) 5.6 mm. Greatest width (measured at the
widest part of the cephalon) 3.8 mm. Cephalon somewhat rounded,
3.0 by 3.8 mm. First thoracic segment fused to head. Dorsal
thoracic plates present on segments 2-4. Plates of segment 2 extend-
ing beyond those of segment 3 and to about middle of plates of seg-
ment 4. Plates of segment 2 are generally slightly divergent. Plates
of segments 3 and 4 fused at their bases. Genital segment 2.3 by
2.3mm. Abdomen 1-segmented (fig. 70), with dorsal plate. Dorsal
plate 3.0 by 2.3 mm, slightly longer than wide. Caudal ramus (see
fig. 70) 720u long and 270u wide bearing 4 spines and 2 plumose setae.

Oral area as in other members of genus. Adhesion pads associated
with first antennae, second antennae, maxillipeds, and a pair at ventral
distal corners of cephalon. First antenna (fig. 71) 2-segmented.
First segment 600y long and bearing 13 naked setae (fig. 72). Second
antenna (fig. 73) 3-segmented. Terminal segment with 2 spines, in
form of stout claw. Figure 73 shows adhesion pad (dotted line) in
relation to second antenna. Mouth tube of typical pandarid type.
Mandible (fig. 74) within tube. First maxilla (fig. 75) with a terminal
claw, 3 short setae, and located near base of mouth tube. Second
maxilla (fig. 76) armed asin figure. Maxilliped (fig. 77) terminal claw
with spatulate tip and opposed by 1 large adhesion pad on basal
segment.

Legs 1-4 biramose, with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 0 0:0 1:0 0:0 VII I
seg. 2 VI III xX IV VI II

Leg 1 (fig. 78) with both rami 2-segmented. First exopod segment

with outer fringed spine. Last exopod segment with 6 terminal

spines. First endopod segment with adhesion pad. Last endopod
221-534—67__2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

segment with 3 inner spines. Leg 2 (fig. 79) with both rami 2-seg-
mented. First exopod segment with outer spine. Last exopod
segment with 10 spines (outer 4 articulated, inner 6 not). First endo-
pod segment unarmed. Last endopod segment with 4 naked spines.
Leg 3 (fig. 80) with both rami 2-segmented. First exopod segment
with outer spine. Last exopod segment with 6 spines. First endopod
segment unarmed. Last endopod segment with 2 terminal spines (all
spines on rami of leg 3 naked). Leg 4 (fig. 81) with both rami 1-segmented.
Exopod with 7 spines. Endopod with 1 short spine. All spines
naked. Leg 5 (fig. 82) with outer single spine and inner lobe with 3
spines. Leg 5 located near point of attachment of abdomen (see
fig. 70). Area of attachment of spermatophore at junction of abdo-
men and genital segment, composed of a heavily sclerotized area with
shelf to receive neck of spermatophore. Spermatophores crossing
with neck penetrating opening of the seminal receptacle of opposite
side. Egg strings long and composed of single strands of eggs.

Color in life creamy yellow with light brown pigmentation as in
figure 69.

Maur.—Body form as in figure 83. Total length, based on 5 speci-
mens, 7.4mm. Greatest width 4.2 mm measured at widest part of
cephalon. First thoracic segment fused with head. Segments 2-4
free and without dorsal plates. Genital segment somewhat longer
than wide, 1.98 by 1.72 mm with spermatophores visible within.
Abdomen 2-segmented. First segment 300u long. Second segment
410u long. Caudal ramus (fig. 84) with 6 setae, the inner 4 long and
plumose. Inner margin of caudal ramus bearing row of hairs.

Oral area similar to that of female. As in P. floridanus, adhesion
pad of second antenna considerably smaller than one associated with
first antenna (204:516y) (see fig. 85). First antenna (fig. 86) as in
female except spines more heavily barbed. Second antenna (fig. 87)
4-segmented with adhesion pad on basal segment. Terminal segment
in form of claw with 2 median spines. Mouth tube and mandible as
in female. First maxilla (fig. 88) similar to that of female except
terminal claw has a more striated surface. Second maxilla (fig. 89)
as in female. Maxilliped (fig. 90) with terminal claw pointed rather
than spatulate as in female. Opposing adhesion area composed of
3 pads instead of 1 as in female.

Legs 1-4 not modified as in female and with spine and setal formula
as follows:

leg 1 leg 2 leg 3 leg 4
end. exp. end. exp. end. xp. end.
seg. 1 1:0 0:0 esi O:1 fei 0:1 Tey O:1
seg. 2 IV 3 Li Gas TVs5)) 56 IN Say 8)

Legs 1-4 biramose and each ramus 2-segmented. All setae plumose.
Leg 1 (fig. 91) with adhesion areas as in the figure. Leg 2 (fig. 92)

No. 3570 PANDARIDAE—CRESSEY 19

with outer edge of exopod first segment with long hairs and short
spinules. Leg 3 (fig. 93) with outer edge of exopod first segment
bilobed. Leg 4 (fig. 94) with outer edge of exopod first segment
similar to that of leg 3. Leg 5 (fig. 95) situated on midlateral margin
of genital segment and consisting of a lobe bearing 4 spines, 1 naked
and 3 smaller and plumose. Leg 6 (fig. 96) situated at point of
attachment of abdomen to genital segment and consisting of lobe
bearing 3 spines (1 plumose and 2 naked). Color in life cream,
generally devoid of pigment.

Discussion.—This species thus far has been collected only from
carcharinid sharks of the western central Pacific Ocean. It is closely
related to P. zygaenae but can be separated from this species by the
nature of the dorsal thoracic plates. The plates of the second thoracic
segment of P. katoi extend only to about the middle of the plate of
the fourth segment, whereas in P. zygaenae these plates extend to the
distal end of the plate of segment 4. Since the appendages of these
2 species are identical, it seems probable that they have evolved
from a common ancestor and thus are very closely related. Because
I was able to examine several collections of each species (10 collections
of P. katoi, 4 collections of P. zygaenae), a good sample for comparison
was available. In each case the copepods are separated easily on the
aforementioned characters.

I have named this species for Mr. Susumu Kato, who originally
collected the type material and who has generously collected in the
Pacific area for me.

Pandarus zygaenae Brady, 1883
Figures 97-99

Pandarus zygaenae Brady, 1883, p. 134.—Bassett-Smith, 1899, p. 467.— Wilson,
1907, p. 416.

SPECIMENS STUDIED.— Collections from the following areas: Off Sao
Paulo, Brazil, from Sphyrna zygaena (Linnaeus); Pacific (9°47’ N,
85°48’ W) from Sphyrna species; Pacific (18°31’ N, 109°34’ W) from
Sphyrna zygaena.

FrmaLe.— Body form as in figure 97. Total length (based on an
average of 2 specimens) 7.6 mm. Greatest width (measured at the
widest part of the cephalon) 3.9mm. Cephalon somewhat truncated
and measuring 3.7 by 3.9 mm, somewhat wider than long. Thoracic
plates present on segments 2-4. Plates of segment 2 extending
beyond those of segment 3 and to distal border of plate of segment
4. Genital segment somewhat longer than wide (2.7 by 2.3 mm).
Abdomen 1-segmented and with dorsal plate. Caudal rami as in
P. katot except that they are generally held parallel to each other.

Oral area and all appendages like those of P. katoi with the follow-

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

ing 2 exceptions: tip of maxilliped of P. zygaenae (fig. 98) shaped
somewhat differently; endopod of P. zygaenae unarmed; spine and
setal formula, otherwise, same as P. katov.

Color in preserved material creamy white except for some light
brown pigmentation in mature females as indicated in figure 97.

Mauz.— Body form as in figure 99. Male indistinguishable from
male of P. katoi. Only 1 male of each species was collected. More
material of both species might show some differences between the
males of these 2 species. Total length of male of P. zygaenae (based
on a single specimen) 8.1 mm (length not including setae on caudal
rami). Greatest width 4.6 mm.

Discusston.—Brady described this species in 1883 from Zygaena
malleus collected near Cape Verde Islands. In 1907 Wilson placed
the species in synonomy with Pandarus satyrus. Wilson examined
2 females from Brady’s original collection but noted that the cope-
pods were so covered with ‘‘fish slime’ as to be difficult to study.
I received 2 collections of copepods from the same host with enough
material to ascertain that these were not P. satyrus but clearly a
separate species. A comparison of this new material with Brady’s
original description and figures prove his species to be valid. The
figure of the female by Brady appears to be of an immature form but
shows some features consistent with my recent collections, namely,
the relative lengths of the dorsal thoracic plates and the fact that
the caudal rami are generally held parallel to each other rather than
divergent as in most other members of the genus (in preserved
specimens).

This species is closely related to P. katoi but can be distinguished
from it by the relationship of the plates of segment 2 to those of
segement 4. P. zygaenae so far has been found only on the genus
Zygaena and may well be confined to species of hammerhead sharks.

Pandarus bicolor Leach, 1816
Figures 100-108

Pandarus bicolor Leach, 1816, p. 405; 1819, p. 535.—Desmarest, 1825, p. 339.—
Burmeister, 1833, p. 331.—Kr¢yer, 1837, p. 202; 1838, p. 34; 1863, p. 261.—
Milne-Edwards, 1840, p. 470.—Baird, 1850, p. 288.—Beneden, 185la, p. 94;
1861, p. 148.—Norman, 1868, p. 301.—Olsson, 1868, p. 21.—Richiardi, 1880,
p. 149.—Carus, 1885, p. 362.—Bassett-Smith, 1896, p. 156; 1899, p. 466.—
Brian, 1898a, p. 12; 1899, p. 3; 1906, p. 55; 1914b, p. 7; 1940, p.11.—Scott, T.,
1900, p. 157.—Seott, A., 1904, p. 40.—Norman and Scott, T., 1906, p. 211.—
Wilson, 1907, p. 400; 1932, p. 486; 1935b, p. 778.—Scott, T., and Scott, A.,
1913, p. 95.—Hansen, 1923, p. 36.—Scott, A., 1929, p. 95.—Leigh-Sharpe,
1934b, p. 112.—Pesta, 1934, p. 29.—Oorde and Schuurmans Stekhoven,
1936, p. 141.—Stephensen, 1940, p. 5.—Rose and Vaissiere, 1953, p. 86.—
Barnard, 1955, p. 257.—Nunes-Ruivo, 1956, p. 17.—Causey, 1960, p. 331.—
Heegaard, 1962, p. 177.

NO. 3570 PANDARIDAE—CRESSEY 21

Pandarus bosciz Leach, 1816, p. 406; 1819, p. 535.—Guérin-Meneville, 1829-
1843, p. 41.—Burmeister, 1833, p. 331.—Desmarest, 1825, p. 339.—Krgyer,
1837, p. 202.—Baird, 1850, p. 289.

Caligus bicolor—Lamarck, 1818, p. 137.

Pandarus fissifrons.— Milne-Edwards, 1840, p. 470.

Pandarus lividus—Frey and Leuckart, 1847, p. 166.

SPECIMENS STUDIED.—Three collections of females from Squalus
acanthias Linnaeus caught at the following locations: in the North Sea,
eastern North Atlantic (53°04’ N, 04°02’ E), and off the coast of the
Netherlands.

FrmaLe.—Body form as in figure 100. Total length (based on an
average of 2 specimens) 9.1 mm. Greatest width (measured at the
widest part of the cephalon) 3.8mm. Cephalon only about one-third
body length, measuring 3.6 by 3.8mm. Dorsal thoracic plates present
on segments 2-4. Plates of segment 2 short and not extending beyond
the posterior edge of plates of segment 3. Plate of segment 4 fully
exposed. Abdomen 1-segmented (see fig. 101) and covered by broad
dorsal plate. Caudal ramus (fig. 101) broad and much shorter than
in other species of genus. Caudal rami not obvious in dorsal view.
Each ramus armed with 5 short spines.

Oral area as in other members of genus. Since this species has been
well reported and amply figured in the literature, only salient features
included here. Second antenna (fig. 102) with short claw at tip
separated from last segment. In other species of genus these 2
elements are often fused. Maxilliped (fig. 103) with spatulate process
at tip of claw.

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
p. end. exp. end. exp. end. exp. end.
seg. 1 1-0) * 0:0 1:0 0:0 I:0 0:0 Vii oe
seg. 2 VI Ill VS Vi VI II

Legs not notably different from other species of genus with exception
of reduction in number of spines on last segment of exopod of leg 2
(8 instead of 10) and elongated nature of exopod of leg 4. Legs 1-4
as in figures 104-107. Leg 5 (fig. 108) with outer single spine and
inner lobe with 3 spines.

Color cream yellow to dark brown in pigmented areas. Cephalon
usually heavily pigmented with eye spots not separated and often
joined by median clear stripe extending distally.

Maure.—No material available for study. Apparently male of this
species is rare, as no collection examined contained one. It is interest-
ing to note that males of P. carcharini and P. niger (probably closely
related to P. bicolor) are unknown.

T. and A. Scott (1913) illustrate male of P. bicolor and their figures
show it to be much the same as other males of the genus.

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Spine and setal formula (based on T. and A. Scott’s illustrations)

are as follows:
leg 1 leg 2 leg 3 leg 4
p.

end. exp. end. exp. end. exp. end.
seg. 1 1:0 OO pea al. 0:1 (not illustrated by the Scotts)
seg. 2 IV:3 3 IV:6 8

Remarks.— This copepod seems to be limited to the waters of the
eastern Atlantic and is the primary species of Pandarus found in the
coastal waters of Europe. It seems to be restricted to smaller inshore
species of sharks and, as far as recorded in the literature, is found only
on the body surface of its host.

Pandarus bicolor can be easily separated from the other known
species of the genus on the basis of the caudal rami. The relative
lengths of the dorsal thoracic plates of segments 2 and 3 separate it
from all species of the cranchit group. This copepod has twice been
confused by American authors (Wilson, 1932, and Causey, 1960) with
P. sinuatus because of the presence of a median stripe from the eye
spots to the distal border of the cephalon of both species. I have
examined specimens in the U.S. National Museum identified by Wilson
as P. bicolor and they are clearly P. sinuatus.

Finally, I have placed Pandarus lividus Frey and Leuckart in
synonymy with P. bicolor. Wilson (1907) commented that it was
impossible to separate the 2 species on the basis of Frey’s and
Leuckart’s description but he did not place it in synonymy. Frey
and Leuckart gave no figures of P. lividus.

Pandarus niger Kirtesinghe, 1950
Fiaures 109-118
Pandarus niger Kirtesinghe, 1950, p. 83.

SPECIMENS sTupDIED.—A female paratype loaned to me by Kirte-
singhe, and a single collection of 6 females from Galeorhinus species
in Formosa.

FEMALE.—Since this species has been described recently by Kirte-
singhe (1950), I will be concerned with only those features taxonom-
ically important to the genus. Body form as in figure 109. Total
length 8.2 mm (based on an average of 2 specimens). Greatest
width (measured at the widest part of the cephalon) 3.8mm. Cephalon
somewhat truncated only about one-half as wide across anterior third
as across posterior third. First thoracic segment fused to head.
Dorsal thoracic plates present on segments 2-4. Plates of segments
2 and 3 fused basally with posterior borders more or less in a straight
line across proximal portion of plate of segment 4. Plate of segment 4
with posterior border nearly straight with only a very slight medial
indentation. Genital segment as in other species of genus. Abdomen

NO. 3570 PANDARIDAE—CRESSEY 23

1-segmented and with dorsal plate. Dorsal plate small, extending
only slightly beyond posterior border of abdomen (see fig. 110).
Caudal ramus (fig. 110) long (1.8 mm) and well exposed dorsally.

Oral area as in other members of genus. First antenna (fig. 111)
2-segmented, bearing 22 spines on first segment and 10 naked setae
on second segment. Second antenna (fig. 112) with tip produced into
a short claw similar to that of P. bicolor. First maxilla (fig. 113) with
group of short spines on basal third composed of only 2 spines rather
than usual 3. Second maxilla (fig. 112) with distal setae in form of
fringed claw.

Legs 1-4 (figs. 115-118) biramose and with spine and setal formula
as follows:

leg 1 leg 2 leg 3 leg 4
xp. end. exp. end. exp. end. exp. end.
seg. l 1:0 0:0 1:0 0:0 1:0 0:0 VEE TLE
seg. 2 VI Ill x VII VI Ill

Leg 5 (see fig. 110) with an inner lobe and an outer long seta.

Color dark brown except where unpigmented. Eye spots separated
in older females.

Maue.—Unknown. Kirtesinghe (1950) illustrates and briefly de-
scribes an immature male but not enough is shown to be of value in
an analysis of the adult.

ReMArRKS.—So far this copepod has not been reported from areas
other than Ceylon and Formosa. It may well be a parasite restricted
to the Indian Ocean and western Pacific. Not enough is known to
ascertain its host preferences but inshore carcharinid sharks seetn to
be preferred hosts.

This copepod is closely related to P. bicolor, but the nature of the
caudal ramus separates it easily from all other members of the genus.
Kirtesinghe (1950) notes the relationship between this species and
P. bicolor; he also suggests an affinity with P. satyrus on the basis of
the caudal ramus. A comparative survey of the genus shows this
not to be so, especially on that basis since the caudal ramus of P.
satyrus does not extend beyond the tip of the abdominal plate.

Pandarus carcharini Ho, 1960
Figures 119-124
Pandarus carcharini Ho, 1963, p. 93.

SPECIMENS STUDIED.—Paratype female from Formosa loaned to
me by Ho. Also 2 more collections of females from Formosa from
Galeorhinus species and Carcharinus gangeticus. Three collections
from Nosy Bé, Madagascar, from Carcharinus leucas, C. sorrah, and
. limbatus. A single collection from Durban, South Africa, from C.
eucas.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Frema.e.—A full description of this species will not be given here
since it has been well described recently by Ho (1960). Body form
as in figure 119. ‘Total length 9.8 mm (based on an average of 3
specimens). Greatest width 4.9 mm (measured at the widest part of
the cephalon). Cephalon somewhat truncated but not as narrowed
anteriorly as P. niger. First thoracic segment fused with head.
Dorsal thoracic plates on segments 2-4 with plates of segments 2 and
3 fused basally, posterior borders forming a straight line as in P.
bicolor and P. niger. Some of the salient features have been figured
for the sake of comparison. Caudal rami (fig. 120) intermediate
between that of P. bicolor and P. niger, measuring 1.26 mm in length
and extending posteriorly only about as far as the dorsal abdominal
plate.

Oral area as in P. niger. Appendages of cephalon as in P. niger
except that first antenna (fig. 121) has 24 spines on first segment and
10 naked setae on last.

Legs 1-4 biramose, with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4

exp. end. exp. end. exp. end. exp end.
seg. 1 1:0 0:0 1:0 0:0 LO 00: AVAL Tet
seg. 2 VI Ill xX VI VII II

Legs 2-4 as in figs. 122-124. Leg 5 as in P. niger.

Color creamy yellow with dark brown pigmentation as in figure 119.
Eye spots fused. Egg strings of usual type.

Mate.— Unknown.

Remarks.—This species described by Ho (1960) from Formosa has
been collected by the author in Nosy Bé, Madagascar, from carchari-
nid sharks. Its range may well be the same as P. niger but not enough
material has been collected on which to base a definitive conclusion;
nevertheless, from what is known, it appears to be a parasite of in-
shore species of sharks in the Indian Ocean and western Pacific coast.

Pandarus carcharini may be separated from its closely related
species (P. bicolor and P. niger) on the basis of the caudal ramus and
the spine and setal formula. Ho (1960) has already cited the similar-
ities between those 3 species. His comparison and description
implies the presence of a ventral abdominal plate. I could find no
evidence of a ventral plate and, after personal communication with
Ho, he agrees that this is synonomous with the abdomen and that
the use of the term “‘plate’’ is invalid in this case.

Genus Phyllothereus Norman 1903

Phyllophora Milne-Edwards, 1840, p. 471. [Type-species: P. cornutus.]
Nogagus.—Steenstrup and Lutken, 1861, p. 386. [Refers to N. grandis only.]
Laminifera Poche, 1902. [Cite Wilson, 1907, p. 361.]

NO. 3570 PANDARIDAE—CRESSEY 95

Phyllothereus Norman, 1903, p. 368.
Parapandarus Wilson, 1924a, p. 7.

Fremate.—Frontal plate distinctly separate. First thoracic seg-
ments 2-4 free and possessing dorsal plates. Dorsal plate of segment
3 extending well beyond distal margin of plates of segment 2. Plates
of segment 4 separated. Abdomen 1-segmented and with a dorsal
plate. Abdomen or its plate visible dorsally and attached to distal
end of genital segment. Caudal rami lateral to abdomen. First
antenna 2-segmented. Claw of maxilliped pointed at tip. Adhesion
pads absent or much reduced. Legs 1-4 biramose, rami of legs 1-3
2-seemented, those of leg 4 1-segmented. All legs without plumose
setae. Leg 5 present. Egg sacs long and straight.

Maue.—No dorsal plates present. Cephalic appendages in general
like those of female. Legs 1-4 biramose, all rami 2-segmented bearing
plumose setae. Fifth and sixth legs present. Leg 3 endopod un-
modified. Abdomen 2-segmented. Caudal rami held at distal end
of abdomen. Adhesion pad with first antenna.

Discussion.—A great deal of confusion has existed over the
synonymy of this genus. The genus was first described by Milne-
Edwards in 1840 as Phyllophora with P. cornutus as the type-species.
Later, in 1861, Steenstrup and Lutken described the male as Nogagus
grandis. Norman (1903) cited the fact that Phyllophora is thrice
preoccupied and changed the generic name to Phyllothereus. Wilson
(1907) discussed the preoccupation of the name Phyllophora and
stated that Poche in 1902 suggested changing the name to Laminzfera.
This reference to Poche (1902) is in error since there is no reference
to the name Laminifera by this author in 1902 or in subsequent works
(see Brian, 1946). Brian synonymized his Laminifera doello-jurador
with P. cornutus in 1946; consequently, the valid name of the genus
should be Phyllothereus as proposed by Norman in 1903. Wilson
in 1932 states that his Parapandarus is so close to Phyllothereus that
the 2 may be identical. I have examined Wilson’s types of
Parapandarus and have concluded that they are indeed the same;
consequently, P. nodosus Wilson 1924 is synonymous with Phyllothereus
cornutus (Milne-Edwards, 1840).

Phyllothereus cornutus (Milne-Edwards, 1840)
Figures 125-138

Phyllophora cornuta Milne-Edwards, 1840, p. 372.

Nogagus grandis Steenstrup and Lutken, 1861, p. 386.

Phyllophorus cornutus—Bassett-Smith, 1899, p. 465.

Phyllothereus cornutus—Norman, 1903, p. 368 Norman and Scott, 1906, p. 212.—
Scott, T., and Scott, A., 1913, p. 92.— Wilson, 1932, p. 440.—Monod and Doll-
fus, 1938, p. 196.—Brian, 1946, p. 142.

Parapandarus nodosus Wilson, 1924a, p. 7.

Laminifera doello-juradoi Brian, 1944, p. 193; 1946, p. 142.

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

SPECIMENS sTUDIED.—Three collections from Prionace glauca in the
Indian Ocean (0°58’N, 55°00’E; 0°14’N, 55°00’E; 6°37’S, 55°00’E).
Five collections from Prionace glauca at various points in the western
North Atlantic.

FEMALE.—Body form as in figure 125. Average length of North
Atlantic specimens 14.6 mm (based on an average of 5 specimens).
Greatest width of North Atlantic specimens 10.3 mm. Average
length of Indian Ocean specimens 10.3 mm (based on 4 specimens).
Greatest width of Indian Ocean specimens 5.6 mm. Dorsal thoracic
plates on segments 2-4. All thoracic plates paired and often greatly
inflated in preserved specimens. Genital segment (fig. 126) pear
shaped, being widest in posterior third. Abdomen 1-segmented,
large, and bearing a dorsal plate that does not cover it when viewed
dorsally. Caudal rami as in figure 126 with 4 short spines.

Oral area without adhesion pads. First antenna 2-segmented, first
segment with 22 naked spines and last segment with 12 naked spines.
Second antenna (fig. 127) with long terminal claw. When second an-
tenna is flattened against oral area, claw extends posteriorly to maxil-
lipeds. Terminal claw bearing 2 short spines. First maxilla (fig. 128)
with short stout terminal spine and group of 3 short setae on basal
third. Second maxilla (fig. 129) with usual fringed claw and 2 setae at
base, 1 large and fringed, other shorter and plumose at tip. Maxil-
liped (fig. 130) bearing terminal claw with pointed tip and opposed on
penultimate segment by a ridge.

Legs 1-4 (fig. 131-134) biramous with rami of legs 1-3 2-segmented
and those of leg 4 1-segmented. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 1:0 0:0 ESO 0:0 WAN EE JOBE
seg. 2 VII Iil VII VII WAGne. In

Leg 5 (fig. 135) consisting of an inner lobe and single outer seta.

Egg strings long and each composed of a single strand of disclike
egos.

Color whitish yellow.

Mate.—Since the male of this copepod has been well figured and
described by Monod and Dollfus (1938), only a few salient features
will be mentioned here. Body form as in figure 136. Oral area as
in female with the following differences: second antenna smaller, in
relation to rest of oral area, than in female; maxilliped with a claw at
tip but opposed by 2 adhesive areas rather than 1 as in female (see
Monod and Dollfus, 1938, p. 198, fig. 5).

Legs 1-4 biramose and not as highly modified as in female. Spine
and setal formula as follows:

NO. 3570 PANDARIDAE—-CRESSEY D7

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 30 0:1 1:0 O:1 Irate ea
seg. 2 TVeSie ie LVE'Ga as VirSyn G LV i500

Leg 5 (fig. 137) located at beginning of distal third of genital segment
and consisting of a ventral lobe bearing 1 outer plumose seta, a group
of 2 plumose setae and a naked spine at distal end of lobe. Leg 6
(fig. 138) located on posterior border of genital segment near junction
of abdomen and consisting of an outer plumose seta and an inner
naked spine.

Discussion.—Phyllothereus cornutus is a parasite on the gills and
gill arches of a number of species of sharks, but it is most commonly
found associated with Prionace glauca. It is cosmopolitan in distribu-
tion and has been collected by the author in both the Indian Ocean
and the North Atlantic. The Indian Ocean specimens were smaller
than those collected in the North Atlantic, but a detailed examination
of their appendages showed no important differences. Since the same
host was involved in both cases, I have concluded that they are of
the same species. More collections in intermediate areas might well
yield intermediate sizes. The nature of the inflated dorsal plates
seems to be a function of the preserving fluid (70 percent alcohol or
10 percent formalin) as this condition has not been observed by me
in life.

Genus Gangliopus Gerstaecker, 1854
Gangliopus Gerstaecker, 1854, p. 189. [Type-species: G. pyriformis.]

Frema.e.—Frontal plate distinctly separate. First thoracic segment
fused with cephalon. Thoracic segments 2-4 free and possessing
dorsal plates. Dorsal plates of segment 2 small. Plate of segment 3
extending beyond tip of plate of segment 2. Plates of segment 4
partially separated. Abdomen 1-segmented, with a dorsal plate.
Abdomen or its plate visible dorsally and attached to distal end of
genital segment. Caudal rami lateral to abdomen. First antenna
2-segmented. Claw of maxilliped pointed at tip. Adhesion pads of
the first and second antenna absent or reduced. Pad of maxilliped
well developed. Legs 1-4 biramose, rami of legs 1-3 2-segmented,
those of leg 4 1-segmented. All legs without plumose setae. Leg 5
present. Egg sacs long and held straight.

Mauze.—No dorsal plates present. Cephalic appendages like those
of female. Adhesion pad with first antenna. Legs 1-4 biramose,
all rami 2-segmented and bearing plumose setae. Fifth and sixth
legs present. Leg 3 endopod unmodified. Caudal rami held at
distal end of abdomen.

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Gangliopus pyriformis Gerstaecker, 1854
Figures 139-146

Gangliopus pyriformis Gerstaecker, 1854, p. 192.—Wilson, 1907, p. 350.—Monod
and Dollfus, 1938, p. 204.

Nogagus angustulus Gerstaecker, 1854, p. 193.

Gangliopus tetrapturt Yamaguti and Yamasu, 1960, p. 142.

SPECIMENS sTuDIED.—Three collections from Prionace glauca in
the Indian Ocean (0°58’N, 55°00’E; 0°14’N, 55°00’E; 6°37’S, 55°00’E).
One collection from the same host in the North Atlantic (39°32’N,
28°02/W).

FrmaLte.—Body form as in figure 139. Total length 9.1 mm
(based on an average of 3 specimens). Greatest width 4.9 mm
(measured at widest part of the cephalon). Dorsal thoracic plates
on segments 2-4. Plates of segment 2 small and not extending to tip
of plates of segment 3. Plates of segments 3 and 4 each with a deep
median incision. Genital segment nearly square. Abdomen 1-
segmented and joining genital segment at posterior border. Abdomen
with a small dorsal plate. Caudal rami small and held at lateral
angles of abdomen.

Oral area of usual pandarid type. No adhesion pad with first
antenna. Adhesion pad of second antenna small and inconspicuous.
Adhesion pad of maxilliped large and L-shaped (fig. 140).

Monod and Dollfus (1938) have given a detailed description of
female appendages.

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 1:0 0:0 30 0:0 VII II
seg. 2 VII Ill xX VI Ve 1CT eee Tene

Leg 5 (fig. 141) consisting of a single finger-like spine with 2 short
setae. Fifth leg near junction of abdomen and genital segment.

Egg strings long and each composed of a single strand.

Color in life pale yellow and without pigment.

Matzr.—Body form as in figure 142. Total length 7.4 mm (based
on 1 specimen). Greatest width 3.5 mm (measured at widest part of
cephalon). Thoracic segments without dorsal plates except for a
winglike expansion on the posterolateral corners of segment 2.
These extend nearly to end of segment 3. Genital segment (fig.
143) nearly square, 1.75 mm long by 1.85 mm wide. Genital segment
with each posterior corner produced into broad lobe. Abdomen
2-segmented. Caudal rami prominent, with 4 long plumose setae
and 2 smaller ones at each distal corner.

Oral area as in female except a small adhesion pad associated
with first antenna. Adhesion pad of maxilliped small and incon-

NO. 3570 PANDARIDAE—CRESSEY 29

spicuous. Cephalic appendages like those of female except for
maxilliped (fig. 144).
Legs 1-4 biramose and with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
end. exp. end. exp. end. exp. end.
seg. 1 I:0 0:0 ot 0:1 et 0:1 | ia 0:1
seg. 2 LVS. 3 IV:6° "8 IV:5 6 TV 25". +5

Leg 5 (fig. 145) located on midlateral edges of genital segment and
consisting of 3 plumose setae and 1 short stout spine. Leg 6 (fig.
146) located near junction of abdomen and genital segment and
composed of a single plumose seta and a short spine.

RemarKks.—Gangliopus pyriformis is apparently cosmopolitan in
distribution and occurs commonly on the gills of the blue shark,
Prionace glauca. In 1960 Yamaguti and Yamasu described a new
species Gangliopus tetraptert. Since they do not refer to the work of
Monod and Dollfus (1938), it is assumed that they were not aware
of it. They based their new species on morphological differences
which do not exist. Gerstaecker originally described the female
as having a 2-segmented abdomen. His original description was very
superficial Monod and Dollfus (1938) updated the work and
provided a working description. Gangliopus tetrapteri does not differ
from this later description nor from my own collections in the Indian
Ocean. The male was described under the name Nogagus angustulus
by Gerstaecker at the same time as the female. No description of
the male has appeared since then. I have added salient features
to this original description.

Genus Pseudopandarus Kirtesinghe, 1950

Pseudopandarus Kirtesinghe, 1950, p. 84. [Type-species: P. gracilis.]
Pandarus.—Gnanamuthu, 195la, p. 1245. [Refers to P. longus only.]

Frma.te.—Frontal plate distinctly separate. First thoracic segment
fused with cephalon. Thoracic segments 2-4 free and possessing
dorsal plates. Dorsal plates of segment 2 small and widely separated.
Plate of segment 3 extending beyond tip of plates of segment 2.
Abdomen incompletely divided into 2 segments and with a dorsal plate.
Abdomen completely or partially concealed beneath genital segment.
Abdomen attached to genital segment ventrally. Caudal rami
attached terminally to abdomen. First antenna 2-segmented. Claw
of maxilliped with a spatulate tip. Adhesion pads present on cephalon.
Legs 1-4 biramose. Rami of legs 1-3 2-segmented, those of leg 4
l-segmented. Leg 5 present and consisting of a free segment. Egg
sacs long and straight.

Mauz.—I had no material of the male of this genus to study but

30) PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

the male has been described by Gnanamuthu (1951) and my diagnosis
is based on this description.

Body form of typical pandarid configuration. Cephalic appendages
like those of female except setae on first antenna are more plumose
in male. Legs 1-4 biramose with each ramus composed of 2 segments.
All setae plumose. Legs 5 and 6 present. Abdomen 2-segmented.
Caudal rami attached distally.

Remarks.—This genus is known only from the Indo-Pacific area
and consists of 2 species from the body surface of sharks of the genera
Carcharinus and Triakis. This genus may well be restricted to inshore
sharks.

Key to Females of Genus Pseudopandarus

Abdomen concealed in dorsal view, genital segment with posterior corners pro-
ducedinto long ‘pointed | processes! J.2)-2 6252 2) ee SO gracilis
Abdomen partially visible in dorsal view, genital segment with posterior corners
roundedwand notrereatly, produced. as <5 jase Een eee longus

Pseudopandarus gracilis Kirtesinghe, 1950
Figures 147-150

Pseudopandarus gracilis Kirtesinghe, 1950, p. 84.
Pseudopandarus scyllii Yamaguti and Yamasu, 1959, p. 124.

SPECIMENS STUDIED.—Paratypes on loan from P. Kirtesinghe.
6 females ex Scoliodon palasorrah, Nosy Bé, Madagascar.

FrmMaLe.—Body form as in figure 147. Total length 4.8 mm
(based on an average of 3 specimens). Greatest width 1.75 mm
(measured at widest part of the cephalon). Thoracic segments 2—4
with dorsal plates. Plates of segments 2 and 3 fused basally. Plates
of segment 2 small and entirely lateral to those of segment 3. Plates
of segment 4 covering proximal part of genital segment and fused
basally. Posterior edges of plates of segments 3 and 4 often irregular
in shape (I noticed this in both samples from Ceylon and Madagascar)
Genital segment long (3 mm) and posterior border produced to form
2 lateral projections and median finger-like process. Abdomen (fig.
148) joined ventrally to genital segment and concealed in dorsal view.
Abdomen indistinctly divided into 2 segments. A small dorsal plate
present on first segment of abdomen and not extending beyond
distal tip of second segment. Caudal ramus triangular in shape,
widest at distal end, bearing 3 terminal naked spines and outer
plumose seta. Two naked subterminal spines located at distal
corners (see fig. 148).

Since this species has been well described by Kirtesinghe (1950)
and Yamaguti and Yamasu (1959), only a few taxonomically im-
portant features of the appendages will be discussed here.

NO. 3570 PANDARIDAE—CRESSEY 31

Oral area with prominent adhesion pads with first and second
antenna. Adhesion pad of maxilliped small. Second maxilla (fig.
149) with shortest spine feathered at tip. Maxilliped (fig. 150)
spatulate at tip.

Legs 1-4 biramose and with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 1:0 0:0 1:0 0:0 V:3 2
seg. 2 Vis As BV5 2% IvV:4 3

Surface of segments of legs 1-4 generously covered with patches of
spinules. Leg 5 (see fig. 148) consisting of a free segment bearing
4 naked setae (in one specimen the 2 innermost setae were fused to
form a thick spine).

Egg strings long and straight.

Color in preserved specimens cream white with no pigmentation.

Maur.—Unknown.

Remarks.—Pseudopandarus gracilis was described in 1950 by P.
Kirtesinghe and was not reported again until 1959 by Yamacuti
and Yamasu. They determined their species to be new, based
primarily on the structure of the tip of the maxilliped. Unfortunately,
Kirtesinghe’s figure of this was misleading. Iwas fortunate to be able
to examine type material of P. gracilis, in which I found that there ac-
tually were no differences between the two species on this or any other
basis and thus have placed P. scyllii in synonymy. Yamaguti and
Yamasu define the fourth leg as having 2-segmented rami. Their
figure shows no articulation between segments. J have interpreted
the rami as being 1-segmented. They also refer to the plate of the
fourth segment as the structure that I have called the genital segment.
The plate of segment 4 actually covers the proximal portion of the
genital segment and is the same plate referred to by them as the plate
of segment 8. The plates of segments 2 and 38 referred to as one
plate belong to segment 2. A careful study of the copepod shows
this interpretation to be in error and not consistent with the usual
pandarid situation.

To date, this parasite has been found in the Indo-Pacific area only
(Madagascar, Ceylon, Japan), parasitic on the body surface of small
inshore sharks (Carcharinus and Triakis).

Pseudopandarus longus (Gnanamuthu, 1951)
Ficures 151-161
Pandarus longus Gnanamuthu, 1951a, p. 1245.—Kurian, 1955, p. 114.
SPECIMENS sTUDIED.—Four females ex Carcharinus obesus, 2 fe-

males ex Rhizoprionodon acutus, both sharks caught off Durban, South
Africa.

oo PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Frmaute.—Body form as in figure 151. Total length 4.3 mm
(based on 1 specimen). Greatest width 1.5 mm. Thoracic plates
on segments 2-4. Plates of segment 2 small and extending only to
about middle of plates of segment 3. Plate of segment 4 covering
proximal end of genital segment. Genital segment long (2.6 mm),
over one-half the length of body. Posterior border of genital segment
with a broad midian sinus. Abdomen (fig. 152) joined ventrally to
genital segment and partially concealed dorsally. Abdomen consist-
ing of 2 incompletely separated segments. Caudal rami held at
distal end of abdomen and bearing 4 terminal setae and 2 subterminal
setae on outer corners. Outer terminal seta finely plumose.

Oral area with prominent adhesion pads associated with first and
second antennae. Adhesion pad of maxilliped evident but some-
what reduced. First antenna (fig. 153) 2-segmented and of typical
pandarid type. First segment with 24 spines, most of which are
plumose. Last segment with 13 naked setae. Second antenna (fig.
154) with small claw at tip and a large adhesion pad. Mouth tube,
mandible, and first maxilla of usual type. Second maxilla (fig. 155)
with usual 3 spines at or near tip. Maxilliped (fig. 156) of usual
type and with broad spatulate tip on last segment.

Abdomen composed of 2 segments. Caudal rami attached distally.

Remarks.—This copepod was first described by Gnanamuthu in
1951 and assigned to the genus Pandarus. On the basis of the
diagnostic features of the genus Pandarus I have removed this species
and placed it in the genus Pseudopandarus Kirtesinghe, 1950. A
comparison of P. gracilis Kirtesinghe and P. longus (Gnanamuthu)
shows the following common features: the arrangement of the dorsal
thoracic plates, the spatulate process of the maxilliped, the prominent
first and second antennal adhesion pads with a reduction of the
maxilliped pad, the nature of the abdomen and its position in relation
to the genital segment, and the free segment of leg 5.

This species, like P. gracilis, seems to be Indo-Pacific in distribution
(Ceylon and Durban, South Africa). So far, it has been reported
only from carcharinid sharks.

Pannosus, new genus

Type-species: Gangliopus japonicus (Shiino, 1960).

Fremate.—Frontal plate distinctly separate. First thoracic seg-
ment fused with cephalon. Thoracic segments 2-4 free and possessing
dorsal plates. Dorsal plate of segment 3 extending well beyond
plates of segment 2. Plates of segment 4 extends well beyond end
of plate of segment 3. Plates of segments 3 and 4 fused basally.
Abdomen l-segmented. Caudal rami held laterally on abdomen.
Adhesion pads of cephalon well developed. First antenna 2-seg-

No. 3570 PANDARIDAE—CRESSEY a

mented. Claw of maxilliped spatulate. Legs 1-4 biramose, rami of
legs 1-3 2-segmented, those of leg 4 1-segmented. Egg sacs long
and straight.

Maur.—Unknown.

Remarks.—This copepod was described by Shiino in 1960 from a
single ovigerous female taken off a hammerhead shark (Sphyrna
zygaenae) by him. He assigned this new species to the genus Gang-
hopus. After a redescription of the generic diagnosis of this genus,
it became apparent that this new copepod did not belong to that
genus. Since I have had no material of this species to work with,
no further diagnosis of Shiino’s species can be made. JI have removed
this species from the genus Gangliopus for the following reasons:
(1) Shino’s species show well-developed adhesion pads associated with
the first and second antennae; (2) the dorsal thoracic plates of the
second segment are well formed in Pannosus; (3) the maxilliped of
japonicus has a spatulate tip whereas in Gangliopus the tip is pointed.
Shiino does not mention the presence of fifth legs but more material
would probably reveal their nature.

Since this copepod does not conform to any known genus, I propose
placing it in a new genus Pannosus.

The name Pannosus, from Latin, meaning “covered with rags,” refers
to the appearance of the dorsal plates.

Genus Perissopus Steenstrup and Liitken, 1861

Perissopus Steenstrup and Liitken, 1861, p. 393. [Type-species: P. dentatus.]
Chlamys Beneden, 1892a, p. 227.
Achtheinus Wilson, 1908, p. 450.

Frmaty.—Frontal plate not distinctly separate. First thoracic
segment fused with cephalon. Thoracic segments 2-4 free and
possessing dorsal plates. Dorsal plate of segment 2 extending to
posterior border of plates of segment 3. Plates of segment 4 covering
anterior portion of genital segment. Abdomen l-segmented and
attached ventrally to genital segment. Caudal rami rudimentary.
First antenna 2-segmented. Second antenna armed with teeth at tip.
Mouth parts of typical pandarid type. Adhesion pads present.
Maxilliped with small claw and large adhesion pad. Legs 1-4 biramose
but much reduced. Rami of legs 1 and 2 2-segmented. Rami of legs
3 and 4 1-segmented. All legs without plumose setae. Legs 5 and 6
present. Keg strings long and straight.

Maur.—Frontal plate separate. No dorsal plates present. Oral
area similar to female but with adhesion pads reduced. Maxilliped
with a strong terminal claw. Legs 1-4 biramose with each ramus
2-segmented. All setae plumose. Fifth and sixth legs present. Leg 3

221-534—67__3

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

endopod unmodified. Abdomen 2-segmented. Caudal rami attached
distally to abdomen.

Discussion.—This genus is represented by two species, Perissopus
dentatus Steenstrup and Liitken 1861 and P. oblongatus (Wilson, 1908).
A discussion of the history of the synonomy of these species is included
in the species descriptions.

Perissopus oblongatus (Wilson, 1908)

Achtheinus oblongatus Wilson, 1908, p. 450.
Achtheinus dentatus Wilson, 1911, p. 630.
Achtheinus pinguis Wilson, 1912, p. 235.

Achtheinus japonicus Wilson, 1922, p. 4,

Achtheinus parvideus Wilson, 1923, p. 7.

Achtheinus intermedius Kurtz, 1924, p. 614.
Achtheinus galeorhini Yamaguti, 1936, p. lett
Achtheinus platensis Thomsen, 1949, p. 20.
Achtheinus chinesis Thomsen, 1949, p. 23.
Achtheinus impenderus Shen and Wang, 1958, p. 27.

Discussion.—In 1908 Wilson described a new copepod and erected
for it a new genus, Achtheinus. Since then, 10 species assigned to this
venus have been described. A comparison of the description and
figures of these species show that they should be included in the genus
Perissopus. This is apparent when one compares the appendages of
the two groups. In both, the second antenna has a hoodlike process
bearing spines. The maxillipeds of each bears a reduced claw and the
basal segment is in the form of a flattened pad. Legs 1-4 are much
reduced in both and are similar in form.

I have grouped all species described in the genus Achtheimus as
one species. These had been separated on the basis of overall form
and not on details of the appendages. It is apparent that we have
here a situation like that found in Perissopus dentatus—a single species
with variation in body form. I have examined Wilson’s material of
A. oblongatus, A. dentatus, and A. pinguis, and I could find no good
basis for keeping them as separate species. On the basis of the
descriptions of other species, there is no valid evidence to justify
more than one species. Probably the best description of the ap-
pendages of this species can be found in Yamaguti’s (1936) description
of A. galeorhint.

This species can be separated from P. dentatus on the basis of 2
characters. In P. dentatus the posterior corners of the genital seg-
ment are sharply angular whereas in P. oblongatus they are rounded.
In P. dentatus the endopods of legs 1-4 are unarmed. In P. ob-
longatus the endopods of legs 1 and 2 and sometimes 3 are armed
with short setae.

I did not collect this copepod and more material would certainly

NO. 3570 PANDARIDAE—CRESSEY 35

be desirable to clarify this situation. It may be that future de-
scriptions will warrant resurrecting some of the previously described
species but, on the basis of existing descriptions, this is not justified.

So far, this species has been reported from a number of inshore
species of sharks, especially of the genera Triakis and Acanthias. It
is a parasite on the body surface of its host.

Perissopus dentatus Steenstrup and Liitken, 1861
Figures 162-189

Perissopus dentatus Steenstrup and Liitken, 1861, p. 393.—Richiardi, 1880,
p. 148.

Perissopus communis Rathbun, 1887, p. 560.

Chlamys incisus Beneden, 1892a, p. 227.

Perissopus crenatus Leigh-Sharpe, 1930, p. 7.

Perissopus manuelensis Gnanamuthu, 195la, p. 1252.

Perissopus travancosiensis Kurian, 1955, p. 108.

Perissopus serratus Heegaard, 1962, p. 175.

SPECIMEN sTuDIED.—Five females and 1 male ex Carcharinus
milbertt from Sarasota, Fla., 1 female ex C. leucas, Sarasota, Fla.,
4 females ex C. maculipinnis, Sarasota, Fla., and 3 females ex
“shark” from Siboga Expedition, 2 females ex Mustelus species
Durban, South Africa.

All descriptions and figures refer to specimens from Carcharinus
milberti unless otherwise stated.

Frmatr.—Body form as in figures 162, 187, 188, and 189. Total
length 4.9 mm (based on 1 specimen). Greatest width 3.8 mm
(measured at widest part of cephalon). Frontal plate not completely
separated from cephalon. First thoracic segment fused with cephalon.
Thoracic segments 2-4 free and with dorsal plates. Plates of segment
2 widely separated and extending only to distal tip of plates of seg-
ment 3. Plates of segment 4 extending only slightly over proximal
portion of genital segment. Genital segment large, comprising
about one-half total body length. Shape of genital segment variable
(see figs. 162, 187, 188, 189). Abdomen (fig. 163) 1-segmented, nearly
hidden in dorsal view. No dorsal plate evident. Caudal rami
joined distally to abdomen, articulated on dorsal surface only. Each
ramus with 7 short spines along posterior border.

Oral area of usual pandarid type. Adhesion pads moderately
developed and associated with first and second antenna and maxilliped.
No pad on outer distal corners of cephalon. First antenna (fig. 164)
2-segmented. First segment with 18 spines, armed as in figure.
Second segment with 11 naked setae. Second antenna (fig. 165) with
small adhesion pad at base, last segment in form of claw with very
small spine near base. Tip of claw (fig. 166) with curious hooded
appearance. Mouth tube of usual type, labium fringed at tip (fig.

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

167). Mandible of usual form. First maxilla (fig. 168) consisting of
broad basal lobe with short spine at tip and group of 3 short setae.
Second maxilla with tip armed as in figure 169. Maxilliped (fig. 170)
large. Terminal spine small but opposed by a very broad adhesion
area. Surface of adhesion area shows imprint of host denticles sug-
gesting a secretion of a cement-like substance on surface of the
maxilliped. The maxilliped can be easily pulled off the copepod
when removing the parasite from the host.

Legs 1-4 (figs. 171-174) biramose. Spine and setal formula as
follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.

seg. 1 E:0 0:0 I:0 0:0 IV 0 IV 0

seg. 2 IV 0 IV 0
Legs 1-4 small and weakly developed. Endopod of all legs unarmed
except for patches of spinules. Segmentation reduced in legs 3 and 4.
Leg 5 (fig. 175) consisting of a single lobe with 4 naked setae, situated
near the ventral distal corner of genital segment (see fig. 163). Leg 6
(fig. 176) consisting of 2 unarmed lobes located near junction of
abdomen and genital segment (see fig. 163).

Egg strings long and straight.

Color in life cream white without pigmentation.

Ma.s.—Body form as in figure 177. Total length 2.9 mm (based
on 1 specimen). Greatest width 1.4 mm (measured at widest point
of cephalon). Cephalon rounded. Frontal plate distinctly separate.
Thoracic segments 2-4 free. No dorsal plates present. Genital seg-
ment longer than wide with posterior corners rounded and scarcely
projecting posteriorly. Spermatophores visible within. Abdomen
2-segmented. Caudal ramus (fig. 178) about as long as wide with 4
long terminal plumose setae and 2 short subterminal ones on distal
corners of each ramus. Four terminal setae all nearly equal in length.

Oral area similar to that of female. Adhesion pads reduced. First
antenna as in female. Second antenna as in figure 179. Oral ap-
pendages as in female. Maxilliped (fig. 180) with a strong terminal
claw opposed by an area of heavy ridges on basal segment.

Legs 1-4 biramose. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4

Dp. end. exp. end. exp. end. exp. end.
seg. 1 I:0 0:0 esi Oral fecal Od ett 0:1
seg. 2 Ii:4 3 TIT6) 8 JULES) WiI:5 4

Leg 1 (fig. 181) with each ramus 2-segmented but exopod only weakly
divided into 2 segments. Legs 2-4 (figs. 182-184) with each ramus
strongly divided into 2 segments. All setae plumose. Leg 5 (fig.

NO. 3570 PANDARIDAE—CRESSEY 37

185) located on midlateral edge of genital segment and consisting of a
group of 3 setae and 1 stout spine. All setae naked, spine finely
barbed. Leg 6 (fig. 186) located near junction of abdomen and genital
segment and consisting of a single naked seta and a finely barbed
stout spine.

Color in life cream white, no pigment.

Remarks.—To date, 7 species of Perissopus have been described.
I have examined material of this genus from several hosts and noted
wide variation in body form. A closer examination of the append-
ages showed no differences between one form and another. Un-
fortunately, I had only a few specimens of each type available. Figure
187 shows an adult female from Carcharinus leucas, Sarasota, Fla.
Its measurements are 4.4 by 2.2 mm. Figure 188 is an adult female
from Carcharinus maculipinns, Sarasota, Fla. (3.9 by 2.4 mm).
Figure 189 is an adult female from the Siboga Expedition to the Indo-
Pacific, host recorded only as a “‘shark.”” It measures 3.8 by 2.2 mm.
A cursory examination of these would indicate more than one species,
but I must, on the basis of the few specimens available, concur with
Capart (1953), who noticed this same variation and concluded that
they should be treated as 1 species.

More material from these and other hosts may favor splitting the
genus into a number of species but, in the face of so small a sample, I
do not feel justified in doing this at the moment. One would expect
to find at least minor differences in the finer details if they are indeed
separate species. None seem to exist. I have thus placed all known
species in synonymy as Perissopus dentatus. Apparently Heegaard
(1962) did not know of the new species of Gnanamuthu or Kurian
when he described Perissopus serratus.

Perissopus dentatus seems to be a parasite of inshore species of
sharks. The copepod is often found externally attached near the
nares and less often near the posterior border of the fins. It is the
only pandarid that attaches to its host by the use of cement. As
pointed out above, the maxilliped of the female bears a very large pad
that, upon removal of the copepod from the host, often shows denticles
or the impression of denticles adhering to the surface of this pad.
This adaption may be due to the rather hazardous area of attachment
on the host (nose). Undoubtedly the nose of the host is often bumped,
thus requiring an effective hold-fast structure if the parasite is to
remain on the host.

The male is described here for the first time. It was attached to a
female when found.

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38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Genus Dinemoura Latreille, 1829

Caligus Miiller, 1785, p. 182. [Refers to C. productus only.]
Dinemoura Latreille, 1829, p. 197. [Type-species: D. producta.]
Binoculus Nordman, 1832, p. 38.
Dinematura Burmeister, 1833, p. 284.
Pandarus.—Baird, 1850, p. 286. [Refers to P. lamnae only.]
Nogagus.— Milne-Edwards, 1840, p. 460. [Refers to N. gracilis only.]
Frema.te.—Frontal plate distinctly separate. Thoracic segments
2-4 free. Dorsal thoracic plates on segment 4. Genital segment
large. Abdomen 2-segmented and joined to genital segment ventrally.
Caudal rami broad. Oral area with or without adhesion pads. Adhe-
sion pad of first antenna double when present. Second maxilla with
a patch of long setules at base of terminal claw. Maxilliped with
small terminal claw. Legs 1-4 biramose. Rami of leg 1 2-sezgmented.
Rami of legs 2 and 3 3-segmented. Rami of leg 4 1-segmented and in
form of broad lamellae. Egg strings long and straight or folded.
Ma.z.—Body form of usual pandarid type. Dorsal plate of seg-
ment 4 reduced. Abdomen 2-segmented. Caudal rami large. Oral
area as in female. Legs 1-4 biramose. Legs as in female with the
following exceptions: terminal segment of endopod of leg 3 modified;
leg 4 rami 2-segmented; legs 5 and 6 present.
Discusston.—The genus Dinemoura occurs on the body surface of
large pelagic sharks. The genus is represented by 4 valid species.
There has been much confusion in the literature over the generic
name of this group. In 1829 Latreille separated Caligus productus
Miiller from the genus Caligus and assigned it to a new genus, Dine-
moura. In 1832 Nordman used the name Binoculus but this was pre-
occupied by Geoffroy in 1792 for a phyllopod genus. Burmeister
changed the name Dinemoura to Dinematura in 1833 because the
etymology of the word Dinemoura was incorrect. Since then, the
2 names have appeared with almost equal frequency. Yamaguti
(1963) suggested that the original name be the proper one. Accord-
ing to Article 33a of the “International Code of Zoological Nomen-
clature,” this viewpoint is correct. Burmeister’s change was an
“unjustified emendation” and cannot stand. Not enough is known
of the males of the 4 species on which to base a key, but one is pro-
vided below for the females.

Key to Females of Genus Dinemoura

1. Adhesion pads conspicuous on oralarea ..... ohekeaho. Mee nis pcunes MLlCt en ere 2
Adhesion pads absent or peas onoral area. . . See ema

2. Genital segment about % body length. Dorsal nleiies bak segment 4 wider
than long... . 5 4 datifolia

Genital segment eon eads ieee een ae a segment 4 longer
Ghar WAG se ae ke) arse en ee Shoe. eroygl Re sie cis pea . . . .producta

NO. 3570 PANDARIDAE—CRESSEY 39

3. Posterior corners of genital segment not greatly produced and rounded. ferox
Posterior corners of genital segment greatly produced and truncated.

discrepans
Dinemoura producta (Miiller, 1785)

Fieures 190, 191

Caligus productus Miiller, 1785, p. 132.

Dinemoura producta.—Latreille, 1829, p. 197.—Krgyer, 1837, p. 202.—Scott, T.,
1901, p. 124.—Scott, T., and Scott, A., 1913, p. 86—Norman and Scott, fl be
1906, p. 211——Norman and Brady, 1910, p. 404.—Fage, 1923, p. 281.—
Pesta, 1934. p. 27.—Oorde and Schuurmans Stekhoven, 1936, p. 139.—
Matthews and Parker, 1950, p. 568.—Barnard, 1955, p. 262.—Delamare-
Deboutteville and Nunes-Ruivo, 1958, p. 223.

Binoculus productus—Nordman, 1832, p. 38.

Dinematura gracilis Burmeister, 1833, p. 284.

Dinematura producta.—Burmeister, 1833, p. 331.—Steenstrup and Liitken, 1861,
p. 370.—Norman, 1868, p. 301.—Bassett-Smith, 1899, p. 463.—Scott, T.,
1900, p. 156.—Brian, 1906, p. 52; 1911, p. 197; 1944, p. 202.— Wilson, 1907,
p. 380; 1923, p. 8; 1932, p. 431.—Hansen, 1923, p. 35.—Scott, A., 1929,
p. 95—Pesta, 1934, p. 27.—Heegaard, 1943b, p. 26; 1945, p. 15.—Delamare-
Deboutteville, 1948, p. 446—Matthews and Parker, 1950, p. 568—Rose and
Vaissiére, 1953, p. 86.— Yamaguti, 1963, p. 117.

Pandarus lamnae Johnston, 1835, p. 203.

Nogagus gracilis—Milne-Edwards, 1840, p. 460.

Dinemoura lamnae Baird, 1850, p. 286.

Nogagus productus.—Gerstaecker, 1853, p. 63.

Dinematura lamnae.—Kr¢yer, 1863, p. 179.

Dinematura affinis Thomsen, 1949, p. 14.—Shiino, 1957, p. 365.

SPECIMENS STUDIED.—Single collections from each of the following
hosts: Isurus oxyrhynchus Rafinesque (35 00’N , 20 00’W), Prionace
glauca (Linnaeus) (39 32’N, 28 02’W), Carcharodon carcharias (Lin-
naeus) off Boothbay Harbor, Maine, Cetorhinus maximus (Gunnerus)
on loan from Zoological Museum in Amsterdam.

FrmaLe—A good description of the female of this species has been
given as recently as 1957 by Shiino. His description is of Dinematura
afinis but, since this is synonymous with Dinemoura producta, the
figures and description apply. I will therefore only point out salient
features.

Body form as in figure 190. Total length (based on an average of
2 specimens) 18.5 mm. Greatest width 8.0 mm. Dorsal thoracic
plates, present only on segment 4 fused basally and covering the
anterior third of the genital segment. Genital segment about % body
length. Caudal rami broad and with setae placed laterally. (Wilson,
1907, showed them as terminal, but in all specimens I examined they
are laterally displaced.) Oral area as in figure 191. First antenna
2-segmented and bearing stout spinose spines on first segment and
short naked setae on terminus of last segment. Second antenna with
a stout claw, sharply bent at tip. Mandible of usual type with 12

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

teeth on tip and projecting within mouth tube as in other members of
group. First maxilla of 2 segments with a palplike process bearing 3
short setae on first segment. Second maxilla with usual fringe claw
at tip, a stout subterminal spine and a subterminal patch of long
hairs. Maxilliped with a short claw at tip.

Legs 1-4 biramose. Rami of leg 1 2-segmented. Rami of legs
2 and 3 3-segmented. Rami of leg 4 1-segmented and modified to
form broad lamellae. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 Be 0:1 Tid 0:1 V 0
seg. 2 IV33) 33 Tet 0:2 I:1 0:2
seg. 3 - - TERS, '6 RE55 4

Leg 5 consisting of a small process bearing 2 short spines, outer
plumose the inner naked. Leg 5 located near junction of abdomen
and genital segment.

Egg strings long and straight.

Color in life cream, generally devoid of pigment.

Matr.—The male of this species was first described by Wilson
(1923). Because I had no material of the male of this genus, J am
unable to expand Wilson’s work. He failed to mention any modi-
fication of the endopod of leg 3. Based on evidence in other members
of the genus, this modification should be present. On the basis
of Wilson’s work the male appendages appear to be very similar to
those of the female, except that the fourth leg of the male is not
modified into broad lamellae as in the female but rather each ramus
is 2-segmented and bears spines and setae.

Remarxs.—This copepod has a very long and confusing history.
It has been described or recorded under 11 different names. Many
of the synonyms have been noted in the literature previously (mainly
Wilson, 1907) but I have, in addition to these, placed D. affinis,
Thomsen, 1949, in synonymy with D. producta. Shiino (1957) de-
scribed and illustrated D. afinis from Japan. These descriptions
agree with material I have collected and identified as D. producta
from the Atlantic from 4 different hosts, including the same host
from which Thomsen described D. affinis. In view of this, I feel
certain that only one species of copepod is involved here.

D. producta is found on several species of pelagic sharks but is
most commonly associated with sharks of the genera Lamna and
Isurus. It is probably worldwide in distribution.

NO. 3570 PANDARIDAE—CRESSEY 4]

Dinemoura ferox (Krgyer, 1838)
Figures 192, 193

Dinematura ferox Kréyer, 1838, p. 40.—Steenstrup and Liitken, 1861, p. 376.—
Olsson, 1868, p. 17.—Meirs, 1880, p. 71.—Bassett-Smith, 1899, p. 463.—
Wilson, 1907, p. 377; 1920, p. 7.—Hansen, 1923, p. 33.—Stephensen, 1940,
p. 5.—Yamaguti, 1963, p. 117.

Dinemoura ferox.—Milne-Edwards, 1840, p. 465.

Dinemoura elongatus Beneden, 1857, p. 226.

Dinematura carcharodonti Thomson, 1889, p. 360.

SPECIMENS STUDIED.—Two collections from the U.S. National
Museum: USNM 12036 from a shark caught off Iceland, USNM
37783 (no collection data).

FrEMALE.—Body form as in figure 192. Total length 32 mm (based
on 1 specimen). Greatest width 10 mm. The female of this species
has been well illustrated by Krgyer (1838) and Steenstrup and Liitken
(1861) so that I will not repeat the figures here. Dorsal thoracic
plates on segment 4. Plates extending slightly over proximal portion
of genital segment. Genital segment large, about one-half body
length. Abdomen 2-segmented. First segment bearing lateral wing-
like projections. Caudal rami large and with 4 short spines along
distal margin. Oral area as in figure 193. No adhesion pads asso-
ciated with first and second antenna. A small pad with maxilliped.
Oral appendages as in D. producta. The claw of the maxilliped is
larger in D. ferox than in D. producta.

Legs 1+4 as in figure 193. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 I:0 0:0 esi 0:1 Teal 0:1 IV IV
seg. 2 IVs a} Tat 0:2 ileal 0:2
seg. 3 HO TTC: ya:

Legs 5 and 6 represented by a series of lobes on posterior ventral
surface of genital segment (see fig. 193). Lobes nearest to attachment
of abdomen probably represent leg 6. Single most posterior lobe
undoubtedly represents leg 5.

Egg strings long and straight.

Color in preserved specimens cream yellow and lacking pigment.

Maue.—Hansen (1932) described a copepod identified by Kroyer
as the male of D. ferox. He did not mention the presence of spermato-
phores which should be the foremost character relating to its sex.
The fourth legs are not modified as in the female. The description
in general is insufficient; that this is actually the male of D. ferox

42 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

is inconclusive. It appears to be of the genus, however. Conse-
quently, the male of this species still remains poorly known and
perhaps even yet to be described.

Discuss1on.—Dinemoura feroz is the largest pandarid copepod yet
described. It is almost twice as long as its next largest relative.
This alone easily separates it from other members of the group. It
has been reported only from sharks from the North Atlantic in the
area of Greenland. The known hosts are Somniosus microcephalus
(Bloch and Schneider), reported by Wilson (1920), and Centrophorus
squamosus Miller and Henle, reported by Hansen (1923). Miers
(1880) reported this copepod taken off the ‘‘Greenland shark” and
noted that the copepods were usually but not always found attached
to the eyes of the host.

Dinemoura latifolia (Steenstrup and Lutken, 1861)
Figures 194-196

Dinematura latifolia Steenstrup and Litken, 1861, p. 378.—Heller, 1868, p. 199.—
Richiardi, 1880, p. 148.—Valle, 1880, p. 60.—Carus, 1884, p. 390.—Brian,
1898b, p. 14; 1899, p. 4; 1902, p. 17; 1906, p. 52; 1944, p. 201.—Bassett-Smith,
1899, p. 463.—Wilson, 1907, p. 383; 1923, p. 15; 1982, p. 432; 1935b, p. 778.—
Yamaguti, 1936, p. 9; 1963, p. 117.—Shiino, 1954, p. 318; 1957, p. 365.—
Delamare-Deboutteville and Nunes-Ruivo, 1954, p. 204.—Barnard, 1955,
p. 263.—Heegaard, 1962, p. 177.

SPECIMEN STUDIED.—Two collections from Jsurus oryrinchus Rafi-
nesque in the North Atlantic (42°18’N, 64°02’W; 35°00’N, 70°00’W).
A single collection from the same host in the Indian Ocean (8°55’S,
55°08’E).

FreMALE.—Body form as in figure 194. Total length 14.5 mm
(based on an average of 5 specimens). Greatest width 8.2 mm.
Dorsal thoracic plates on segment 4 projecting posteriorly over the
anterior portion of genital segment. Genital segment about one-half
body length. Abdomen 2-segmented, each segment with a dorsal
plate. Caudal rami large, each bearing 4 setae. Oral area as in
figure 195.

Since the female of this species has been well described and figured
by Yamaguti (1936) and Shiino (1954), I will only discuss salient
features. Adhesion pads associated with first and second antenna
and maxilliped. Oral appendages similar to those of D. producta.

Legs 1—4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 8 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 ea 0:1 Liat! 0:0 V 0
seg. 2 IMVesir 33 a 0:2 ye. 0:2
seg. 3 Teh: Te 5) 4

Shiino (1947) stated that the fifth legs are still undiscovered. I have

NO. 3570 PANDARIDAE—CRESSEY 43

figured the posteroventral surface of the genital segment (fig. 196).
This region shows 2 areas representing legs 5 and 6. I have interpreted
the single, broad, spatulate process as belonging to leg 5 and the curved,
clawlike process at the junction of the genital segment and abdomen
as leg 6.

Matr.—The male of this species has been described with some
figures by Wilson (1907). Because I had no material of the male of
this species, I am unable to amplify this description. Wilson did show
some modification on the endopod of leg 3 (Wilson, 1907, p. xxv,
fig. 103) which supports my contention that this is characteristic of
the males of the entire group II of the family. The figures and
description are incomplete and more material would certainly be
desirable to complete the male description of this species. He stated
that the fourth legs are not laminate as in the female but each ramus
is 2-jointed and bearing spines and setae. He did not figure this
appendage.

Discussion.—This is a fairly common species occurring on the
body surface of a number of pelagic sharks of the genera Jsurus,
Lamna, and Carcharodon. These 3 genera of sharks compose the
family Jsuridae (after Bigelow and Schroeder, 1948). The parasite
may well be restricted to that group. It is easily separated from
members of the genus by the shape of the dorsal thoracic plates.

Dinemoura discrepans, new species
Figures 197-217

SPECIMENS STUDIED.—Highteen females and 13 males from the body
surface of Alopias vulpinus (Bonneterre) from the Indian Ocean
(9°24’N, 54°58’E). Holotype female (USNM 113592), allotype
male, and 10 paratypes (59 9, 50%”) deposited in alcohol in the
U.S. National Museum, 10 paratypes (59 9, 50’) in the British
Museum (Natural History), and the remaining paratypes in the
author’s collection.

Other specimens studied: a single collection from Alopias vulpinis
in the Indian Ocean (7°17’N, 55°00’E); a single collection from
Alopias superciliosus (Lowe) from Nosy Bé, Madagascar; a single
collection from Alopias superciliosus from the Pacific Ocean (0°38/N,
124°23’W).

Frmate.—Body form as in figure 197. Total length 13.9 mm
(based on an average of 5 specimens). Greatest width 6.7 mm (meas-
ured at widest part of cephalon). Cephalon rounded, 6.7 mm by
6.2 mm, slightly wider than long. Thoracic segments 2-4 free.
Posterior corners of segment 2 produced to form winglike expansions.
Dorsal plates present on segment 4. Plate of segment 4 extending
only slightly over anterior portion of genital segment with their

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

posterior borders serrate. Plates widely separated. Genital segment
longer than wide (5.7 by 3.9 mm). Posterior corners of genital
segment produced and truncated. Abdomen 2-segmented. Each
segment bearing a dorsal plate. Plate of segment 1 bilobed. Plate
of segment 2 single. Caudal ramus large (1.6 by 0.8 mm) and bearing
6 short, naked spines.

Oral area as in figure 198. Adhesion pads reduced. First antenna
(fig. 199) 2-segmented. First segment with 29 short, stout spines
(majority of the spines covered with spinules). Second segment with
14 naked setae. Second antenna (fig. 200) with a terminal claw
recurved at tip, with 2 short spines along outer edge. Small adhesion
areas on the penultimate segment. Mandible (fig. 201) of the usual
type with 11 teeth at tip (fig. 202). First maxilla (see fig. 201) a
broad lobe with a group of 3 setae near base and a broad terminal spine.
Second maxilla (fig. 202) with terminal claw subdivided into 2 seg-
ments (fig. 204). Terminal segment with short rows of fringe.
Penultimate segment with group of broad bladelike setae on inner
distal corner and surface of segment covered with stout setules.
Antepenultimate segment with large spine and patch of hairs at inner
distal corner. Maxilliped (fig. 205) with strong terminal claw.

Legs 1-4 biramose. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 TO) 0:0 Neca 0:1 Lea 0:0 4 0
seg. 2 [V3.3 | | 0:2 TV:6) 10:2
seg. 3 Til: 5 4

Leg 1 (fig. 206) with both rami 2-segmented. No adhesion pads on
the rami. Leg 2 (fig. 207) with each ramus 3-segmented. Outer
edge of first exopod segment striated as an adhesion area. Outer
edges of other exopod segments serrated. Leg 3 (fig. 208) with exopod
of only 2 clearly separated segments. Terminal segment shows some
evidence of being subdivided but indicates a fusion of last 2 segments.
Leg 4 with each ramus in shape of a broad lamella (see fig. 198).
Exopod bearing 4 short spines. Endopod unarmed. Leg 5 (see fig.
209) a single lobe with 1 small setae. Leg 6 (see fig. 219) located near
junction of abdomen and genital segment and composed of 2 spatulate
processes projecting under lateral extensions of first abdominal seg-
ment. ‘Two sclerotized areas located on the midline of genital seg-
ment. ‘These probably function in attachment of spermatophores.

Egg strings long and folded (fig. 210) forming 3 strands.

Color in life cream and without pigmentation.

MaueE.—Body form as in figure 211. Total length 9.7 mm (based
on an average of 2 specimens). Greatest width 5.1 mm (measured
at the widest part of the cephalon). Cephalon about one-half body

NO. 3570 PANDARIDAE—CRESSEY 45

length. Dorsal thoracic plates present on segment 4 and similar to
those of female. Genital segment with posterior corners produced to
form lobes serrate on posterior border (see fig. 212). Genital segment
slightly longer than wide (2.8 by 2.3 mm). Abdomen 2-segmented
and without dorsal plates. Caudal rami as in female.

Oral area as in female. Legs 1-3 as in female except for a modifi-
cation on endopod of leg 3 (fig. 213). This appendage may be used
in the transfer of spermatophores to female. Leg 4 (fig. 214) with
rami 2-segmented and not modified in form of lamellae. Spine and
setal formula of legs 1-4 as follows:

eg 1 leg 2 leg 3 leg 4
exp. end. exp. end, exp. end. exp. end.
seg. 1 0 0:0 gil @eal eel 0:0 LOo0: f
seg. 2 TVS) 3 Tet -Os2, IV26 0:2 Vii Mat
seg. 3 JOULE 6 6 4

Leg 5 (fig. 215) located at the midpoint of the lateral margin and
composed of an outer lobe with 2 setae (outer plumose, inner naked)
and an inner lobe with a single naked setae. Leg 6 (fig. 216) near
Junction of abdomen and genital segment and consisting of a small
process bearing a single naked spine.

Spermatophores (fig. 217) attach to female and cross to opposite
seminal receptacle as in other members of the family. It is less
obvious in this group since the spermatophores lie side by side in
close proximity to each other. Figure 217 shows one spermatophore
removed to reveal neck of the other spermatophore crossing to recep-
tacle of other side.

Color cream white as in female and devoid of pigmentation.

Discussion.—This parasite seems to be specific to sharks of the
genus Alopias. I have examined material from the Indian Ocean
and Pacific and it may well be found throughout the range of the
host genus. This species is easily separated from the known members
of the genus by the shape of the genital segment and nature of the
dorsal thoracic plates. It is found on the body surface of the host
and, from my own experience, seems to favor the undersurface of
the host in the region of the cloaca. The description of the male
should give pertinent information regarding the nature of the lesser
known males of other species of this genus. The modification of the
endopod of leg 3 is quite distinct. It is interesting to note that the
male and female are much more alike than in members of group I
of this family. This may prove to be a consistent difference between
these two groups.

The word discrepans is from Latin, meaning “‘to differ.’

46 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Genus Demoleus Heller, 1865

Caligus.—Otto, 1821, p. 15. [Refers to C. heptatus only.]
Binoculus—Nordman, 1832, p. 32. [Refers to B. sexsetaceus only.]
Dinematura.—Burmeister, 1833, p. 331. [Refers to D. sexsetaceus only.]
Nogagus.— Gerstaecker, 1853, p. 63. [Refers to N. productus only.]
Demoleus.—Heller, 1868, p. 199. [Type-species: D. paradozus.]

Frmatz.—Frontal plate distinctly separate. Thoracic segments
2-4 free. Dorsal thoracic plates on segment 4 only. Abdomen 1-
segmented and with a dorsal plate. Caudal rami large. Oral area
with adhesion pads associated with first and second antenna, maxilliped
and an additional pad between bases of the maxillipeds. Oral
appendages of usual type. The second antenna with terminal hook-
like spine reduced. Penultimate segment of second maxilla with
large spine and patch of stout setules on distal corner. Legs 1-4
biramose; all rami 2-segmented. Legs 5 and 6 present. Egg strings
folded.

Maux.—Body form of usual type. No dorsal plates present.
Oral area similar to that of female. Appendages similar to female
except for a modification on endopod of third leg of male. Males
of this genus are poorly known.

Discussron.—At present there are 2 species in this genus. D.
heptatus, female, is well known and is amply recorded in the literature
principally as an external parasite of Hexanchus. LD. latus is reported
by Shiino (1954) from Acanthidium eglantina (Jordan and Snyder)
and I have examined material from Squalus acutipinnis Regan. Both
of these hosts are members of the family Squalidae.

The adhesion process between bases of maxillipeds is not found in
any other genus in this family. This character alone sets it apart
from other pandarid copepods.

Key to Females of Genus Demoleus

Genital segment covering the dorsal abdominal plate. Egg strings exposed.

heptatus
Genital segment only covering about of the dorsal abdominal plate. Egg
ptrimgs hidden iAte et Gh te. \a, RPL, Re. A RARE SRG Salton a Aha Monin latus

Demoleus heptatus (Otto, 1821)
Figures 218-237

Caligus heptatus—Otto, 1821, p. 15.

Caligus paradozus Otto, 1828, p. 352.

Binoculus sexsetaceus Nordman, 1832, p. 32.
Dinematura sexsetaceus—Burmeister, 1833, p. 331.
Nogagus productus Gerstaecker, 1853, p. 64.

NO. 3570 PANDARIDAE—-CRESSEY 47

Demoleus paradoxus.—Heller, 1868, p. 199.—Carus, 1884, p. 361.—Bassett-
Smith, 1899, p. 460.—Pearson, 1905, p. 166.—Brian, 1906, p. 50.—Wilson,
1907, p. 349; 1935b, p. 778.—Scott, T., and Scott, A., 1913, p. 79.—Rose
and Vaissiere, 1953, p. 85.

Demoleus heptatus—Dollfus, 1943, p. 1—Yamaguti, 1963, p. 115.

SPECIMENS STuDIED.—Eleven females and 1 male USNM 60465
from Hezanchus species from Monterey Bay, California.

FrmaLe.—Body form as in figure 218. Total length 12.5 mm
(based on a single specimen). Greatest width 4.5 mm (measured
at widest part of cephalon). Carapace rounded about as long as
wide. Dorsal thoracic plates present on segment 4. Plates extend-
ing only slightly over anterior portion of genital segment. Genital
segment 5.8 mm long and 3.1 mm wide, about one-half body length.
Posterior corners produced dorsally and extending as rounded lobes
over abdomen and caudal rami. Abdomen (see fig. 219) 1-segmented
and with a large dorsal plate. Caudal rami large (2.6 by 1.3 mm),
joined to the abdomen distally (see fig. 219). Each ramus with 6
naked setae, innermost and outermost very short.

Oral area (fig. 220) with adhesion pads associated with first and
second antenna and maxilliped. Adhesion pads small and not well
developed. A padlike process located between bases of maxillipeds
(fig. 221). This process is directed posteriorly and has a pad at tip.
Pad is divided by a median line suggesting a fusion of 2 pads. First
antenna (fig. 222) 2-segmented. First segment with 28 spines and
setae armed as in figure. Last segment with 9 naked setae. Second
antenna (fig. 223) small. Terminal claw not well developed. Ad-
hesion area near base. Mouth tube of the usual form. Mandible
with 12 teeth at tip. First maxilla (fig. 224) composed of a lobe with
articulated process at tip and basal process bearing 3 short spines.
Second maxilla of usual form. Terminal claw (fig. 225) subdivided.
Distal segment with rows of fringes asin figure. Subterminal segment
with patch of spines. Penultimate segment bearing prominent
spinose spine and group of long hairs. Maxilliped (fig. 226) with
terminal claw opposed by sclerotized protuberances on basal segment.

Legs 1-4 (figs. 227-230) biramose. Spine and setal formula as
follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 LO 0:0 io! 0:1 eat 0:1 ligil, Meu
seg. 2 LV:3" "3 TVi25.. «8 PV 25° 6 We S58

Leg 5 (fig. 231) located on ventral surface of genital segment near
lateral margin (see fig. 219) and composed of single lobe with 2 naked
spines. Leg 6 (fig. 232) at area of spermatophore attachment and
modified to assist in this function.

4S PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Egg strings folded to form 3 strands.

Ma.e.—Body form as in figure 233. Total length 9 mm (based on
1 specimen). Greatest width 3.4 mm. No dorsal plates present.
Oral area as in female. First antenna as in female. Second antenna
(fig. 234) with clawlike tip. Penultimate segment with stout hooklike
spine at midpoint. Other oral appendages as in female. Maxilliped
(fig. 235) with long terminal claw opposed by 4 adhesive areas on
preceding segments. Legs 1-4 as in female except for modification
of last segment of endopod of leg 3 (fig. 236). Leg 5 single lobe
with 2 or 3 setae. In the only specimen I was able to observe one
leg was broken and the other was not clear. Leg 5 located along
margin of genital segment near midpoint. Leg 6 (fig. 237) located
near junction of abdomen and genital segment and composed of
a single lobe with 2 naked setae.

Discussion.—Demoleus heptatus has been described and its synon-
ymy considered recently by Dollfus (1943). The male is poorly
known and has been elaborated here on the basis of a single
specimen. This parasite seems to be most common on sharks of the
genus Hexanchus.

Wilson designates Nogagus grandis Steenstrup and Liitken as the
male. This has been shown to be the male of Phyllothyreus cornutus
Milne-Edwards. The true male is not well known and the single
specimen that I studied may be the first record of the true male of
this species.

Demoleus latus Shiino, 1954
FIGURES 238-242
Demoleus latus Shiino, 1954, p. 325.

SPECIMENS sTuDIED.—A single collection of 12 females from the
Discovery Collections, collected off Cape Trawler, July 8, 1927,
from Squalus acutipinnis.

FrmaLtE.—This species has been well described and figured by
Shiino (1954). Except for the inclusion of a few details omitted in
the original description, I will only present a superficial overall
descripton here.

Body form as in figure 238. Total length 9.2 mm. (based on an
average of 3 specimens). Greatest width 3.1 mm. (measured at
widest part of genital segment). Dorsal thoracic plates present on
segment 4. Genital segment about one-half body length. Abdomen
l-segmented and with broad dorsal plate. Caudal ramus broad,
with 4 terminal naked setae.

Oral area (fig. 239) and associated appendages well described by
Shiino except that it is worth noting that there is an adhesive pad
between the bases of the maxillipeds as in D. heptatus.

NO. 3570 PANDARIDAE—CRESSEY 49

Legs 1—4 biramose, with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 O20). yet Or Tet On Tee 0:1
seg. 2 Wess LV eS IV:5 5 Pb) 4

Leg 5 (fig. 240) located ventrally near outer distal corner of genital
segment and composed of a signle lobe bearing a short plumose seta.
Leg 6 (fig. 241) located near junction of abdomen and genital segment
and represented by a finger-like lobe.

Egg strings coiled and not protruding (fig. 242), remaining hidden
between dorsal abdomen plate and abdomen.

Color in preserved specimens cream and devoid of pigmentation.

Matze.— Unknown.

Discusston.—I have been able to supplement the original descrip-
tion on a few points, particularly the nature of the 5th and 6th legs
and the egg strings. The specimens I received were still attached
to pieces of fin from the host shark. I noticed that the copepods
were attached to the lighter pigmented surface (presumably lower)
of the fin. This would indicate that in this instance the pectoral or
pelvic fins were involved.

This is only the second record of this copepod; consequently, too
little is known about the species to draw any conclusions regarding
its host specificity or geographic distribution.

This species is easily separated from D. heptatus by the nature of
the abdomen and general configuration of the body.

Genus Pagina Cressey, 1964
Pagina Cressey, 1964, p. 285. [Type-species: P. tunica.]

Frmaute.—Frontal plate distinctly separate. First thoracic seg-
ment fused with cephalon. Thoracic segments 2-4 free. Second
and third thoracic segments without dorsal plates. Fourth segment
with a dorsal plate. Abdomen 2-segmented, each segment with a
dorsal plate. Abdomen attached to distal end of genital segment
and visable dorsally. Cephalic appendages of typical pandarid
type. Legs 1-4 biramose; rami of leg 1 2-segmented, those of legs
2-4 3-segmented; all bearing plumose setae. Fifth and sixth legs
present. Egg sacs long.

Maur.—Body of typical pandarid form. No dorsal plates present.
Appendages with same generic characters as the female.

The name Pagina, from Latin, meaning “a page,” refers to the
relationship of the abdominal plates to each other.

221-534—67—_+4

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Pagina tunica Cressey, 1964
Figures 243-267
Pagina tunica Cressey, 1964, p. 285.

SPECIMENS STUDIED.—21 specimens (18 females and 3 males) col-
lected from Alopias superciliosus (Lowe) caught at Majunga, Mada-
gascar. Holotype female, allotype male, and 8 paratype females in
alcohol deposited in the U.S. National Museum, 3 paratype females
in alcohol deposited in the Collection of the Centre d’Oceanographic
et des Peches de Nosy Bé, Madagascar, and the remaining paratypes
in the author’s collection.

OTHER SPECIMENS STUDIED.—A single collection from Alopias
superciliosus from Nosy Bé, Madagascar; a single collection from the
same host in the Pacific (0°38’N, 124°23’W).

FrMALe.—Body form as in figures 243 and 244. Total length,
based on an average of 4 specimens, including caudal rami but not
setae, 17.9 mm. Greatest width (measured at the widest part of
cephalon) 5.8 mm.

Cephalon nearly round, slightly longer than wide, measuring 6.0 by
5.8 mm (measurements including marginal fringe). Posterior corners
of cephalon projecting distally. First thoracic segment fused with
head. Second segment distinct with 2 lateral lobes extending to
posterior margin of third thoracic segment, thus incorporating the
smaller third segment within its posterior border. Fourth thoracic
segment with dorsal plate consisting of 2 conspicuous winglike lobes.
The distal corner of this plate extends only slightly over anterior
corner of genital segment. Genital segment large, its greatest length
4.6 mm and its greatest width at the posterior corners 3.8 mm. Pos-
terior corners produced to form 2 short lobes. Abdomen 2-segmented,
both segments possessing a conspicuous dorsal plate. From the dorsal
aspect abdominal plates covering rest of abdomen. Ventrally the
first abdominal segment (fig. 248) as wide as long, measuring 1.5 by
1.5mm. Dorsal plate of this segment extending over rest of abdomen
and with a deep median sinus dividing it into 2 long lobes (see figs.
243, 244). Second abdominal segment also as long as wide and
measuring ventrally 2.1 by 2.1 mm. Its dorsal plate composed of
only a single lobe and extending over proximal ends of caudal rami
(see figs. 243, 245). Caudal ramus (fig. 245) large, comprising almost

one-fourth of total body length. Each ramus about 4 times as long |
as wide measuring 4.2 by 1.3 mm. Distal end of ramus bearing 6 ©

setae. The outermost and innermost small and naked, but median 4
plumose and all nearly equal in length, longest measuring 0.4 mm.
Outer border heavily sclerotized while inner only weakly so and often
appearing wrinkled in preserved specimens.

—_—_—-—

NO. 3570 PANDARIDAE—CRESSEY 51

Oral area as in figure 246. A single pair of adhesion pads near base
of first antenna. First antenna (fig. 247) 2-segmented. First segment
0.77 mm long and bearing 23 stout setae along anterior distal border,
all of which are covered with spinules. Four smaller finely plumose
setae internal to outer spines. Terminal segment 0.42 mm long and
bearing 13 naked setae. Second antenna (fig. 249) 3-segmented.
Terminal segment in form of a stout, heavily sclerotized claw bearing
2 setae, 1 basal and the other median. No adhesion pad associated with
this appendage. Mouth tube (fig. 250) about twice as long as basal
width with labium extending beyond tip of labrum. Tip of labrum
somewhat expanded and weakly trilobed with a pair of subterminal
processes projecting within tube (fig. 251). The labium expanded at
tip and fringed as in figure 251. The mandible attached to head near
base of tube (see fig. 250) and bearing a long process with extends be-
tween labrum and labium. Distal end of mandibular process with an
inner row of about 11 teeth (fig. 252). First maxilla (fig. 253) indis-
tinctly divided into 2 segments. Proximal segment with a group of 3
setae near base. Distal segment short and terminating as a blunt
process. Second maxilla (fig. 254) 3-segmented. Second segment
bearing a group of stout setules and a single seta at inner distal corner.
Third segment short, with 2 transverse rows of setules and bearing
a claw ornamented on proximal two-thirds of concave margin with
transverse rows of spinules and on convex surface with longitudinal
rows of spines. Maxilliped (fig. 255) apparently 4-segmented. First
segment bearing padlike process on anteroventral surface, which
undoubtedly serves as an adhesion pad. Third segment bearing a
heavily sclerotized spinelike process opposed by claw of fourth seg-
ment, thus forming a chela. Fourth segment bearing a single seta
near base of claw.

Legs 1-4 biramose, with spine and setal formula as follows:

leg 1 eg 2 leg 8 leg 4
xp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 T= 0:1 Isat 0:1 eat 0:1
seg. 2 LE: 43 Iai 0:2 Tea 0:2 lice 0:2
seg. 3 - - TIL25,..6 TETs5), 4 IDES, VS

Leg 1 (fig. 256) with both rami 2-segmented. First exopod segment
outwardly greatly inflated and bearing a single outer spine. Last
exopod segment with 3 outer spines and 4 inner setae. First endopod
segment with no setae but second segment bearing 3 setae. Basipodite
bearing a short outer seta; inner margin naked except for a short seta.
No setae on coxopodite. Leg 2 (fig. 257) with both rami 3-segmented.
First exopod segment with an outer spine and an inner seta. Second
segment same. Third segment with 3 outer spines and 5 terminal
setae. First endopod segment with an inner seta. Second segment

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

with 2 inner setae. Third segment with 6 terminal setae. Basipodite
with only an outer seta; inner margin with a row of hairs. Coxopodite
seta stout and densely plumose. Leg 3 (fig. 258) with both rami 3-
segmented. First and second exopod segments with an outer spine
and an inner seta. Third segment with 3 outer spines and 5 terminal
setae. First endopod segment with an inner seta. Second segment
with 2 inner setae. Third segment with 3 terminal setae. Inner
portion of basipodite expanded with a marginal fringe as in figure.
No seta on the coxopodite. All setae on legs 1-4 plumose and all
spines fringed. Leg 5 located on ventral surface near posterior corner
of genital segment (see fig. 248), bearing 3 naked spines. Leg 6 modi-
fied to form a hooklike process that holds spermatophores in place
(see fig. 248). Egg strings long, 1.5 times as long as body.

Maure.—Body form as in figure 260. Total length, based on an
average of 2 specimens, including caudal ramus but not setae 11.7 mm.
Greatest width 4.6 mm measured at widest part of cephalon. Cepha-
lon nearly round somewhat wider than long (4.6 by 4.2 mm) with
posterior corners projecting. Lateral dorsal edge of segment 2 bearing
on each side a clear membrane that extends posteriorly to fourth seg-
ment. Fourth segment with only a suggestion of winglike plate
found in female. Genital segment (fig. 261) 2.6 by 2.1 mm, somewhat
longer than wide. Spermatophores visable through posterior half of
genital segment. Abdomen 2-segmented, without dorsal plates.
First segment measuring 0.88 by 0.88 mm. Second segment longer
than wide (1.4 by 1.08 mm) with widest part in distal portion of seg-
ment. Caudal ramus armed as in female; about 4 times as long as
wide (1.9 by 0.49 mm). Inner margin bearing a row of short hairs.

Oral area as in female. First antenna like that of female. Second
antenna (fig. 262) 4-segmented. Second segment with striated areas
as shown in figure. Claw shorter and stouter than in female and com-
posed of 2 segments (in the female these segments are fused to form
1). Other oral appendages like those of female. Maxilliped (figs.
263, 264) heavily sclerotized with achela at tip. When chela is closed,
claw of last segment fits into a bifurcation on tip of spinelike process on
penultimate segment. A bossed area present between these claws.
In addition to claw, last segment bearing a single seta (see fig. 264).
Adhesion process near base of maxilliped as in female.

Legs 1—4 as in female except for last endopod segment of leg 3 (fig.
265). Ventral surface of this segment bearing a heavily sclerotized
process that extends out over an embossed area near edge (this seems
to be modified for holding, but exact function is yet unknown). In
addition to ventral process, a more weakly sclerotized dorsolateral
process. Leg 5 (fig. 266) located ventrolaterally in middle of genital
segment (see fig. 261) and bearing 4 setae, 3 plumose and 1 fringed with

NO. 3570 PANDARIDAE—CRESSEY 53

spinules. Outer seta not greatly displaced from other 3, as in female.
Leg 6 (fig. 267) located internal to distal corner of genital segment and
consisting of a small process bearing short setae.

Discussion.—The genus Pagina is closely related to Dinemoura
Latreille, 1829, of which there are 4 known species. These 2 genera
have the following characteristics of the female in common: a wing-
like dorsal plate on the fourth thoracic segment; a 2-segmented abdo-
men, each abdominal segment bearing a dorsal plate, the first of which
is bilobed and the second single-lobed, and legs 1-3 similar and rela-
tively unmodified. Pagina can be separated from Dinemoura by the
fact that in Dinemoura the fourth leg is broad and conspicuously
lamelliform, whereas the fourth leg of Pagina is unmodified.

Pagina is unlike all other known genera of this family in having
the rami of legs 2-4 3-segmented.

Genus Echthrogaleus Steenstrup and Liitken, 1861

Dinemoura.— Guerin-Meneville, 1837, pl. 35. [Refers to D. alata only.]

Dinematura.—Dana, 1852, p. 60. [Refers to D. braccata only.]

Echthrogaleus Steenstrup and Liitken, 1861, p. 380. (Type-species: EH. coleop-
tratus.]

Pandarus—Thomson, 1889, p. 363. [Refers to P. armatus only.]

Frmatx.—Frontal plate distinctly separate. First thoracic segment
fused with cephalon. Dorsal thoracic plates on segment 4. Abdomen
1-segmented. Abdomen concealed beneath genital segment. Caudal
rami joined to abdomen terminally. Oral adhesion pads present but
somewhat reduced. First antenna 2-segmented. Legs 1-4 biramose.
Leg 4 lamelliform. Legs 5 and 6 present. Egg strings long and
straight.

Matz.—No dorsal thoracic plates present. Abdomen 2-segmented
Legs 1-4 biramose. Leg 3 with modification on endopod. Leg 4
not lamelliform. Legs 5 and 6 present and not as reduced as in female.
Other oral and thoracic appendages as in the female.

Discussion.—This genus is cosmopolitan as a parasite on the body
surface of elasmobranch fishes. LZ. coleoptratus and denticulatus may
be restricted to larger pelagic sharks whereas torpedinis has been re-
ported only from the ray, Torpedo occidentalis. The copepod is para-
sitic on the body surface of the host.

This genus is closely related to Dinemoura but differs from it princi-
pally in having a 1-segmented abdomen. The males of this genus
have not been well described. The descriptions of coleoptratus and
denticulatus males have been amplified here.

In 1899 Thomson described a copepod under the name of Dinematura
hamiltoni. This is obviously a member of the genus Echthrogaleus,
but the figures and description are too poor to assign it to any species.

54 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Key to Females of Genus Echthrogaleus

1. Posterior border of dorsal thoracic plate smooth. Leg 5 concealed in dorsal
ViCW fs ers ..... . coleoptratus
Posterior border of dorsal ihoracie plate nee a EN eR Sieh ced ces

2 Leg 5 visible in dorsal view. Dorsal thoracic plate covering not more than
Ff PENAL SOLMENG ao case. bs inst bs Hey eee dot ead Sets tes | Webbe te denticulatus

ree 5 concealed in dorsal view. Daral thoracic plate covering at least %
Conital BeGMEDG oo. whe ae ws vey ene, act ceric. egies am eee ner ea OEE OATES

Echthrogaleus coleoptratus (Guerin-Meneville, 1837)
Figures 264-280

Dinemoura alata Guerin-Meneville, 1837, p. 42.

Dinemoura coleoptrata Guerin-Meneville, 1837, p. 42.

Dinemoura affinis Milne-Edwards, 1840, p. 465.

Dinematura braccata Dana, 1852, p. 60.

Echthrogaleus coleoptratus Steenstrup and Liitken, 1861, p. 380.—Olsson, 1868,
p. 20.—Norman, 1868, p. 301.—Rathbun, 1884, p. 488.—Brian, 1899, p. 4;
1902, p. 8; 1906, p. 53; 1908, p. 4; 1912, p. 12; 1914b, p. 148; 1944, p. 202.—
Bassett-Smith, 1899, p. 464.—Scott, T., 1900, p. 156; 1901, p. 125; 1902,
p. 292.—Scott, T., and Scott, A., 1913, p. 89.—Norman and Scott, T., 1906,
p. 214.— Wilson, 1907, p. 367; 1908, p. 452; 1920, p. 12; 1922, p. 5; 1923,
p. 138; 1932, p. 427.—Norman and Brady, 1910, p. 404.—Stebbing, 1910,
p. 559.—Hansen, 1923, p. 33—Marukawa, 1925, p. 1242; 1947, p. 926.—
Yamaguti, 1936, p. 7; 1963, p. 119.—Oorde and Schuurmans Stekhoven, 1936,
p. 1389.—Rose and Vaissiere, 1953, p. 86.—Shiino, 1954, p. 291; 1957, p. 364.—
Delamare-Debouteville and Nunes-Ruivo, 1954, p. 204.—Barnard, 1955, p.
264.—Capart, 1959, p. 97—Stuardo and Fagetti, 1961, p. 58—Heegaard,
1962, p. 177.

Echthrogaleus braccatus—Heller, 1868, p. 197.—Thomson, 1889, p. 361.—Norman
and Scott, 1906, p. 213.—Wilson, 1907, p. 366.

Nogagus lutkenit Norman, 1868, p. 300.

Echthrogaleus perspicax Olsson, 1868, p. 18.—Wilson, 1907, p. 457.

Echthrogaleus lutkenti—Norman and Scott, 1906, p. 213.—Scott, T., and Scott,
A., 1913, p. 90—Oorde and Schuurmans Stekhoven, 1936, p. 139.

Echthrogaleus affinis—Brady, 1883, p. 133.—Bassett-Smith, 1899, p. 465.—
Wilson, 1907, p. 363.

SPECIMENS STUDIED.—Twenty-six collections from Prionace glauca
from the western North Atlantic (35°-45°N, 20°-80°W). A single
collection from Lamna ditropis Hubbs and Fallett sent to me by
Dr. P. Gilbert from the North Pacific. Four collections from Prionace
glauca from the Indian Ocean (42°23’S, 74°56’E) and 2 collections
from Lamna nasus (Bonnaterre) at the same locality.

This copepod has been extensively collected and reported in the
literature. The female has been well described and figured (most
recently by Shiino, 1954); however, the male is poorly known and
will be dealt with here in more detail.

FrmaLr.—Body form as in figure 268. Total length 9.7 mm (based
on an average of 10 specimens). Greatest width 4.8 mm. Dorsal

NO. 3570 PANDARIDAE—CRESSEY 55

thoracic plates on segment 4. Oral area and associated appendages
described by Shiino (1954). First maxilla as in figure 269. Adhesion
pads present but small.

Leg 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 EON Os0 Ea Ora Tere 0:1 LV
seg. 2 IV=3) 3 1:0 8 Te 6
seg. 3 TEES TEES

Leg 1 endopod (fig. 270) with usual 3 setae but inner and outer
short. Sternal plate between legs 1 with a bilobed adhesion pad
(fig. 271). Leg 5 (fig. 272) a single lobe with 3 stout spines near tip.
Leg 6 incorporated into area of spermatophore attachment and not a
separate element.

Egg strings long and straight.

Color in life cream with light brown pigment on cephalon.

Mate.—Body form as in figure 273. Total length 6.4 mm (based
on an average of 2 specimens). Greatest width 3.6 mm (measured
at widest part of the cephalon). Cephalon rounded, about as long
as wide. No dorsal plates present. Genital segment (fig. 274)
1.6 by 1.4 mm, slightly longer than wide. Posterior corners not

markedly produced. Abdomen 2-segmented. Caudal ramus with
4 moderately long, plumose setae and plumose along inner margin.

Oral area and associated appendages as in the female. Adhesion

| pads somewhat smaller.

Legs 1-4 biramose. Spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 i000 ifeil eal Isiah @sal 1g ays
seg. 2 IV:3 3 Teen 8 Eater ce er eo
seg. 3 III:5 TSS TSS

_ Leg 1 (fig. 275) as in the female except that setae on endopod are
of equal length in male. Leg 2 (fig. 276) as in female but without
_ patches of stout spinules. Leg 3 with a modification of last endopod
segment (fig. 277). Leg 4 as in figure 278.

Leg 5 (fig. 279) located on midlateral margin of genital segment

_ and consisting of a single lobe with 1 stout spine and 3 plumose setae.

Leg 6 (fig. 280) situated near junction of genital segment and abdomen

_ and represented by a single spine and plumose seta.

Color in life cream, devoid of pigment.
Discussion.—Echthrogaleus coleoptratus is a widely distributed

_ species of copepod and has been reported from a wide variety of

ee

sharks. It is generally found on pelagic rather than inshore varieties.

it occurs on the body surface of the host, commonly on the fins.

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

The male shows the modified endopod of leg 3 placing this copepod
in group II.
Echthrogaleus denticulatus Smith, 1874
Figures 281-290

Echthrogaleus denticulatus Smith, 1874, p. 282.—Rathbun, 1884, p. 488.— Wilson,
1907, p. 369; 1932, p. 428.—Shiino, 1954, p. 297; 1959b, p. 352.

Dinematura neozealanica Thomson, 1889, p. 359.—Bassett-Smith, 1899, p. 464.—
Wilson, 1907, p. 363.

Pandarus armatus Thomson, 1889, p. 363. [Nogagus male.]

SPECIMENS sTUDIED.—A single large collection from Alopias
pelagicus Nakamura from Majunga, Madagascar. Three collections
from Alopias vulpinus from the Indian Ocean (16°13’N, 63°29’E;
9°24’N, 54°58’E; 7°17’N, 55°00’E). A _ single collection from
Eulamia floridanus from the Indian Ocean (14°36’N, 55°23’E),
Two collections from Alopias vulpinus from the Pacific Ocean (7°47’N,
102°377 WW; 212154N, TA3°267VV):-

Frma.LE.—Body form as in figure 281. Total length 7.8 mm
(based on an average of 5 specimens). Greatest width 4.3 mm
(measured at the widest part of cephalon). The female of this
species has been recently redescribed and figured by Shiino (1954)
and a complete description will not be repeated here. Oral area of
usual type, adhesion pads present. Cephalic appendages as shown
by Shiino except terminal claw of second maxilla (fig. 282) is indis-
tinctly subdivided.

Legs 1-4 biramose with spine and setal formula as follows:

leg 2 leg 8 leg 4
Dp end. exp. end. exp. end. exp. end.
seg. 1 1:0 OO. e 0s Wie Braet ab 0:1 Xe, Vi
seg. 2 IV:3 3 T:1 O52). 1st 6
seg. 3 IV:4 6 Tih:5

Leg 5 (fig. 283) a long process projecting beyond distal corners of
genital segment, bearing a stout terminal spine and 2 subterminal
setae. Leg 6 (fig. 284) located at junction of abdomen and genital
segment and composed of a bilobed process. Leg 6 covered by
abdomen in ventral view.

Egg strings long and straight.

Color in life cream, no pigmentation.

Maur.—Body as in figure 285. Total length 5.2 mm (based on
an average of 2 specimens). Greatest width 2.8 mm (measured at
widest point of cephalon). Frontal plate separate. Cephalon
rounded, somewhat wider than long. No dorsal thoracic plates.
Genital segment (fig. 286) with posterior corners produced only
slightly. Two patches of spinules on ventral surface of segment.
Genital segment longer than wide (1.2 by 1.0 mm). Abdomen 2-

|
|

—

NO. 3570 PANDARIDAE—CRESSEY 57

segmented. First segment 218u long. Second segment 360y long.
Caudal rami long, measuring 720u by 144y (about 5 times as long
as wide). Each ramus bearing 4 plumose terminal setae and 2
shorter subterminal ones. Each ramus plumose along inner edge.
Oral area as in female. Setae on first and second antennae slightly
longer than in female.
Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end, exp. end.
seg. 1 I:0 0:0 Tea 0:1 faa O:1 tea Oz
seg. 2 LY 23h) 23 eal 8 Te REG wrt 5
seg 3 IIL: 5 TITs5 III:4

Leg 1 as in female. Leg 2 endopod of only 2 segments. Leg 3 with
modification as in figure 287. Leg 4 (fig. 288) bearing a prominent
spine on outer distal corner of first exopod segment. Leg 5 (fig. 289)
located on posterior corners of genital segment, composed of a process
bearing a stout terminal spine and 3 setae on inner margin. Distal 2
setae plumose. Proximal one naked. Leg 6 (fig. 290) located at
junction of abdomen and genital segment, composed of an inner stout
spine and an outer plumose seta.

Color in life cream and devoid of pigment.

Discussion.—This copepod was recently redescribed by Shiino
(1954). He described a male of this species also, but a closer exami-
nation shows this to be actually the male of Pandarus satyrus. A
comparison of his description and figures leaves no doubt as to its
true identity. He comments that there is ‘‘a remarkable dimorphism
found between the sexes.”” This is not the case when one considers
the true male of this species, described here for the first time. The
modification on the third leg is well formed, establishing the position
of this species in group II. I have synonymized D. neozealanica
Thomson, 1889, with this species since Thomson’s original description
fits in every way.

This copepod is found generally on the body surface of a wide
variety of pelagic sharks. It is occasionally recovered from the gills
also. I collected large numbers of this species from Alopias vulpinus
in the Indian Ocean. On this host the copepod often occurred in
great clusters (200-300 copepods) around the opening of the cloaca.
Females far outnumber the males in such cases. This copepod has
been reported from the Atlantic (Smith, 1874), North Pacific (Shiino,
1954), and I have collected it from the Indian Ocean and received
material from the Central Pacific. This seems to indicate a cosmo-
politan distribution. It is apparently a common parasite of the genus
Alopias.

58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Echthrogaleus torpedinis Wilson, 1907
Fiaures 291-294
Echthrogaleus torpedinis Wilson, 1907, p. 371; 1932, p. 429.

SPECIMENS STUDIED.—USNM 11350, syntypes. 3 females from
Torpedo occidentalis from Provincetown, Mass.

The only material available for study were type specimens from the
U.S. National Museum. Because I did not dissect any of these types,
a complete redescription is not possible here. This will have to wait
until more material can be collected; nevertheless, I have figured some
details and have added here to the original description.

Frema.Le.—Body form as in figure 291. Total length 12.8 mm (based
on a single specimen). Greatest width 8.2 mm (measured at widest
part of the dorsal thoracic plates). Cephalon rounded, about as wide
as long (6.2 by 6.2mm). Dorsal thoracic plates on segment 4. These
plates very conspicuous and serrated along their posterior borders.
Plates extending over the proximal two-thirds of the genital segment.
Genital segment with its posterior corners produced to form inwardly
directed lobes. Abdomen 1-segmented, hidden in dorsal view. A
small dorsal plate with abdomen. Caudal ramus (fig. 292) bearing 6
naked setae on posterior border. Rami with fine spinules along mner
margins.

Oral area with adhesion pads reduced. Pad associated with maxil-
liped in form of a posteriorly directed process. Oral appendages like
those of H. denticulatus.

Legs 1-4 (fig. 293) biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 Teal O:1 est O:1 EX, TV
seg. 2 Wie S ees ell of est 6
seg. 3 IV:4 IV:4

Leg 4 lamelliform as in figure. Leg 5 (fig. 294) a process not projecting
beyond tip of genital segment, bearing a single stout spine and 3 setae
armed as in figure. Leg 6 as in F. denticalatus.

Color in preserved specimens cream tan, devoid of heavy pigmen-
tation.

Egg strings long and straight.

Mate.— Unknown.

Discussion.—This species seems closely related to EL. denticulatus,
but it can be separated from that species by the nature of the dorsal
thoracic plates of segment 4, by the relative length of leg 5, and by
the differences in the armature of the legs. This copepod has been
collected twice from Torpedo occidentalis off the coast of Massachu-
setts. Further collecting would be necessary before concluding that

NO. 3570 PANDARIDAE—CRESSEY 59

it is specific to that host. It was reported by Wilson (1907) from the
pectoral and ventral fins of the host.

Genus Nesippus Heller, 1868

Nesippus Heller, 1868, p. 193. [Type-species: N. orientalis.]

Nogagus.—Beneden, 1892b, p. 246. [Refers to N. augustatus only.]

Frema.te.—Frontal plate distinctly separate. First thoracic seg-
ment fused with cephalon. Thoracic segments 2-4 free. Dorsal
plate may or may not be present on segment 4. Abdomen 1-seg-
mented and joined to genital segment ventrally. Caudal rami at-
tached distally to abdomen. Adhesion pads present on cephalon.
First antenna 2-segmented. Oral appendages of usual pandarid type.
Maxilliped with a claw pointed or rounded at tip. Legs 1-4 biramose.
Rami of legs 1-3 2-segmented, those of leg 4 l-segmented. Leg 5
reduced to 1 or 2 setae. Leg 6 absent. Egg strings straight.

Maur.—The same generic characters of female with following
exceptions. A reduced modification on endopod of leg 3. Leg 6
present but much reduced. Abdomen 1|-segmented. The males of
this genus can be separated from all other pandarids by the 1-seg-
mented abdomen.

Discussion.—Members of this genus seem to be restricted to inshore
species of sharks and are generally not found on the body surface of
the host. The usual sites of infestation are the mouth, gill arches,
and nasal passages.

Three species of copepods have been described and assigned to the
genus Nesippus that are now certainly not members of this genus:
Nesippus curticaudis Dana, 1852, N. borealis Steenstrup and Lutken,
1861, and N. bengalensis, Gnanamuthu, 1949. These are described
as males of this genus. On the basis of the descriptions of the males
of 2 species of Nesippus in this paper, it has been shown that the above
3 species are not males of this genus. None of these species shows
any indication of sexual maturity. None has been reported in copu-
lation with a female. All have been reported from the plankton.
These 3 species, therefore, should be removed from this genus. Their
taxonomic position remains in doubt owing to the fact that they
appear to be immature. It cannot be established that they are even
members of the family Pandaridae.

Key to Adult Females of Genus Nesippus

Beermconod.or leg 4.UNATMCd, 2 i) Gis yeu ie seimy ie ey sen # mye seas 8H 2
Preacpod ole + withlong Setde. 24: koe cr uek s. gels cGy As 3

2. Genital segment conspicuously narrowed anteriorly, caudal rami with promi-
MENU SCUAC)) a cacy Sabb bthiey Sutsatet ch Gant tale. oii ehcan to per S060 5% crypturus

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Genital segment not conspicuously narrowed anteriorly, caudal rami with

small reduced:setaes .. .° = 4s. caw 2 iy fo ceesri. lotto ie feetignis
3. Fourth thoracic segment with alate plates. . ...... . .. . orientalis
Fourth thoracic segment without plates. ........... =. + «Vespa

Nesippus orientalis Heller, 1868
Figures 295-304

Nesippus orientalis Heller, 1868, p. 194.—Bassett-Smith, 1899, p. 459.—Brian,
1906, p. 49; 1924, p. 33.—Wilson, 1907, p. 457.—Capart, 1953, p. 658; 1959,
p. 96.—Rose and Vaissiere, 1953, p. 86.—Barnard, 1955, p. 265.—Nunes-
Ruivo, 1956, p. 22.— Yamaguti, 1963, p. 123.

Nogagus angustatus Beneden, 1892b, p. 245.

Nesippus alatus Wilson, 1905, p. 130; 1907, p. 426; 1932, p. 438.—Bere, 1936,
p. 595.—Heegaard, 1943b, p. 27.—Pearse, 1952b, p. 213.—Capart, 1953,
p. 659.—Rose and Vaissiere, 1953, p. 86.—Barnard, 1955, p. 265.— Yamaguti,
1963, p. 123.

Nesippus ornatus Thomsen, 1949, p. 17.—Yamaguti, 1963, p. 124.

Nesippus incisus Heegaard, 1962, p. 179.

Nesippus australis Heegaard, 1962, p. 178.

SPECIMENS STUDIED.—Five collections from Sarasota, Fla., from
the following hosts: Ginglymostomum cirratum, Galeocerdo cuvier,
Carcharinus leucas, and Carcharinus maculipinnis. A single collec-
tion from Sphryna zygaenae from Durham, South Africa. From Nosy
Bé, Madagascar, the following: 8 collections from Carcharinus maculi-
pinnis, 4 collections from Sphyrna lewini, 4 collections from Car-
charinus leucas, 1 collection from Galeocerdo cuvier, and 1 collection
from Scoliodon palasorrah.

FEMALE.—Body form as in figure 295. Total length 5.6 mm (based
on an average of 5 specimens). Greatest width 3.1 mm (measured
at widest point of cephalon). Cephalon rounded about as long as
wide. Thoracic segments 2—4 free with dorsal plates on segment 4.
Plates extending posteriorly slightly over anterior portion of genital
segment. Genital segment 2.1 mm long and 1.5 mm wide. Genital
segment of equal width throughout with posterior border trilobed.
Abdomen (fig. 296) 1-segmented. Caudal rami (see fig. 296) with 6
terminal setae, inner 4 plumose.

Oral area with adhesion pads associated with first and second
antennae and maxilliped. The pad of first antenna with posterior
margin produced to form a hooklike process (fig. 297). First antenna
2-segmented (see fig. 297). Second antenna clawlike; of usual
pandarid form. Mandible with 11 teeth at tip and of usual type.
First maxilla (fig. 298) with a broad process on outer distal corner.
Second maxilla (fig. 299) with tip produced to form a clear bulblike
tip. A short plumose spine and a small patch of hairs near base of
claw. Maxilliped (fig. 300) with a short blunt claw.

NO. 3570 PANDARIDAE—CRESSEY 61

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
end. exp. end. exp. end. exp. end.
seg. 1 1:0 Os Oe Dad Orda ste0 0:1 V:4 4
seg. 2 IV:3 3 IV:5 a IV:5 4

Leg 1 (fig. 301) with an interpodal adhesion pad as in figure. Leg 2
(fig. 302) armed as in figure. Leg 3 (fig. 303) with innermost seta of
last exopod segment reduced. Leg 4 (fig. 304) with each ramus
1-segmented and armed as in figure. Leg 5 small process bearing 2
short setae (see fig. 296). Leg 6 absent.

Egg strings long and straight.

Color in life cream.

Mate.—Wilson in 1905 described a male of Nesippus alatus
(=orientalis). I have examined this type specimen and have con-
cluded that it is immature and does not indicate the nature of the
adult male; consequently, the true adult male of this species is
unknown.

Discussion.—This copepod has been reported from a number of
sharks and is probably cosmopolitan in distribution. It seems to be
restricted to inshore species and is usually found in the mouth and
gill arches of the host.

Capart (1953) states that more material will eventually show
alatus and angustatus to be synonymous with orientalis. As a result
of my collections and the material I have examined, I am convinced
that this interpretation is correct and have placed the two former in
synonymy. Nesippus ornatus described by Thomsen in 1949 is also
the same as orientalis. Heegaard described 2 new species (incisus
and australis) in 1962. He noted the similarity to alatus and orientalis
and I believe that these 2 species should be placed in synonymy. I
have examined material from the North Atlantic, Caribbean, and
Indian Ocean and conclude that my own collections represent a single
cosmopolitan species that is found on a wide variety of hosts. The
exact shape of the dorsal thoracic plates of segment 4 should not be
regarded by itself as a specific character. The presence of this plate
separates it from the other known species. It also differs from other
species in the nature of the adhesion pad of the first antenna.

Nesippus crypturus Heller, 1868

Figures 305-324

Nessipus crypturus Heller, 1868, p. 196.—Bassett-Smith, 1899, p. 459.— Wilson,
1907, p. 425; 1935a, p. 3—Barnard, 1955, p. 265.— Yamaguti, 1963, p. 124.

Nesippus occultus Wilson, 1924b, p. 214— Yamaguti, 1963, p. 124.

Nesippus costatus Wilson, 1924b, p. 213.—Yamaguti, 1963, p. 124.

Nesippus gracilis Wilson, 1935a, p. 4.—Bere, 1936, p. 595.— Yamaguti, 1963, p. 124.

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

SPECIMENS STUDIED.—Single collections from Sarasota, Fla.,
from the following hosts: Sphryna mokarran, Galeocerdo cuvier,
Carcaharinus milberti, and Carcharinus leucas. <A single collection
from Socorro Island (Pacific) from Carcharyunus galapagensis. From
Nosy Bé Madagascar: Sphyrna lewina (4 collections), Carcharinus
maculipinnis (8 collections), C. leucas (4 collections), Scoliodon
palasorrah (1 collection), and Galeocerdo cuvier (1 collection).

FrmMALE.—Body form as in figure 305. Total length 6.8 mm (based
on an average of 5 specimens). Greatest width 3.5 mm (measured at
widest part of cephalon). Cephalon rounded, somewhat wider than
long (2.5 by 2.7 mm). Thoracic segment 2 expanded laterally.
Segment 3 small. Segment 4 expanded laterally somewhat but no
plates present. Genital segment 2.6 mm long and narrowed an-
teriorly; posterior border with deep median sinus. Abdomen (fig.
306) 1-segmented Caudal rami (see fig. 306) attached to abdomen
distally and with 6 terminal setae, inner 4 plumose and longer than
outer 2.

Oral area with adhesion pads associated with first and second
antennae and maxilliped (see fig 307). First antenna (fig. 308)
2-segmented, armed as in figure. Adhesion pad produced to form
a hooklike process. Second antenna (see fig. 307) of usual form.
Mandible and mouth tube of usual form. First maxilla (fig. 309) a
broad lobe with a small outer spine and an anterior group of 3 setae.
Second maxilla of usual type. Maxilliped (fig. 310) with a short
terminal claw rounded at tip. Tip of claw is opposed by a raised
area with a central crater-like depression.

Legs 1-4 biramose with the spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 I:0 OO leat Oe ei 0:1 V 0
seg. 2 EVES 3 VES 7 IV:4 4

Terminal exopod segments of legs 1-3 (figs. 311-313) with spines
armed as in figures. Setae of these legs pinched near base and often
broken off at this point (see figs. 311-313). Leg 4 (fig. 314) with
each ramus of 1 jomt. Endopod unarmed. Leg 5 (fig. 315) con-
sisting of 2 short setae, 1 naked, the other with a serrate fringe.
Leg 5 located lateral to caudal rami on ventral surface of genital
segment (see fig. 306). Leg 6 absent.

Eeg strings long and straight.

Color in life cream.

Maur.—Body form as in figure 316. Total length 5.4 mm (based
on an average of 2 specimens). Greatest width 3.1 mm (measured at
widest part of cephalon). Lateral margins of thoracic segment 2
somewhat alate. No dorsal plates present. Genital segment slightly

NO. 3570 PANDARIDAE—CRESSEY 63

longer than wide. Abdomen 1-segmented. Caudal rami (fig. 317)
with 4 terminal setae and 2 subterminal. Each ramus joined distally
to abdomen.

Oral area with adhesion pads as in female. First antenna as in
female. Second antenna (fig. 318) with terminal hook more recurved
than in female. An adhesion area present on antepenultimate seg-
ment. Remaining oral appendages as in female. Maxilliped with
tip of terminal claw produced as in figure 319. Legs 1—4 (figs. 320-323)
biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exp. end. exp. end. exp. end. exp. end.
seg. 1 1:0 0:0 1:0 O:1 Ted O:1 PV cary he 4.
seg. 2 IV:3 3 We Ded IV:4 4

Leg 5 (fig. 324) located near outer distal corner of genital segment
and consisting of 2 short, naked setae. Leg 6 absent.

Discussion.—This species, like N. orientalis, seems to be cosmo-
politan in distribution and occurs in the mouth and gill arches of a
variety of inshore species of sharks.

The copepod was originally described by Heller in 1865 and has

since been redescribed as N. occultus, N. costatus, and N. gracilis by
C. B. Wilson (1924; 1924b; 1935). I have examined Wilson’s types
of N. gracilis and determined this to be the same as N. crypturus.
Nesippus occultus from the published description also appears to be
‘synonymous. Nesippus costatus was described from an immature
female and was collected together with the material described as
N. occultus by Wilson. It appears to be merely a young female of
the type described as NV. occultus.
__ This copepod is characterized by a lack of dorsal plates, the nature
of the fourth leg of the female, and the deep median sinus on the pos-
terior portion of the female genital segment. The form of the male
maxilliped should separate this sex from other species of the genus.

Nesippus tigris, new species
Figures 325-345

SPECIMENS STUDIED.—T'welve females and 4 males from Galeocerdo
cuvier from Sarasota, Fla. Holotype female, allotype male, 3 para-
| bypes (299, 1o) deposited in alcohol in the U.S. National Museum.
Four paratypes (females) deposited in the British Museum (Natural
History). Remaining paratypes in the author’s collection. Addi-
tional specimens studied from Galeocerdo cuvier from Nosy Bé,
Madagascar (699).
| Fermate.—Body form as in figure 325. Total length 8.2 mm
(based on an average of 2 specimens). Greatest width 4.3 mm
(measured at widest part of cephalon). Cephalon rounded, slightly

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

wider than long (4.1 by 4.3 mm). Thoracic segments 2-4 free. No
dorsal plates present but lateral margins of segments 2 and 4 expanded
laterally somewhat. Genital segment 3.9 by 2.9 mm, widest at its
midpoint, its lateral margins somewhat irregular. Genital seement
with a median sinus posteriorly. Abdomen (fig. 326) 1-segmented.
Caudal rami (see fig. 8326) round, bearing 3 weak spines as in figure.

Oral area with adhesion pads associated with first and second
antennae and maxilliped. First antenna (figs. 327, 328) 2-seemented.

Each segment armed as in figures. All spines and setae of first
antenna naked. Adhesion pad of first antenna produced to form
a sharp, posteriorly directed spine (see fig. 327). Second antenna
(fig. 329) of usual form with 2 short setae on distal claw. Mouth
tube and mandible as in other members of genus. First maxilla
(fig. 330) with a median short, stout spine and an anterior group of
3 short setae. Second maxilla (fig. 331) with a short terminal claw
bearing rows of fringe. The penultimate segment with a stout
distal spine and a small patch of setae. Maxilliped (fig. 332) with a
stout terminal claw opposed by a raised area with a central depression
on penultimate segment. Maxilliped with an adhesion pad near base.
Pad produced in form of a sharp, posteriorly directed process (fig.
333). This process seemingly subdivided.

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 8 leg 4
exp. e

exp. end. exp. end. exp. end. nd.
seg. 1 1:0 0:0 Led Oia Til Ort IV 0
seg. 2 IVs3. 43 DWi5) 7 37 IV:4 4

Leg 1 (fig. 334) with both rami 2-segmented. Setae constricted
near their bases as in N. crypturus. First exopod segment with an
outer spine. Last exopod segment with 4 outer spines and 3 inner
setae. First endopod segment unarmed. Last endopod segment
with 3 terminal setae. Leg 2 (fig. 335) with both rami 2-segmented.
First exopod segment with an outer spine and an inner seta. Last
exopod segment with 4 outer spines and 5 inner setae. First endopod
segment with an inner seta. Last endopod segment with 7 terminal
setae. Leg 3 (fig. 336) with both rami 2-segmented. First exopod
segment with an outer spine and an inner seta. Last exopod segment
with 4 outer spines and 4 inner setae. First endopod segment with
an inner seta. Last endopod segment with 4 terminal setae. Leg 4
(fig. 337) with both rami 1-seemented. Exopod with 4 terminal
spines. Endopod unarmed. Leg 5 a single seta located on ventral
surface of genital segment near posterior corner (see fig. 326).

Eee strings long and straight.

Color in life cream.

Matr.—Body form as in figure 338. Total length 5.7 mm (based

NO. 3570 PANDARIDAE—CRESSEY 65

on an average of 2 specimens). Greatest width 3.6 mm (measured
at widest part of cephalon). No dorsal plates. Posterior corners of
segment 2 produced. Genital segment about as long as wide (1.6
by 1.6 mm). Abdomen 1-segmented. Caudal ramus (fig. 339)
somewhat rounded with 4 terminal plumose setae and 2 short sub-
terminal ones. Inner margin with short hairs.

Oral area as in female. Appendages of cephalon as in female except
for maxilliped. Maxilliped (fig. 340) with tip of claw papillose
(fig. 341). Adhesion areas as in figure 340. Adhesive pad of maxil-
liped (fig. 342) not pointed as in female but with posterior portion
produced as a rounded process with heavy striations as in figure.

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
xp. end. exp. end. exp. end. exp. end.
seg 1 1:0 0:0 Tet 0:1 Est 0:1 LVS ie 54
seg 2 TVicsy 3 UVic ie EVc5e "4:

Legs 1 and 2 asin female. Leg 3 with a modification on last endopod
segment as in figure 343. This is consistent with group II males,
although this modification is much reduced in this species. Leg
4 (fig. 344) with each ramus 1-segmented but not as reduced as the
female. Leg 5 (see fig. 345) a single short seta on margin of genital
segment. Leg 6 (see fig. 345) a single short seta near junction of
genital segment and abdomen.

Discussion.—This species has been collected twice from Galeocerdo
cuvier. The specimens from Nosy Bé were recovered from the nasal
passages of the host. Those from Sarasota were not collected by the
author, and there is no information as to their location on the host.

This species may be separated from N. orientalis by the nature of
the fourth legs of the females. It seems to be more closely related
to NV. erypturus but can be separated from it on the basis of the
caudal rami and by the fact that in N. crypturus the genital seg-
ment is considerably narrowed anteriorly whereas in N. tigris it is
not markedly so.

The name tigris, from Latin, meaning ‘‘tiger,” refers to the host,
the tiger shark.

Nesippus vespa Kirtesinghe, 1964
Figures 346-356
Nessipus vespa Kirtesinghe, 1964, p. 91.

SPECIMENS STUDIED.—Eleven females form Rynchobatus djeddensis
(Forskal) from Nosy Bé, Madagascar.

Frmate.—Body form as in figure 346. Total length 3.8 mm (based
on an average of 2 specimens). Greatest width 1.5 mm (measured at
widest part of cephalon). Cephalon rounded, slightly wider than

long (1.2 by 1.5mm). Thoracic segments 2 and 3 indistinctly divided
221-534-675

66 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

in dorsal view. No dorsal plates present. Genital segment globose,
about as long as wide (1.3 mm long by 1.2 mm wide). Genital
segment deeply incised posteriorly. Abdomen (fig. 347) joined distally
to abdomen and bearing 6 terminal setae, inner 4 long and plumose.
Inner margin of ramus with a row of hairs.

Oral area with adhesion pads as in other species of genus. Pad
associated with first antenna not produced to form a hooklike process
as in other species of the genus (see fig. 348). First antenna (fig. 348)
2-segmented, armed as in figure, and bearing only 12 spines on first
segment and 6 setae on the second. Second antenna (fig. 349) of
usual form. The 2 setae near base of claw longer than in other
species of genus. Mouth tube and mandible of usual pandarid type.
First maxilla (fig. 351) with a short spine and a small patch of setules
near base of the claw. Claw with rows of fringe as in figure. Maxil-
liped (fig. 352) bearing a blunt terminal claw opposed by a crater-like
area on opposite segment.

Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 8 leg 4
exp. end. exp. end. exp end. exp. end.
seg. 1 10 0:0 el OFT cat Ole FIVES 4
seg. 2 TV 3y 3 LV 550 4 IV:4 4

Leg 1 (fig. 353) with both rami 2-segmented. First exopod segment
with an outer spine. Last exopod segment with 4 outer spines and
3 inner setae. First endopod segment unarmed. Last endopod
segment with 3 terminal setae. Leg 2 (fig. 354) with both rami
2-sezmented. First exopod segment with an outer spine and an
inner seta. Last exopod segment with 4 outer spines and 5 inner seta.
First endopod segment with an inner seta. Last endopod segment
with 7 terminal setae. Leg 3 (fig. 355) with each ramus 2-segmented.
First segment of the exopod with an outer spine and an inner seta.
Last exopod segment with 4 outer spines and 4 inner setae. First
endopod segment with an inner seta. Last endopod segment with 4
terminal setae. Leg 4 (fig. 356) with each ramus 1-segmented.
Exopod with 4 outer spines and 3 inner setae. Endopod with 4
setae. Setae on legs 1-4 constricted near their bases (see fig. 355)
and often broken off at this point. Leg 5 (see fig. 347) a single
plumose seta located ventrally near distal margin of genital segment.
Leg 6 absent.

Egg strings long and straight.

Color in life cream.

Discusston.—This copepod has been collected only from Rhyn-
chobatus djeddensis in Madagascar and Ceylon. It was found in
the mouth of the host in Madagascar and on the body surface of the
Ceylon host.

NO. 3570 PANDARIDAE—CRESSEY 67

This species can be separated from N. orientalis by the lack of
the dorsal plate on fourth segment. It can be separated from N.
crypturus by the nature of adhesion pads of the cephalon and the
nature of the fourth leg of the female. It differs from N. tigris by
the nature of the fourth thoracic segment (wide in N. tigris, narrow
in N. vespa) and also by the nature of the caudal rami.

Genus Paranesippus Shiino, 1955
Paranesippus Shiino, 1955, p. 340. [Type-species: P. incisus.]

In 1955 Shiino described a new genus and species of parasitic
copepod on the basis of a single female specimen taken from the
body surface of Acanthidiwm eglantina (Jordan and Snyder).

In general, this copepod appears to be a member of group II and,
as the name implies, seems to be more closely related to Nesippus
than to other members of the family. Since Shiino has provided
a good description with illustrations, I shall not repeat his description
here. Because I had no material to study, I cannot add any more
details to the existing description.

Adhesion pads are associated with first and second antennae and
maxilliped.

The spine and setal formula of legs 1-4 (based on Shiino, 1955)
as follows:

leg 1 leg 2 leg 3 leg 4
xp. end. exp. end. exp. end. exp. end.
seg. 1 I:0 0:0 igi 0:1 ey O:1 ia Os
seg. 2 IVS menS Tea 0:2 I:1 0:2 ee 5
seg. 3 T5506 TE Aa +4 III:4

It is interesting to note that the egg strings remain recurved and
hidden beneath the genital segment as in some other members of group
II (emoleus latus and Dinemoura discrepans).

The male is unknown.

Comparative External Morphology and Taxonomic
Relationships Within the Family Pandaridae

It seems appropriate to examine the family Pandaridae as a group
and discuss the taxonomic features on a comparative basis. During
the course of the study, it became apparent that certain features
were quite stable, while others showed variations that should be
pointed out. In some cases these variat’ons can be explained rea-
sonably on the basis of the ecology of the parasite.

Body form: The body of the adult is composed of a cephalon (head
and first thoracic segment fused together), 3 free thoracic segments
(nos. 2-4), a genital segment (thoracic segments 5 and 6 fused), and
an abdomen (of 1 or 2 segments) that bears 2 caudal rami. In gen-

68 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

eral, the body of the female is more compact than that of the male.
The thoracic segments tend to be more broadly joined to each other,
making the females less flexible and adapted for a more sedentary
existence. The male is not modified in this way and is more mobile
than the female (compare figs. 1 and 16).

Cephalon: The cephalon is usually rounded in both sexes and
concave on the ventral surface. This configuration is quite stable
and found in most caligoid copepods. This is an obvious modifica-
tion to aid in attaching to the host, the cephalon forming a sucking
disc. The lateral borders usually bear a membrane. This thin
flexible membrane further enhances the holding ability of the cephalon.

Thoracic segments: In all cases in both sexes segment 1 is fused to
the head and segments 2—4 are free. The thoracic segments of the
female may bear dorsal plates in one of two arrangements, plates
present on segments 2-4 as in Pandarus, Pseudopandarus, Gangliopus,
Phyllothereus, Perissopus, and Pannosus, or plates present on segment
4 only as in Echthrogaleus, Demoleus, Pagina, Dinemoura, Nesippus,
and Paranesippus. This feature separates the females easily into
2 distinct groups.

Genital segment: The genital segment in both sexes is well formed.
In the female the genital segment may comprise more than one-half
of the total length (as in Pseudopandarus) and in all cases is at least
one-third of the total length of the copepod.

Abdomen: The abdomen of the female is 1-segmented in all genera
except Pagina and Dinemoura. The segments of the abdomen bear
dorsal plates except in the genera Nesippus and Paranesippus. ‘The
abdomen may be joined to the genital segment ventrally or terminally.
In the male the abdomen is of 2 segments except in the genus Nesippus,
in which there is only 1. None of the males have dorsal abdominal
plates.

Caudal rami: There is a great deal of variation in the form of the
rami in the females of this family. In the genus Pandarus the rami
are attached laterally to the abdomen and are often strongly sclero-
tized and elongated, terminating in a point (see figs. 2, 28, 35, 110).
In Pandarus bicolor (see fig. 101) the rami are not elongated but are
attached laterally to the abdomen.

It is interesting to note that the 2 genera Phyllothereus and Gang-
liopus, which are found only on gill filaments of the hosts, show a
great reduction in adhesion pads. Dinemoura ferox and D. discrepans
also have no adhesion pads. Both these species are found on the
body surface, and the loss of pads is unexplained in these cases.

First antenna: In both sexes of all members of this family the
first antenna is 2-segmented. The first segment bears terminally
a number of spines that are often covered with spinules or hairs.

NO. 3570 PANDARIDAE—CRESSEY 69

The last segment bears a smaller number of naked setae in all cases.
This appendage is not of great taxonomic importance on the generic
or specific level.

Second antenna: This appendage is well developed in all members and
serves as a principal means of attachment. The tip is always in the
form of a claw and this reaches its greatest development in the females
of the genera Phyllothereus and Gangliopus (see fig. 127). These
genera, as previously pointed out, show a reduction in adhesion pads
and also are the only pandarids found routinely on the gill filaments.
In view of this location on the host, adhesion pads would not be as

TaBLE 2.—Arrangement of female cephalic adhesion pads in genera of the family
Pandaridae (+ =present, — =absent)

Cephalic Adhesion Pads

Genus Al A2 base of between distal
Mxpd Mxpd corner of
cephalon

Pandarus
Pseudopandarus
Perissopus
Paranesippus
Pannosus
Nesippus hooklike
Dinemoura (part) double
Demoleus
Pagina
Echthrogaleus
Phyllothereus
Dinemoura (part) =
Gangliopus a

+++++
+tt+++4++
|
|

hooklike — —

P+ i ++t+++4++

+++

|

|
|
|

useful whereas a well-developed claw to envelope the gill filament
would be. The terminal claw always bears 2 setae.

Mouth tube: The mouth tube is composed of the labrum and labium,
forming a conelike structure, within which is housed the stylus of the
mandible. The tip of the labium is fringed (see figs. 6, 167, 251).
The labium extends beyond the tip of the labrum. The labrum bears
2 accessory spinelike structures at its tip (see fig. 251).

Mandible: The mandible is composed of a basal segment located
near the base of the mouth tube and a long stylet projecting within
the tube. The tip of the stylet is armed with 10-12 teeth. ‘This
appendage is of little taxonomic value and shows little variation from
species to species in both sexes.

First maxilla: This appendage is attached laterally to the base of

70 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

the mandible (see fig. 6) and is composed of a basal segment bearing
a group of 3 short setae anteriorly and a posteriorly directed process.
Within the family this appendage takes 2 basic forms. In members
of group I, plus Nesippus, the first maxilla is represented by a basal
segment fused to the cephalon along its entire length and bearing a
group of 3 short setae and a more prominent spine, generally directed
posteriorly. The appendage does not appear to have much mobility
(see figs. 6, 75, 113, 128, 168). In the genera Pagina, Echthrogaleus,
Dinemoura, and Demoleus (all of group II), the first maxilla is attached
to the cephalon at the anterior margin of the basal segment and gives
the appearance of greater freedom of movement (see figs. 224, 253,
269).

Second maxilla: The second maxilla in all species bears a clawlike
tip that usually bears rows of fringe or spinules. Group I can be
separated from group II by the nature of this appendage. In group I
there are 2 prominent spines near the base of the claw (see figs. 9,
76, 129, 149, 169). In group II the smaller spine is replaced by a
patch of setules or spinules (see figs. 204, 225, 254, 282). This
distinction is present in both sexes.

Maxilliped: The maxilliped is in all species a strongly developed
appendage. It undoubtedly aids in holding the copepod to the host
or is used by the male to hold the female during copulation. It is
interesting to note that, in group I, 3 genera found on the body surface
of the host possess a maxilliped with a spatulate tip on the claw
(Pandarus, Pseudopandarus, and Pannosus). This modification would
seem to enable the parasite to grip the denticles of the host better.
The remaining genus (Perissopus) of group I found on the body surface
of the host attaches by cementing the maxilliped to the host. The 2
genera of group I found on the gills have pointed claws on the maxilli-
ped. All members of group II have sharply pointed or rounded tips
on the claw of the maxilliped. None are spatulate.

Leg 1: This appendage is relatively stable in pandarid copepods,
but 2 genera can be separated from the rest of the family on the basis
of the endopod. In Pandarus the endopod is 1-segmented. In all
other known members of the family the endopod is 2-segmented.
In Perissopus the endopod is unarmed. In all other members of the
family the endopod bears 8 setae.

Leg 2: In group I the exopod of all species is 2-segmented. In
group IT all genera have 3-segmented exopods except Demoleus and
Nessipus, which have only 2.

Leg 3: The situation regarding this appendage is like that of leg 2.
All members of group II (except Demoleus, with only 2) have 3-
segmented exopods. Copepods of group I have a 2-segmented
exopod except Perissopus, which has only 1.

NO. 3570 PANDARIDAE—CRESSEY 71

Leg 4: In many genera of both groups this leg in the female is
lamelliform (Pandareus, Phyllotherus, Echthrogaleus, and Dinemoura).
The females of these genera can be separated from other female mem-
bers of the family on this basis alone. In the male the leg is not
modified and is of more usual form.

Leg 5: This leg is reduced in both sexes and is represented by a
group of 2-4 setae. This appendage is not useful in separating
members of the family.

Leg 6: This leg is incorporated in the female into the area of
spermatophore attachment and is more obvious in group II species
(see figs. 209, 284). In the male this leg is represented by 2 setae
located near the union of the abdomen and genital segment in all
members of the family in which the male is known.

It should be noted here that certain observations were made
regarding the determination of adult males. It was noted (see
page 14) that often, in collections, 2 sizes of males showing sperma-
tophores were present. A closer examination indicated that the larger
was more mature than the smaller (based on relative development of
the spermatophores). It was obvious that in a collection where
only the smaller ones were present that these might be erroneously
considered as mature. I found that in the larger forms the pad of
the second antenna was only about one-half the size of the pad of the
first antenna. In the smaller forms they are of about equal size.
The opinion is therefore presented that this difference may be a method
of determining sexual maturity of the male since the spermatophore
is seen in earlier stages and is not a dependable criterion. It was
observed that in mature males of other species of Pandarus this re-
duction of the pad of the second antenna was also present (see figs.
59, 60).

72 PROCEEDINGS OF THE

NATIONAL MUSEUM VOL. 121

A List of Sharks Examined with the Pandarid Copepods Recovered

from Them

(Number in parenthesis indicates collections made)

Hexanchus species

Demoleus paradoxus (1)
TIsurus oxyrhynchus

Pandarus smithii (8)

Dinemoura producta (1)

Dinemoura latifolia (3)

Echthrogaleus denticalatus (3)
Lamna nasus

Pandarus floridanus (1)

Echthrogaleus coleoptratus (1)
Lamna ditropis

Echtrogaleus coleoptratus (1)
Carcharodon carcharias

Pandarus smithit (1)

Pandarus floridanus (4)

Dinemoura producta (1)
Cetorhinus maximus

Dinemoura producta (1)
Alopias superciliosus

Dinemoura discrepans (2)

Pagina tunica (3)
Alopias pelagicus

Echthrogaleus denticulatus (1)
Alopias vulpinus

Pandarus smithii (2)

Dinemoura discrepans (2)

Echthrogaleus denticulatus (8)
Ginglymostomum cirratum

Nesippus orientalis (1)
Mustelus species

Perissopus dentatus (1)
Carcharinus milberti

Perissopus dentatus (1)

Nesippus orientalis (1)

Nesippus crypturus (1)
Carcharinus maculipinnus

Perissopus dentatus (1)

Pandarus sinuatus (2)

Pandarus carcharini (2)

Nesippus orientalis (5)

Nesippus crypturus (1)
Carcharinus leucas

Perissopus dentatus (1)

Pandarus sinuatus (2)

Pandarus carcharini (2)

Nesippus orientalis (5)

Nesippus crypturus (1)

Carcharinus gangeticus
Pandarus carcharini (1)
Carcharinus sorrah
Pandarus carcharini (1)
Carcharinus limbatus
Pandarus smithii (1)
Pandarus carcharini (1)
Carcharinus obesus
Pseudopandarus longus (1)
Carcharinus malpeloensis
Pandarus cranchii (1)
Pandarus smithii (1)
Pandarus katoi (5)
Carcharinus galapagensis
Pandarus cranchit (1)
Pandarus smithii (1)
Nesippus crypturus (1)
Carcharinus azureus
Pandarus smithii (2)
Pandarus katoi (2)
Carcharinus platyrhynchus
Pandarus katoi (1)
Pterolamiops longimanus
Pandarus cranchii (27)
Eulamia floridanus
Pandarus cranchii (12)
Pandarus smithit (4)
Echthrogaleus denticulatus (1)
Eulamia falciformis
Pandarus cranchii (2)
Pandarus smithii (2)
Eulamia obscura
Pandarus cranchii (2)
Hypoprion signatus
Pandarus smithii (1)
Prionace glauca
Pandarus satyrus (35)
Echthrogaleus coleoptratus (30)
Phyllothereus cornutus (8)
Gangliopus pyriformis (A)
Dinemoura producta (1)
Galeocerdo cuvier
Pandarus cranchit (2)
Nesippus tigris (2)
Nesippus orientalis (2)
Nesippus crypturus (2)

NO. 3570 PANDARIDAE—CRESSEY ie

Negaprion brevirostris Sphyrna lewini
Pandarus sinuatus (1) Nesippus orientalis (4)
Galeorhinus species Nesippus crypturus (4)
Pandarus niger (1) Sphyrna zygaenae
Pandarus carcharini (1) Pandarus cranchii (1)

Pandarus smithii (1)

Pandarus zygaenae (3)

Nesippus orientalis (1)
Squalus acutipinnus

Demoleus latus (1)
Squalus acanthias

Scoliodon palasorrah
Pseudopandarus gracilis (1)
Nesippus orientalis (1)
Nesippus crypturus (1)

Rhizoprionodon acutus

Pseudopandarus longus (1) Pandarus bicolor (1)
Sphyrna mokarran Rhynchobatus djeddensis
Nesippus crypturus (1) Nesippus nana (1)

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Krgyer, H. |
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NO. 3570 PANDARIDAE—CRESSEY Tee

Kurtz, H.
1924. Philodopus (Achtheinus) intermedius und Dissonus glaber, zwei neuy
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1930. Parasitic Copepoda. Fasc. 2 of vol. 3 in Resultats scientifiques du
voyage aux Indes Orientales Néerlandaises . . . , pp. 1-11.
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Norman, A. M., and Scort, T.
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1868-69. Prodromus faunae copepodorum parasitantium Scandinaviae.
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OorDE DE Lint, G. M. van, and SCHUURMANS STEKHOVEN, J. H., JR.

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eee icOupp:
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1828. Beschreibung einiger neuen, in den Jahren 1818 und 1819 im mittel-
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PearseE, A. S.

1952a. Parasitic Crustacea from the Texas coast. Inst. Mar. Sci. Port
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1952b. Parasitic crustaceans from Alligator Harbor, Florida. Quart. Journ.
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PEARSON, J.

1905. A list of the marine copepoda of Ireland, 1: Litoral forms and fish
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Presta, Orro

1934. Krebstiere oder Crustacea, 1: Ruderfiisser oder Copepoda, 6:

Caligoida. Pt. 29 in Dahl, Die Tierweit Deutschlands, 68 pp.

NO. 3570

Pocue, F.
1902.

RATHBUN,
1884.

1886.
1887.

RICHIARDI,
1880.

| Ross, M.,

. 1952a.
1952b.
1953.

SAY. cD.
1817.

Scott, A.
1904.

1929.

Scort, T.
1900.

1901.
1902.

1904.

PANDARIDAE—CRESSEY 79

Bemerkungen zu der Arbeit des Herrn Bassett-Smith: ‘A systametic
description of parasitic Copepoda found on fishes, with an enumera-
tion of the known species.”’ Zool. Anz., vol. 26, pp. 8-20. ([Refer-
ence not seen.]

R.

Annotated list of the described species of parasitic Copepoda
(Siphonostoma) from American waters contained in the United
States National Museum. Proc. U.S. Nat. Mus., vol. 7, no. 31,
pp. 483-492.

Description of parasitic Copepoda belonging to the genera Pandarus
and Chondracanthus. Proc. U.S. Nat. Mus., vol. 9, pp. 310-324.

Description of new species of parasitic copepods belonging to the
genera T'rebius, Perissopus, and Lernanthropus. Proc. U.S. Nat.
Mus., vol. 10, pp. 559-571.

S.

Contribution alla fauna d’Italia: Catalogo sistematico dei crostacei
che vivono sul corpo degli animali aquatici. Jn Catalogo degli
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and VAISSIERE, R.

Catalogue préliminaire des copépodes de |’ Afrique du Nord, 1. Bull.
Soc. Hist. Nat. Afrique Nord, vol. 43, pp. 113-136.

Catalogue préliminaire des copépodes del’ Afrique du Nord, 2. Bull.
Soc. Hist. Nat. Afrique Nord, vol. 48, pp. 164-176.

Catalogue préliminaire des copépodes de |’ Afrique du Nord, 3. Bull.
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An account of the Crustacea of the United States. Journ. Acad.
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Some parasites found on fishes in the Irish Sea. Trans. Biol. Soc.
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The copepod parasites of Irish Sea Fishes. Rep. Lancahire Sea-Fish.
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Notes on some crustacean parasites of fishes. 18th Ann. Rep. Fish.
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Notes on some parasites of fishes. 19th Ann. Rep. Fish. Bd. Scot-
land, pp. 120-153.

Notes on some parasites of fishes. 20th Ann. Rep. Fish. Bd. Scot-
land, pp. 288-299.

On some parasites of fishes new to Scottish marine fauna. 22d Ann.
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Scort, T., and Scort, A.

1913.

SHEN, C. J.

1958.

The British Copepoda, 1: Copepoda parasitic on fishes. Ray Soe.
London, vol. 2, 72 pls., 256 pp.

, and Wang, K. N.

A new parasitic copepod, Achtheinus impenderus (Caligoida, Pandari-
dae) from a shark taken at Peitaiho, Hopei Province. Tung Wu
Hseuh Pao, vol. 10, no. 1, pp. 27-31. [In Chinese with English
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80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Suino, 8S. M.
1954. Copepods parasitic on Japanese fishes, 5: Five species of the family
Pandaridae. Rep. Fac. Fish. Pref. Univ. Mie, vol. 1, no. 3, pp.
291-332.
1955. Paranesippus incisus n. gen., n. sp.: A new parasitic copepod of the
family Pandaridae. Pacific Sci., vol. 9, no. 3, pp. 349-353.
1957. Copepods parasitic on Japanese fishes, 13: Parasitic copepods col-
lected off Kesennuma, Miyagi Prefecture. Rep. Fac. Fish. Pref.
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1959a. Ostpazifische parasitierende Copepoden. Rep. Fac. Fish. Pref.
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1959b. Sammlung der parasitischen Copepoden in der Prafekturuniversitat
von Mie. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 2, pp.
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1960a. Copepods parasitic on fishes from Seto, Province Kii, Japan. Rep.
Fac. Fish. Pref. Univ. Mie, vol. 3, no. 3, pp. 501-517.
1960b. Copepods parasitic on the fishes collected on the Coast of Province Shima,
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Smitu, 8. I.
1874. Invertebrate animals of Vineyard Sound. Rep. Comm. Fish and
Fisher. 1871 and 1872, 478 pp., 38 pls.
STEBBING, T. R. R.
1910. General catalogue of South African Crustacea. Ann. South African
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Srrenstrovp, J. J. S., and Ltrxen, C. F.
1861. Bidrag til Kundskab om det aabne Havs Snyltekrebs og Lernaeer
samt om nogle andre nye eller hidtil kun ufuldstaen digt kjendte
parasitiske Copepoder, vol. 5, pp. 341-342.
SrePpHENsoON, K. H.
1940. Parasitic and semiparasitic Copepoda. Zool. Iceland, vol. 3, no. 34,
pp. 1-24.
Srvuarpo, J., and Facert, E.
1961. Copepodos parasitos chilenos, 1: Una lista de las especies conocidas
y descripcion de tres especies nuevas. Rev. Chilena Hist. Nat.,
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THOMSEN, R.

1949. Copepods parasitos de los pesces marinos des Uruguay. Comm. |

Zool. Mus. Hist. Nat. Montevideo, vol. 3, no. 54, pp. 1-41.
TuomseENn, G. M.

1889. Parasitic Copepoda of New Zealand. Trans. New Zealand Inst.,

vol. 22, pp. 353-376.
Worantiop Xs 1D).

1880. Crostacei parassiti dei pesci del mare Adriatico. Boll. Soc. Adriat.

Sci. Nat., vol. 6, pp. 55-90.
WItson, C. B.

1905. New species of parasitic copepods from Massachusetts coast. Proc. |

Biol. Soe. Washington, vol. 18, pp. 127-131.

1907. North American parasitic copepods belonging to the family Caligidae,
3 and 4: A revision of the Pandarinae and the Cecropinae. Proc. |

U.S. Nat. Mus., vol. 33, pp. 323-490, pls. 17-43.

1908. North American parasitic copepods: A list of those found upon the
fishes of the Pacific coast, with descriptions of new genera and
species. Proc. U.S. Nat. Mus., vol. 35, pp. 431-481.

i

NO. 3570 PANDARIDAE—CRESSEY Sl
Witson, C. B.
1911. North American parasitic copepods: Description of new genera and
species. Proc. U.S. Nat. Mus., vol. 39, pp. 625-634.
1912. Description of new species of parasitic copepods in the collections of
the U.S. National Museum. Proc. U.S. Nat. Mus., vol. 42,
pp. 233-243.
1914. The male of Pandarus satyrus Dana. Sci. Bull. Mus. Brooklyn Inst.
Arts and Sci., vol. 2, no. 4, pp. 71-72.
1920. Report on the parasitic Copepoda collected during the Canadian
Arctic Expedition, 1913-18. Rep. Canadian Arctic Exped.
1913-18, vol. 7, pt. L, pp. 3-16.
1922. Parasitic Copepoda in the collection of the Zoological Museum,
Kristiania. Medd. Zool. Mus. Kristiania, no. 4, pp. 1-7.
1923. Parasitic copepods in the collection of the Riksmuseum at Stockholm.
Ark. Zool., vol. 15, no. 3, pp. 1-15.
1924a. New North American parasitic copepods, new hosts and note on
copepod nomenclature. Proc. U.S. Nat. Mus., vol. 64, no. 2507,
art. 17, pp. 1-22.
1924b. Parasitic copepods from the William Galapagos Expedition. Zoo-
logica, Sci. Contr. New York Zool. Soc., vol. 5, no. 19, pp. 211-217.
1932. The copepods of the Woods Hole region, Massachusetts. U.S. Nat.
Mus. Bull. 158, 635 pp.
1935a. New parasitic copepods (Reports on the collections obtained by the
first Johnson-Smithsonian deep sea expedition to the Puerto
Rican deep). Smithsonian Misc. Coll., vol. 91, no. 3298, art. 19,
pp. 1-9.
1935b. Parasitic copepods from the Pacific Coast. American Midl. Nat.,
vol. 16, no. 5, pp. 776-797.
1936. Parasitic copepods from the Dry Tortugas. Jn Papers from the
Tortugas Laboratory. Carnegie Inst. Washington Publ. 452,
vol. 29, no. 12, pp. 327-347.
YAMAGUTI, S.
1936. Parasitic copepods from fishes of Japan, 3: Caligoida, 2, 21 pp.
1963. Parasitic copepods and Branchiura of fishes, 390 pp.
YamaGotl, S., and Yamasu, T.
1960. New parasitic copepods from Japanese fishes. Publ. Seto Mar,

Biol. Lab., vol. 8, no. 1, pp. 141-152.

82 PROCEEDINGS

OF THE NATIONAL MUSEUM

VOL. 121

SCALE A 4MM

Ficures 1-8.—Pandarus satyrus, female: 1, dorsal (A); 2, abdomen and caudal rami
ventral (D); 3, oral area, ventral (B); 4, first antenna (G); 5, second antenna (G); 6,
mouth tube and first maxilla (G); 7, mouth tube, lateral (G); 8, mandible (G).

No. 3570 PANDARIDAE—CRESSEY 83

SCALE B 4MM

SCALE C 2MM

SCALE D O.5MM

14

Ficures 9-15.—Pandarus satyrus, female: 9, second maxilla (D); 10, maxilliped (D); 11,
leg 1 (D); 12, leg 2 (D); 13, leg 3 (F); 14, leg 4 (F); 15, leg 5 (G).

84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

W Yj;

SSS
SSS
SY

Ti; 3
VU: Le K ES oD

i i

th 4 E
4

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X Oo
wn

Ficures 16-23.—Pandarus satyrus, male: 16, dorsal (A); 17, spermatophore (G); 18, .

caudal ramus (F); 19, second antenna (D); 20, maxilliped (F); 21, leg 1 (F); 22, leg 2)
(F); 23, leg 3 (F).

NO. 3570 PANDARIDAE—CRESSEY 85

E
E
=
re
w
a
<
oO
n

Ficures 24-30.—Pandarus satyrus, male: 24, leg 4 (F); 25, leg 5 (G); 26 leg 6 (E). P.
cranchii, female: 27, dorsal (A); 28, abdomen and caudal ramus (C); 29, maxilliped
(D); 30, endopod of second leg (G).

86

PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

~\
eae \ 3]

SCALE G 0.5mm

Ficures 31-37.—Pandarus cranchii, female: 31, area of spermatohpore attachment (D);
male: 32, second antenna (D); 33, posterior corner of genital segment (D). P. smithii,
female: 34, dorsal (A); 35, abdomen and caudal ramus (D); 36, caudal ramus, ventro-

lateral (F); 37, second antenna (G).

No. 3570 PANDARIDAE—CRESSEY 87

SCALE H O.imm

Ficures 38-43.—Pandarus smithii, female: 38, maxilliped (D); 39, leg 4 (F); 40, area of
spermatophore attachment (D); male: 41, dorsal (A); 42, genital segment and abdomen
(C); 43, leg 1 (D).

88 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

e

Me « «

f .~ IS

Ficures 44-50.—Pandarus smithii, male: 44, leg 2 (D); 45, leg 5 (G). P. floridanus, i
new species, female: 46, dorsal (B); 47, dorsal abdominal plate and rami (F); 48, oral .
area (F); 49, first antenna (E); 50, second antenna (G).

NO. 3570 PANDARIDAE—CRESSEY 89

Figures 51-58.—Pandarus floridanus, new species, female: 51, subterminal spine on
second maxilla (H); 52, leg 1 (D); 53, leg 2 (D); 54, leg 3 (F); 55, leg 4 (F); 56, area of
spermatophore attachment (D). Male: 57, dorsal (C); 58, caudal ramus (G).

221-534—67——_7

90 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

62

Ficures 59-68.—Pandarus floridanus, new species, male: 59, adhesion pads, adult (F); 60,
adhesion pads, young male (F); 61, leg 1 (F); 62, leg 2 (F); 63, leg 3 (F); 64, leg 4 (F);
65, leg 5 (G); 66, leg 6(H). P. sinuatus, female: 67, dorsal, adult female (B); 68, dorsal,
young female (B).

No. 3570 PANDARIDAE—CRESSEY 91

Ficures 69-77.—Pandarus katoi, new species, female: 69, dorsal (A); 70, abdomen and
caudal ramus (F); 71, first antenna (G); 72, detail of last segment of first antenna (H);
73, second antenna (G); 74, tip of mandible (?); 75, first maxilla (H); 76, second maxilla
(D); 77, maxilliped (D).

92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 78-85.—Pandarus katoi, new species, female: 78, leg 1 (D); 79, leg 2 (D); 80,
leg 3 (F); 81, leg 4 (F); 82, leg 5 (H). Male: 83, dorsal (B); 84, caudal ramus (D);
85, adhesion pads, oral area (F).

NO. 3570 PANDARIDAE—CRESSEY 93

Ficures 86-92.—Pandarus katoi, new species, male: 86, first antenna (G); 87, second
antenna (G); 88, first maxilla (E); 89, second maxilla (D); 90, maxilliped (D); 91, leg 1
(D); 92, leg 2 (D).

94 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 93-99.—Pandarus katoi, new species, male: 93, leg 3 (D); 94, leg 4 (D); 95, leg 5
(E); 96, leg 6 (E). P. xygaenae, female: 97, dorsal (A); 98, maxilliped (G); male: 99,
dorsal (B).

NO. 3570 PANDARIDAE—CRESSEY 95

ie
Was
ae
he

Qh
OX,
AN

(Pm

v

Ficures 100-108.—Pandarus bicolor, female: 100, dorsal (A); 101, abdomen and caudal
rami, ventral (C); 102, second antenna (G); 103, maxilliped (D); 104, leg 1 (D); 105,
leg 2 (F); 106, leg 3 (G); 107, leg 4 (F); 108, leg 5 (H).

96 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

110

Ficures 109-116.— Pandarus niger, female: 109, dorsal (A); 110, abdomen and caudal
ramus, ventral (F); 111, first antenna (G); 112, second antenna (G); 113, first maxilla
(E); 114, tip of second maxilla (E); 115, leg 1 (G); 116, leg 2 (G).

NO. 3570 PANDARIDAE—CRESSEY 97

a

a
ORs appre sisig

Ficures 117-123.—Pandarus niger, female: 117, leg 3 (G); 118, leg 4 (G). P. carcharini,
female; 119, dorsal (A); 120, abdomen and caudal ramus, ventral (F); 121, first antenna
(G); 122, leg 2 (F); 123, leg 3 (F).

98 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

oN
ZA

(\

Ficures 124-130.— Pandarus carcharini, female: 124, leg 4 (F). Phyllothereus cornutus,
female: 125, dorsal (A); 126, genital segment, abdomen, and caudal rami, ventral (B);
127, second antenna (C); 128, first maxilla (G); 129, second maxilla (F); 130, maxilliped

CE):

NO. 3570 PANDARIDAE—CRESSEY 99

a
“2

Ficures 131-137.—Phyllothereus cornutus, female: 131, leg 1 (F); 132, leg?2)3(C)54 133;
leg 3 (C); 134, leg 4 (C); 135, area of leg 5 (F). Male: 136, dorsal (A); 137, leg 5 (G).

100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 138-143.—Phyllothereus cornutus, male: 138, area of leg 6 (G). Gangliopus
pyriformis, female: 139, dorsal (A); 140, mouth tube with adjoining adhesion pad (D);
141, leg 5 (G); male: 142, dorsal (B); 143, genital segment and abdomen, ventral (C).

NO. 3570 PANDARIDAE—CRESSEY 101

144

'

Ficures 144-150.—Gangliopus pyriformis, male: 144, maxilliped (D); 145, leg 5 (E); 146,
leg 6 (E). Pseudopandarus gracilis, female: 147, dorsal (C); 148, abdomen and caudal
rami, ventral (G); 149, tip of second maxilla (H); 150, maxilliped (F).

102 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 151-156.—Pseudopandarus longus, female: 151, dorsal (C); 152, abdomen and ~
caudal rami, ventral (C); 153, first antenna (E); 154, second antenna (E); 155, tip of |
second maxilla (H); 156, maxilliped (D).

No. 3570 PANDARIDAE—CRESSEY 103

157

Ficures 157-163.—Pseudopandarus longus, female: 157, leg 1 (G); 158, leg 2 (G); 159,
leg 3 (G); 160, leg 4 (G); 161, leg 5 (H). Perissopus dentatus, female: 162, dorsal (C);
163, abdomen and adjacent area, ventral (D).

104. PROCEEDINGS OF THE NATIONAL MUSEUM vOL. 121

Ficures 164-170.—Perissopus dentatus, female: 164, first antenna (E); 165, second antenna
(G); 166, tip of second antenna (H); 167, tip of labium (I); 168, first maxilla (E); 169,
tip of second maxilla (H); 170, maxilliped (D).

570

PANDARIDAE—CRESSEY

|
|

Ficures 171-176.—Perissopus dentatus, female: 171, leg 1 (E); 172, leg 2 (E); 173, leg 3
(E); 174, leg 4 (E); 175, leg 5 (H); 176, leg 6 (H).

221-534—67—_8

106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Zp 181

Ficures 177-182.—Perissopus dentatus, male: 177, dorsal (F); 178, caudal ramus (E); |
179, second antenna (E); 180, maxilliped (G); 181, leg 1 (E); 182, leg 2 (E). |

—

NO. 3670

186, leg 6 (H).

PANDARIDAE—CRESSEY

187

Ficures 183-188.—Perissopus dentatus, male, 183, leg 3 (E); 184, leg 4 (E); 185, leg 5 (H);

Female: 187, dorsal (C); 188, dorsal (C).

107

108 PROCEEDINGS OF THE NATIONAL MUSEUM yOL. 121

Figures 189-191.—Perissopus dentatus, female: 189, dorsal (C). Dinemoura producta,
female: 190, dorsal (A); 191, oral area (B).

109

CRESSEY

PANDARIDAE

NO. 3570

Ficures 192-193.—Dinemoura ferox, female: 192, dorsal (J); 193, ventral Gj):

1iO PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

SCALE | 0.05mm

196

Figures 194-196.—Dinemoura latifolia, female: 194, dorsal (A); 195, oral area (B); 196,
area of legs 5 and 6 (F).

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LtZ PROCEEDINGS OF THE NATIONAL MUSEUM yOL. 121

201

\\\
WN
OTN

207

Ficures 201-208.—Dinemoura discrepans, new species, female: 201, mandible and first
maxilla (G); 202, tip of mandible (H); 203, second maxilla (F); 204, detail of claw of
second maxilla (G); 205, maxilliped (F); 206, leg 1 (F); 207, leg 2 (F); 208, leg 3 (F).

113

PANDARIDAE—CRESSEY

NO. 3570

MM
MUTI Q
STU a

female: 209, area of legs 5 and 6

new species,

Ficures 209-216.—Dinemoura discrepans,

, genital segment, ventral

212

(F); 210, egg string (no scale) (10X). Male: 211, dorsal (A);
(C); 213, endopod of leg 3 (G); 214, leg 4 (G); 215, leg 5 (H);

|
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|

leg 6 (E).

?

216.

114 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficure 217--221.—Dinemoura discrepans, new species, male: 217, attachment area of
spermatophore (E). Demoleus heptatus, female: 218, dorsal (A); 219, abdomen and |
caudal rami (C); 220, oral area (A); 221, process between maxillipeds (D).

No. 3570 PANDARIDAE—CRESSEY 115

Ficures 222-228.—Demoleus heptatus, female: 222, first antenna (D); 223, second antenna
(D); 224, first maxilla (G); 225, claw of second maxilla (G); 226, maxilliped (F); 227,
leg 1 (D); 228, leg 2 (D).

116 PROCEEDINGS OF THE NATIONAL MUSEUM you. 121

230

Ficures 229-234.—Demoleus heptatus, female: 229, leg 3 (D); 230, leg 4 (D); 23 1, leg 5 (G);
232, leg 6 (G).

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.
.

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.

:
'
:
|
|
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i
}
1
|
|

\
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1

|
|
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No. 3570 PANDARIDAE—CRESSEY 117

Ficures 235-242.—Demoleus hepta us, male: 235, maxilliped (D); 236, endopod of leg
3 (D); 237, leg 6 (H). D. latus, female: 238, dorsal (A); 239, oral area (C); 240, leg 5 (E);
241, leg 6 (E); 242, egg strings, abdomen removed (B).

118 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

245

SCALE _A
SCALE B
SCALE _C 2MM

yh

EE

re

BZ

SESE)
a,
SSD

<<

Sy

SSS

AS

SCALE D O.SMM

SCALE _E _O..MM

Figures 243-247.—Pagina tunica, female: 243, dorsal (A); 244, lateral (A); 245, caudal
ramus (C); 246, oral area, ventral (D); 247, first antenna (FE).

No. 3670 PANDARIDAE—CRESSEY 119

WP

248

MM

SCAU UE:
SCALE G 0.5MM

SCALE _H O,1MM

Figures 248-250.—Pagina tunica, female: 248, posterior end of genital segment and
abdomen, ventral (B); 249, second antenna (E); 250, mouth tube and adjacent append-
ages, postero-lateral (EF).

120 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

254

A Wa
Ae
N
Ny
th

A a \\
FANS

SN

——

bg

\

259 n| 4

Ficures 251-259.—Pagina tunica, female: 251, tip of mouth tube, antero-ventral (F);
252, tip of mandible (G); 253, first maxilla (H); 254, second maxilla (E); 255, maxilliped
(B); 256, leg 1 (B); 257, leg 2 (B); 258, leg 3 (B); 259, leg 4 (B).

No. 3570 PANDARIDAE—CRESSEY 721

Ficures 260-267.—Pagina tunica, male: 260, dorsal (A); 261, genital segment and abdo-
men, ventral (C); 262, second antenna (H); 263, maxilliped, ventral (E); 264, tip of
maxilliped, dorsal (H); 265, leg 3, endopod, 3rd segment, ventral (F); 266, leg 5, left side
(G); 267, leg 6 (G).

221-534—67—_9

122 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

268 Se. oe

269 270

Ficures 268-275.—Echthrogaleus coleoptratus, female: 268, dorsal (A); 269, first maxilla i
(G); 270, endopod of leg 1(G); 271, process between legs 1 (G); 272, leg 5 (E). Male: !
273, dorsal (B); 274, genital segment and abdomen (C); 275, leg 1 (G).

No. 3570 PANDARIDAE—CRESSEY 123

Ficures 276-282.—Echthrogaleus coleptratus, male: 276, leg 2 (G); 277, endopod of leg 3
(E); 278, leg 4 (G); 279, leg 5 (H); 280, leg 6 (H). £. denticulatus, female: 281, dorsal
(B); 282, second maxilla (G).

124 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 283-290.—Echthrogaleus denticulatus, female: 283, leg 5 (G); 284, leg 6 (G).
Male: 285, dorsal (B); 286, genital segment and abdomen (F); 287, endopod of leg 3
(E); 288, leg 4 (G); 289, leg 5 (H); 290, leg 6 (HF).

NO. 3570 PANDARIDAE—CRESSEY 125

Ficures 291-295.—Echthrogaleus torpedinis, female: 291, dorsal (A); 292, caudal ramus
(F); 293, legs 1-4 (F); 294, leg 5 (G). Nesippus orientalis, female: 295, dorsal (B).

126 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 296-302.—Nesippus orientalis, female: 296, abdomen and caudal rami (D); 297,
first and second antennae (D); 298, first maxilla (E); 299, second maxilla (G); 300,
maxilliped (G); 301, leg 1 (G); 302, leg 2 (G).

NO. 3570 PANDARIDAE—CRESSEY 127

Ficures 303-308.—Nesippus orientalis, female: 303, leg 3 (G); 304, leg 4 (G). WN. cryp-
turus, female: 305, dorsal (B); 306, abdomen and caudal rami (D); 307, first and second
antennae (C); 308, first antenna (G).

128 PROCEEDINGS OF THE NATIONAL MUSEUM VvOL, 121

315

Ficures 309-315.—Nesippus crypturus, female: 309, first maxilla (E); 310, maxilliped
(G); 311, leg 1, last endopod segment (E); 212, leg 2, last endopod segment (E); 313,
leg 3, last endopod segment (E); 314, leg 4 (E); 315, leg 5 (1).

No. 3570 PANDARIDAE—CRESSEY 129

321

SCALE K 0.2mm

GLE CCLE UE

ALE

Ficures 316-324.—Nesippus crypturus, male: 316, dorsal (B); 317, caudal ramus (G);
318, second antenna (G); 319, maxilliped (G); 320, leg 1 (K); 321, leg 2 (K); 322, leg 3
(K); 323, leg 4 (K); 324, leg 5 (H).

130 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficures 325-331.—Nesippus tigris, new species, female: 325, dorsal (B); 326, abdomen and
caudal rami, ventral (D); 327, first antenna (D); 328, detail of first antenna (H); 329,
second antenna (F); 330, first maxilla (L); 331, second maxilla (D).

NO. 3570 PANDARIDAE—CRESSEY 13]

<ll.’

ey,
eZ
335 oe

=

Si

Ficures 332-339.—Nesippus tigris, new species, female: 332, maxilliped (D); 333, adhesions
pad at base of maxilliped (K); 334, leg 1 (G); 335 leg 2 (G); 336, leg 3 (G); 337, leg 4 (G).
Male: 338, dorsal (B); 339, caudal ramus (G).

132 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Ficures 340-348.—Nesippus tigris, new species, male: 340, maxilliped (G); 341, tip of
maxilliped claw (H); 342, adhesion pad at base of maxilliped (E); 343, edge of last endopod
segment of leg 3 (H); 344, leg 4 (G); 345, area of legs 5 and 6 (G). N. vespa, female:
346, dorsal (C); 347, abdomen and caudal rami (K); 348, first antenna (L).

No. 3570 PANDARIDAE—CRESSEY ts

\

399

Ss;

SS

X\

0.

SCALE L

Figures 349-356.—Nesip pus vespa, female: 349, second antenna (L); 350, first maxilla
(H); 351, claw of second maxilla (H); 352, maxilliped (L); 353, leg 1 (L); 354, leg 2 (L);
355, leg 3 (L); 356, leg 4 (L).

U.S, GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION . WASHINGTON, D.C.

Volume 121 1967 Number 3571

SUPPLEMENTARY DESCRIPTIONS
OF TWO MYODOCOPID OSTRACODS
FROM THE RED SEA

By Louts S. Kornicker
Associate Curator, Division of Crustacea

This study is based on ostracods collected on the Pola Expedition
to the Red Sea in 1896 and described by Dr. Herbert Graf (1931). In
the original description of Philomedes polae (=Euphilomedes polae),
Graf (1931) did not designate a holotype and omitted some details of
carapace and appendage morphology. Graf’s description may have
been based on more than one species. The species, therefore, is
redescribed herein and a lectotype selected from the available type-
series. Specimens designated as Philomedes species by Graf (1931)
have been identified as females of E. polae and are described as such.
A single specimen included by Graf in the type-series of Philomedes
polae has been identified as Euphilomedes arostrata Kornicker
(1967); this identification is documented with a description and
illustrations of the Red Sea specimen.

The following material was obtained through the courtesy of Dr.
Gerhard Pretzmann from the Naturhistorisches Museum, Vienna,
Austria: (1) One bottle, containing one vial with 3 males, with red
label marked: ‘““Philomedes polae, n. sp. o&*; Rotes Meer: Gulf von Suez;

il

2 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Coll: ‘Pola’ 1895/8; Dr. Graf det.” (2) One bottle with white label
marked: “Philomedes sp. 9; Rotes Meer: Stat. 90; Coll. ‘Pola’ 1895/8;
Dr. Graf det.” This bottle contained one vial with 2 females con-
taining eggs that had not yet been extruded into the brood chamber
and with a label that had been marked ‘‘Philomedes sp. ? (A).”

Since Dr. Graf had not designated a holotype of Philomedes polae
Graf, Dr. Pretzmann suggested that I select a lectotype from the type
material. Examining the material, I found 2 specimens in the first
bottle that I could equate with Philomedes polae Graf, using primarily
as a basis for identification the distinctive secondary appendage
described and figured for this species by Graf (1931, p. 37, fig. 51).
The third specimen in the first bottle I identify as Euphilomedes
arostratus Kornicker, recently described for the first time from the
Maldive Islands, Indian Ocean.

The 2 specimens in the second bottle I equate to specimens
described by Graf (1931) as Philomedes species on his page 38 and
illustrated in his figure 6. The similarity of the distribution of
primary and secondary claws on the caudal rami of the specimens at
hand with the published description of Philomedes species makes it
certain that this identification is correct. After studying these speci-
mens, I have concluded that they are females of Philomedes polae Graf.

I acknowledge with thanks the assistance of Dr. Gerhard Pretzmann
in obtaining specimens for study from the Naturhistorisches Museum,
Vienna, Austria, and of Miss Caroline Bartlett, who inked the final
illustrations. I would like also to thank Mr. I. G. Sohn and Doctors
Raymond B. Manning and W. Duane Hope for reviewing the manu-
script.

Family Cypridinidae Baird, 1850
Subfamily Philomedinae G. W. Miiller, 1912

Genus Euphilomedes Poulsen, 1962

Typn-spectes.—Euphilomedes nodosus Poulsen, 1962, by subsequent
designation, Kornicker (1967).

Euphilomedes polae (Graf, 1931)
FiaureEs 1-5

Philomedes polae Graf, 1931, p. 37, fig. 5.
Philomedes species.—Graf, 1931, p. 38, fig. 6.
LecroryrE.—Specimen designated as specimen B on slide. Natur-

historisches Museum, Zoologische Abteilung, Vienna, Austria.
Srx.— Male.

NO. 3571 RED SEA OSTRACODS—KORNICKER 3

Ficure 1.—Euphilomedes polae, male, specimen C (except a and h, spec. B): a, lateral view
left valve showing some muscle scars, surface reticulation shown only near posteroventral
margin of valve; b, lateral view anterior end of left valve showing surface and marginal
hairs; c, lateral view anterior end of left valve showing flap over antennal sinus, and linear
sclerotized area visible in transmitted light; d, lateral view posterior end of left valve
showing surface hairs and some reticulations; ¢, medial view of part of shell posterior
showing reticulations, normal pore canals and surface hairs; f, lateral view anterior end
of right valve showing marginal denticulations around rostrum, flap over antennal sinus
and linear sclerotized area; g, medial view anterior end of right valve; 4, medial view
anterior end of left valve; 7, medial view of middle section of right valve showing adductor
muscle scars below and dorsal muscle scars above; 7, medial view of dorsal muscle scars
below anterior hinge element and oval lucid area below muscle scars; k, medial view right
valve showing minute reticulations within large reticulations on surface of valve. (Same
scale, in microns: b,¢,d,f,i, ¢,g,h,7-)

4 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

PARALEcTOTYPE.—1 male designated as specimen C, Naturhisto-
risches Museum.

ADDITIONAL SPECIMENS.—2 females from the type-locality desig-
nated as specimens 1 and 2 on slides, N aturhistorisches Museum.

TypE-LOCALITY.—Graf (1931) reported 4 specimens of Philomedes
polae from Pola Expedition Station 90 and 1 specimen from Station 89;
both stations are in the Gulf of Suez; all specimens are males. The
vial containing specimens of P. polae did not bear a station number;
however, as 3 specimens were in the vial, it is probable that all, and
at least 2 of them, are from Station 90 rather than 89. Station 90,
therefore, is considered here to be the type-locality; since Stations 89
and 90 are close to each other, however, whether the lectotype was
collected at one or the other is not of major significance. Females of
P. polae, designated by Graf as Philomedes species, were collected at
Station 90. Data concerning the stations are as follows:

station latitude longitude collecting depth date collected
89 28°40’N 32°57'/E surface March 31, 1896
90 28°00’N 33°36’/E surface April 1, 1896

DESCRIPTION OF MALE.—Shell (figs. 1, 2a-e): oval, elongate with
ereatest height near middle, prominent rostrum and shallow rostral
incisure (fig. 1a); anterior margin of rostrum and anteroventral
marein of shell with scalloped outline formed by cresent-like marginal
denticulations; alate shield projecting laterally and anteroventrally
from behind rostral incisure, partly covering incisure, flap strongly
sclerotized anteriorly (figs. 1¢,f). Surface of valves with irregular
polygons; polygons with pebbly texture appearing as minute polygons
under high magnification (figs. 1e,). Posterior dorsal margin with
linear hinge depressed below outline of shell (figs. 26,c); hinge not
visible in lateral view except by transmitted light. Posterior hinge
element of each valve consisting of angular sclerotized process (figs.
2c,d); medial hinge element straight; anterior hinge element not
prominent. Left valve broadly overlapping right valve along antero-
dorsal margin. About 20 individual muscle scars (partly obscured
by muscles in specimen examined) clustered behind and below middle
of valve; small scar and large oval hyaline spot situated some distance
above others; 2 large and 4 smaller scars situated below anterior hinge
element (figs. 1a,i,j). Inner lamella broad with vestibule; no parallel
striations or line of concrescence observed on inner lamella. Selvage
with wide, corrugated, lamellar prolongation having fringe of slender
spines along outer margin. Six long hairs bearing secondary spines,
forming row on inner lamella behind rostrum (figs. 19,4); inner lamella
ventral to rostral incisure with small plumose hair, followed by wide
space and then about 5 plumose hairs (fig. 2a); about 30 hairs on
posteroventral part of inner lamella (figs. 2d,e).

NO. 3571 RED SEA OSTRACODS—KORNICKER 5

=

1
Bp
1
\

ss

Figure 2.—Euphilomedes polae, male, specimen B (except c, spec. C): a, medial view of
anteroventral shell margin of left valve showing medial hairs; b, medial view of left valve
hinge area (anterior on right); c, medial view of posterior hinge element on right valve;
pustules along middle hinge element are dotted because they could not be seen with
certainty on specimen C and were not observed on specimen B; d, medial view marginal
area at posterior end of left valve showing medial hairs; ¢, medial view posteroventral and
posterodorsal margin showing medial hairs; f, lateral view of right Ist antenna (only
proximal part of c- and f-bristles shown); g, exopodite of 2nd antenna; h, distal end 3rd
joint of endopodite of 2nd antenna; i, protopodite and endopodite of 2nd antenna; 7, medial
view Ist endopodite joint of left mandible, distal bristles are on ventral margin (joint
foreshortened by oblique position of joint in mounting medium); k, medial view 2nd
endopodite joint of left mandible, dorsal margin to right; Z, lateral view end joint of right
mandible, all bristles not shown. (Same scale, in microns: a,b,e,f,g,1; c,d,j,k; hl.)

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Hairs with either pointed or blunt ends distributed on rostral surface
(fig. 1b), some forming row near margin of shell; slender hairs with
pointed ends on posterior surface of shell, some forming row near pos-
terior margin (fig. 1d); a long tapered hair projecting from base
immediately behind posterior hinge element of each valve (figs.
2c-e); all hairs coming through normal pore canals; radial pore
canals not observed.

Dimensions: Lectotype (specimen B) greatest length 1.12 mm,
greatest height 0.63 mm; paralectotype (specimen C) greatest length
1.12 mm, greatest height 0.63 mm. Graf (1931, p. 38) gives length
of this species as 1.05 mm and height 0.60 mm.

First antenna (fig. 2f): First joint with faint clusters of short
hairs on medial surface. Second joint with clusters of long hairs on
medial surface, and distally, a dorsal, ventral, and lateral bristle;
all bristles bare or with short hairs, none with wreaths of long hairs.
Third joint with a few clusters of short hairs on medial surface, and 1
ventral and 2 dorsal bristles, the longer dorsal bristle provided with
wreaths of long hairs near middle and short hairs distally;
other dorsal bristle and ventral bristle with short hairs. Fourth
joint with 2 dorsal bristles, each with wreaths of long hairs and the
more proximal bristle with short hairs distally, and 4 long ventral
bristles subterminally, latter bristles bare or with short hairs distally.
Fifth joint, inferred to be inserted ventrally between 4th and 6th
joints, bearing sensory bristle with broad base and provided with
numerous filaments. Sixth joint with single bristle on 1 limb
and without bristle on other limb (bristle may have been broken off
during dissection). End joints with 5 slender bristles and 2 long
stout c- and f-bristles.

Second antenna (figs. 2g-2): Protopodite triangular with trans-
parent rim (fig. 27). Exopodite (fig. 2g): 1st joimt elongate; 2nd
joint about % length of 1st; 3rd joint about same length as Ist; distal
margins of joints 2-8 with comb of short spines (not shown on fig. 29);
basal spines not observed. Bristle of 2nd joint about % length of
3rd joint, bearing short marginal spines. Bristles of joints 3-8
bearing natatory hairs, without marginal spines. End joint with 4
stout bristles with natatory hairs and 2 short slender bristles. Endop-
odite 3-jointed (figs. 24,7); Ist joint with 5 bare short bristles and
1 long distal bristle with wreaths of long hairs near middle and short
spines distally; 2nd joint elongate, with 2 subterminal bristles pro-
vided with short spines; 3rd joint elongate, broad proximally, with
1 short proximal bristle and 2 longer subterminal bristles; tip with
short spine and about 5 serrate ridges.

Mandible (figs. 2j-l, 3a,b): No coxale endite observed. Basale
(fig. 3b): ventral margin with 3 short slender spinous bristles, and 2

NO. 3571 RED SEA OSTRACODS—KORNICKER a

\\ Ny Ny

ell Hah ght «
NX ANS & N \, \ NINES Greco 7

KY,
ae \\

Ficure 3.—Euphilomedes polae, male, specimen B: a, medial view of exopodite and Ist
endopodite joint without distal bristles of left mandible; }, lateral view of basale and
exopodite of right mandible; c, maxilla, all bristles not shown; d, 5th limb, hairs not
shown on all epipodial bristles; ¢, endites and exopodite of 5th limb, all bristles not shown;
f, 6th limb; g, 7th limb (1 bristle in distal group broken); h, distal end of 7th limb; 7, frontal
organ and median eye; j, lateral view of right furcal lamella; k, copulatory organ; /, detail
of distal tip of copulatory organ; m, lateral eye. (Same scale, in microns: a, e, h, 1;
b, f, i, m3 ¢, g; d, 7, R.)

8 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

long bristles with wreaths of long hairs and short spines distally.
Dorsal margin with 3 bristles, 1 near middle and 2 subterminally.
Medial surface with about 5 short bristles proximally near ventral
corner and 1 longer bristle near ventral margin at middle of joint.
Exopodite reaching middle of Ist endopodite joint, with 2 subequal
terminal bristles with short marginal spines (fig. 3a); tip of joint with
blunt hirsute process bearing short spine. Endopodite: First joint
(fig. 2j) with 1 short spinous bristle and 3 long hirsute posterior
bristles distally. Second joint (fig. 2/) anterior margin with proximal
eroup of 3 bristles and distal group of 5 bristles, all bristles bare or
with short spines; posterior margin with 2 spinous bristles distal and 1
short annulated bristle and 2 clawlike bristles subterminally. End
joint (fig. 27) with 2 large subequal claws, 1 short anterior claw, and 3
annulated bristles. Medial surface of basale and 1st and 2nd joints
of endopodite provided with groups of hairs.

Maxilla (fig. 3c): Very small. It seems to have a structure typical
of the genus, but because of its size I have not attempted to describe
it fully.

Fifth limb (figs. 3d, e): Epipodial appendage with about 47 plumose
bristles. Outer lobe of 3rd exopodite joint with 2 stout plumose
bristles.

Sixth limb (fig. 3f): First endite with 3 bristles, 2nd endite with
4 bristles, 3rd endite with 6 bristles, 4th endite with 6 bristles, 2nd
joint of exopodite with 14 bristles, joint not produced posteriorly ;
epipodial appendage represented by 2 short bare bristles. Surface of
second joint of exopodite with clusters of short hairs.

Seventh limb (figs. 3g, 4): Cleaning bristles: 4 in distal group, each
with 4 or 5 bells; 3 in proximal group, each with 1 to 3 bells; surface
hairs not observed on bristles. ‘Terminal comb with about 7 marginal
teeth, some with marginal spines. Two long curved pegs opposing
comb.

Copulatory organ (figs. 3k, /): Long, slender, divided into 3 lobes,
each with 2 annulated bristles; 1 lobe with large curved sclerotized tip.

Furea (fig. 3): Each lamella with 12 to 13 claws: primary claws
number 1, 2, 6, 10; secondary claws 3, 4, 5, 7, 8, 9, 11, 12, 18 (13th claw
not always present). Primary claws 1 and 2 separated from lamella,
6 and 10 joined to lamella; secondary claws all separated from lamella;
all claws with double row of spines. Primary claws decrease in length
proximally on lamella. Lamella near claws 1 and 2 with clusters of
long hairs. (One specimen has 13 claws on each lamella, another
specimen has 13 claws on the right lamella and only 12 on the left
lamella.)

NO. 3571 RED SEA OSTRACODS—KORNICKER 9

Ficure 4.—Euphilomedes polae, female, specimen 2 (except a-d, j-l, spec. 1): a, lateral
view left valve, dashed area near valve middle contains adductor muscle scars, not all
surface polygons shown; 4, medial view of anterior end of left valve, sclerotized areas
stippled; ¢, medial view anterior of right valve, sclerotized areas stippled; d, lateral view
of anterior end of left valve showing surface and marginal hairs; ¢, medial view of antero-
ventral margin of right valve showing distribution of hairs on inner lamella; f, medial
view of valve near posterior showing large and small reticulations; g, medial view hinge
element of right valve (anterior end on right); h, medial view of muscle scars on right
valves, adductor muscle scars clustered at lower left of figure, anterior of valve to left;
2, medial view of posterior and ventral marginal area of right valve showing distribution
of hairs on inner lamella; 7, lateral view of left 1st antenna; &, lateral view of endopodite,
first 3 joints of exopodite and distal margin of protopodite of left 2nd antenna; /, medial
view of right 2nd antenna, ends of some bristles of exopodite not shown. (Same scale,
in microns: b-d, f, k; e, g-7, l.)

221-535—66——_2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Eyes: Lateral eyes large; about 19 ommatophores visible in side
view, all except 2 divided by a suture into 2 parts (fig. 3m). Medial
eye large, pigmented (fig. 37).

Frontal organ (fig. 37): Elongate, 2-jointed with short spines on
distal half of 2nd joint and subterminally on basal joint.

DeEscrRIPTION OF FEMALE.—Shell (figs. 4a—72): lateral outline more
ovoid than male (fig. 4a); hinge structure (fig. 4g) selvage similar to
male; adductor muscle scars (figs. 4a,h) more anteriorly and ventrally
located on shell than on male; rostrum narrower and incisure shallower
than on male; irregular polygons on surface of shell subdivided by
minute polygons larger than on male shell and visible under low
magnification (fig. 4f/). Inner lamella with parallel striations on
anteroventral part; 9 long hairs with secondary spines forming row
on inner lamella behind rostrum (figs. 40,¢c); inner lamella below rostral
incisure with small hair followed by wide space and then 5 hairs (fig.
4e); posteroventral part of inner lamella with at least 21 hairs (fig. 47);
lateral surface of rostrum and anterior part of shell with numerous
hairs with blunt ends (fig. 4d).

Dimensions: Specimen no. 1 greatest length 1.06 mm, greatest
height 0.86 mm; specimen no. 2 greatest length 1.03 mm, greatest
height 0.73 mm. Graf (1931, p. 38) gives length as 0.90 mm and
1.00 mm and height as 0.70 mm.

First antenna (fig. 47): First and 2nd joints with surface hairs.
Second joint with 3 bristles, 1 dorsal, 1 ventral, and 1 lateral. Third
joint with 1 ventral bristle distally and 1 short bare bristle and 1 long
hirsute bristle on dorsal margin; ventral margin with short spines.
Fourth joint with 2 dorsal subterminal bristles and 1 short and 3 long
ventral subterminal bristles; ventral margin with short spmes. Fifth
joint with stout ventral terminal bristle with filaments. Sixth joint
with medial bristle distally. End joints with 1 short bristle and 2
long bristles without filaments and 4 long bristles with filaments.

Second antenna (figs. 44,1): Protopodite subtriangular without
transparent rim. Exopodite (fig.4l) with 9 joints; 1st joint elongate
with small medial spine; 2nd to 9th joints trapezoidal without basal
spines; distal margins of 2nd to 5th joints with comb of slender spines;
2nd and 3rd joints with bristles without natatory hairs or marginal
denticulations; 4th to 8th joints with bristles having natatory hairs
and marginal denticulations; 9th joint with 2 short bare bristles and
2 subequal long bristles with natatory hairs and marginal denticula-
tions. Endopodite (figs. 44,7) with 2 joints; basal joint with long stout
bristle with wreath of long hairs near middle, and 3 to 5 short bare
bristles; distal joint elongate with small spine at tip and long stout
terminal bristle with wreaths of long hairs near middle and short
distal spines.

NO. 3571 RED SEA OSTRACODS—KORNICKER i

Ficure 5.—Euphilomedes polae, female, specimen 1: a, medial view right mandible (surface
hairs not shown); d, lateral view left maxilla, all bristles not shown; c, posterior view left
5th limb; d, medial view right 6th limb; ¢, 7th limb; f, distal end of 7th limb; g, distal end
frontal organ; h, frontal organ and medial eye; 1, medial view right lamella of furca;
j, lateral view left lamella of furca. (Same scale, in microns: b,d,e,h-j; c,f,g.)

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Mandible (fig. 5a): Coxale endite large, bifurcate, with rows of
spines. Basale: ventral margin with 4 short bristles followed by 1
longer bristle near distal end; dorsal margin with 3 bristles, 1 near
middle and 2 at distal end; medial surface with 1 bristle at middle
and 5 shorter bristles proximally, all near ventral margin. Exopodite
short with 2 bristles and triangular process at tip, inner bristle with
wreath of long hairs near middle. Endopodite: distal end of 1st joint
with 1 short and 3 long bristles ventrally; anterior margin of 2nd joint
with proximal group of 4 and distal group of 5 bristles; posterior margin
with 2 subterminal bristles and terminally with 1 short annulate
bristle and 2 clawlike bristles; distal joint with 2 long stout claws of
subequal length and 3 bristles. Surface of basale and 2nd joint of
exopodite hirsute.

Maxilla (fig. 56): Precoxa and coxa with fringe of fine hair along
dorsal margin; coxa with short bare bristle on dorsal margin; basale
with 3 distal bristles; 1st endopodite joint with marginal hairs, a
hirsute bristle terminal on anterior margin and about 5 bristles on
distal margin; distal end of endopodite joint provided with numerous
bristles; exopodite with 2 long and 1 short bristle. Three endites: 1st
endite with about 8 bristles; 2nd endite with about 4 bristles; 3rd
endite with about 6 distal and 1 proximal bristle (bristles not shown
in illustration).

Fifth limb (fig. 5e): Epipodial appendage with about 49 bristles.
Triangular tooth anterior to main tooth of 1st exopodite joint about
same size as distal tip of 2nd exopodite joint. Distal anterior margin
of 1st jomt with 2 centrally located bristles. Large curved tooth
of 2nd exopodite joint without bristle at distal lateral corner.

Sixth limb (fig. 5d): First endite with 3 bristles; 2nd endite with
4 bristles; 3rd endite with 7 bristles; 4th endite with 6 bristles;
epipodial appendage represented by 2 short bristles. Second joint
of exopodite not produced posteriorly, distal margin with 13 to 14
bristles, medial and lateral surfaces hirsute.

Seventh limb (figs. 5e,f): Five cleaning bristles in distal group,
3 in proximal group, each bristle with 1 to 5 terminal bells. Terminal
comb with about 9 teeth, secondary teeth at base (all teeth not shown
in illustration); 2 elongate pegs opposing comb.

Furea (figs. 57,7): Each lamella with 4 primary claws and 10 to 12
secondary claws. Primary claws 1 and 2 separated from lamella,
7 and 12 (or 11) joined to lamella. Primary claws decrease in length
proximally on lamella. Secondary claws 3 to 6 about same length,
shorter than claw 7, longer than claw 8; claws 8 to 12 (or 11) decrease
in size proximally. Primary claw 12 (or 11) about same size or longer
than preceding secondary claw and longer than following secondary

NO. 3671 RED SEA OSTRACODS—KORNICKER iS

claw. Lamella near basis of claws with long hairs. Primary claws
1 and 2 with double row of teeth, claws 7 and 12 without teeth;
secondary claws with marginal spines. Distribution of primary and
secondary furcal claws on 2 specimens is as follows:

claw number
eS e On Gl KV Seem Om nl Otmulinn Lehn LOnecle ton 316
specimen 1
right DieeDinwSs)s Se ulis*) Se Gp angisee USk She SH Diy HS Shh BVGS

left Dope se CS. .58) {Si PP usmaas. ics S498) os
specimen 2

right Dia p os «Sy (48) IS ~ sph Sis, Ss! sw ep. ssl is

left DMC» Spats. Asa his. SDruisa bse Sia) Sto Spierse aS

Eyes: Large medial eye (fig. 54); lateral eyes not observed.

Frontal organ (figs. 5g,h): 2-jointed with distal joint 3 times length
of proximal joint; tip rounded; surface of 2nd joint and distal end of
1st joint with fine spines.

Remarxks.—Extreme sexual dimorphism among some species of
ostracods creates inherent difficulties in relating males and females
with certainty. Carapace morphology in the region of the rostrum
and rostral incisure between males and females of Huphilomedes polae
identified herein is sufficiently similar to warrant considering them as
conspecific in spite of some differences in appendage morphology.
Both males and females were collected in the same area. The male
and female of FE. arostratus Kornicker, a closely related species, have
similar dimorphism.

Some differences in morphology between males and females of /.
polae and E. arostratus are compared as follows:

E. polae E.. arostratus
male female male female
number of secondary fureal claws 8-9 10-12 7 9
cleaning bristles in distal group of 7th limb oh 5 4 5
number of hairs in row on inner lamella
behind rostrum 6 9 9-11* 11

*The male of E. arostratus described from the Maldive Islands (Kornicker,
1967) has 9 to 10 hairs, whereas the Red Sea form described in this paper has 11.

Comparisons.—EHuphilomedes polae is closely related to E. arostratus
Kornicker but may be differentiated from that species by the shallower
rostral incisure with a lateral shield, the subterminal location of
bristles on the 2nd joint of the endopodite of the 2nd antenna of
the male, and the lower number of bristles on the anteroventral part
of the inner lamella of the carapace: 5 compared to 16-21 on EL.
arostratus.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121
Euphilomedes arostratus Kornicker, 1967

FIGuRE 6

Euphilomedes arostrata Kornicker, 1967, p. 2.

Houoryer.—USNM 112658. Sex: female. Male Atoll, Maldive
Islands, Indian Ocean.

Locauity.—Gulf of Suez, Red Sea. Specimen probably from Sta-
tion 90 but could be from Station 89. Data concerning the stations
is given under ‘““Type-locality” of EL. polae. This specimen was in
vial containing specimens identified as Philomedes polae by Dr.
Herbert Graf.

DESCRIPTION OF MALE.—Shell (figs. 6a—e, h): oval, elongate with
greatest height near middle, prominent rostrum and antennal sinus
(figs. 6a, b, e); anterior margin of rostrum and anteroventral margin of
shell with scalloped outline formed by crescent-like marginal denticu-
lations. Surface of valves with distinct irregular polygons (fig. 6h);
polygons with pebbly texture appearing as minute polygons under
high magnification. Posterior dorsal margin with linear hinge de-
pressed below outline of shell; posterior hinge element of each valve
consisting of angular sclerotized process; medial hinge element
straight; anterior hinge element not prominent. Left valve broadly
overlapping right valve along anterodorsal margin. Numerous
adductor muscle scars (partly obscured by muscles in specimen exam-
ined) clustered behind and below middle of valve; 2 small scars and
large oval hyaline spot situated some distance above other; additional
scars below anterior hinge element (fig. 64). Inner lamella broad with
vestibule; selvage with wide, corrugated lamellar prolongation having
fringe of slender spines along outer margin. Eleven long hairs with
a few secondary spines forming row on inner lamella behind rostrum
(fig. 6b); inner lamella below rostrum with small hair followed by wide
space and then 20 hairs on right valve and 21 on left valve (figs. 6c, d);
about 34 hairs on posteroventral part of inner lamella. Hair with
either pointed or blunt ends distributed on rostral surface, some form-
ing row near margin of shell; slender hairs with pointed ends on
posterior surface of shell, some forming row near posterior margin; a
long tapered hair projecting from base immediately behind posterior
hinge element of each valve.

Dimensions: Greatest length 1.23 mm, greatest height 0.73 mm.
Specimen is larger than male reported by Kornicker (1967) from
the Maldives, length 1.00 mm, height 0.60 mm.

First antenna: First joint with medial hairs. Second joint with
medial hairs and, distally, a dorsal, ventral, and lateral bristle, all
bristles bare or with short hairs distally. Third joint with few medial
hairs and 1 ventral and 2 dorsal bristles, all with short hairs, the

NO. 3571 RED SEA OSTRACODS—KORNICKER 15

Ficure 6.—£Euphilomedes arostratus, male, specimen A: a, lateral view of complete carapace
showing position of lateral eye and some reticulations; b, medial view rostrum left valve;
c, medial view of anteroventral margin of left valve; d, medial view of anteroventral of
margin of right valve; e, medial view of antennal sinus area of left valve; f, endopodite of
left 2nd antenna; g, lateral view protopodite, endopodite, and Ist 3 joints of exopodite of
2nd antenna; A, medial view of reticulations, normal pores, and muscle scars (cross-
hatched) located dorsal to adductor muscle scars on left valve, anterior to right, upper
scars are located below anterior hinge element; 7, medial view right 6th limb (surface
hairs not shown); 7, distal part of 7th limb; &, distal tip of 7th limb; /, detail of distal end
of bristle of 7th limb; m, lateral view left caudal lamella. (Same scale, in microns:
b,e,f,h-7; ¢,d,g,m; k,l.)

16 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

longer dorsal bristle also provided with wreaths of long hairs near
middle. Fourth joint with 2 dorsal bristles, each with wreaths of
long hairs, and 4 long ventral bristles subdistally; bristles bare or with
short hairs distally. Fifth joint, inferred to be inserted ventrally
between 4th and 6th joints, bearing sensory bristle with broad base
and numerous filaments. Sixth joint with single dorsal bristle
distally having short spines. End joint with 1 short dorsal bristle
with short spines, 2 slender bristles with filaments, 2 bristles with
filaments, and usual long, stout c- and f-bristles.

Second antenna (figs. 6f,g): Protopodite triangular without trans-
parent rim (fig. 6g). Exopodite: first jomt elongate; 2nd joint
about % length of 1st; 3rd joint about % length of 1st; distal margins
of joints 2-8 with comb of short spines; basal spines not observed.
First joint without bristle; bristle of 2nd joint about % length of 3rd
joint (fig. 6g), bearing short marginal spines. Bristles of joints 3-8
bearing natatory hairs, without marginal spines. End joint with 4
stout bristles with natatory hairs and 2 short slender bristles without
hairs. Endopodite (fig. 6f) 3-jointed; 1st joint with 5 bare short
bristles and 1 long distal bristle with wreaths of long hairs near
middle and short spines distally; 2nd joint elongate, with 2 bristles near
middle, bristles provided with short spines; 3rd elongate with 1
proximal and 1 distal short bristle, tip with serrated ridges (tip of
appendage partly obscured with the result that presence or absence
of spine at tip could not be ascertained).

Mandible: No coxale endite observed. Basale: ventral margin with
4 short slender bristles and 2 long bristles with wreaths of long hairs
and short spines distally. Dorsal margin with 3 bristles. Medial
surface with about 5 short bristles proximally near ventral corner and
1 longer bristle near ventral margin at middle of joint. Exopodite
reaching middle of 1st endopodite joint, with 2 subequal terminal
bristles; tip of joint with blunt hirsute process.. Endopodite: first
joint with 1 short and 3 long ventral bristles distally. Second joint of
anterior margin with proximal group of 3 and distal group of 5 bristles;
posterior margin with 2 distal spinous bristles and 1 short annulated
bristle and 2 clawlike subterminal bristles. End joint with 2 large
subequal claws, 1 short posterior claw, and 3 annulated bristles. Medial
surface of basale and 1st and 2nd joints of endopodite provided with
groups of hairs.

Maxilla: Very small. It seems to have a structure typical of the
genus, but because of its small size I have not attempted to describe
it fully.

NO. 3671 RED SEA OSTRACODS—KORNICKER 17

Fifth limb: Epipodial appendage with 45 plumose bristles. Outer
lobe of 3rd exopodite joint with 2 stout plumose bristles.

Sixth limb (fig. 67): First endite with 3 bristles, 2nd endite with 4
bristles, 3rd endite with 7 bristles; 2nd joint of exopodite with 14 or
15 bristles, joint not produced posteriorly; epipodial appendage repre-
sented by 2 short bare bristles. Surface of 2nd joint of exopodite with
clusters of short hairs (not shown on illustration).

Seventh limb (figs. 67-/): Cleaning bristles: 4 in distal group, each
with 2 to 4 bells; 3 proximal bristles, each with 1 to 3 bells; some
bristles with short marginal spines. Terminal comb with about 7
marginal teeth, some with spines. Two long curved pegs opposing
comb.

Copulatory organ: Long slender, divided into 3 lobes with bristles.

Furca (fig. 6m): Each lamella with 11 claws: primary claws number
1, 2, 6, 10; secondary claws 3, 4, 5, 7, 8,9, 11. Primary claws 1 and 2
and all secondary claws separated from lamella; all claws with double
row of spines. Primary claws decrease in length proximally on lamella.
Lamella near claws 1 and 2 with clusters of long hairs.

Frontal organ: Elongate, 2-jomted with short spines on surface of
end joint and distal end of basal joint.

Remarks.—Although I have identified the Red Sea specimen as
Euphilomedes arostratus, some differences between this form and that
described from the Maldive Islands (Kornicker, 1967) creates
doubt concerning their conspecificity. A difference of possible
significance is that the Red Sea form has 2 bristles in place of the
epipodial appendage on the 6th limb as contrasted to 3 in this position
on the male of F. arostratus described from the Maldives. The female
of FE. arostratus from the Maldives, however, has only 2 bristles,
suggesting that the species may have either 2 or 3 epipodial bristles.
Also, whereas the male of EZ. arostratus from the Maldives has 9 to 10
bristles on the inner lamella behind the rostrum, the Red Sea form
has 11, the same number as the female from the Maldives. This
suggests that the number of bristles in this position might vary from
9 to 11 for the species. The Red Sea specimen is also larger than the
Maldive form. In most characters, the Red Sea and Maldive Island
forms are so similar that it seems best at this time to include them in
the same species.

Literature Cited
Barrp, W.
1850. The natural history of the British Entomostraca, 364 pp., 36 pls.
[Ostracoda: pp. 138-182, pls. 18-23.]
Grar, H.
1931. Expedition S.M.S. “Pola” in das Rote Meer: Die Cypridinidae des
Roten Meeres. Denkschr. Akad. Wiss. Wiener Math. Naturw.
Klasse, vol. 102, pp. 32-46, figs. 1-10.
Kornicker, L. S.
1967. Euphilomedes arostrata, a new myodocopid ostracod from the Maldive
Islands, Indian Ocean. Proc. U.S. Nat. Mus., vol. 120, no. 3563,
pp. 1-21.
Mi.irr, G. W.
1912. Ostracoda. Pt. 31 in Das Tierreich, 434 pp., 92 figs.
Poutsen, E. M.
1962. Cypridiniformes-cypridinidae. Pt. 1 i Ostracoda-Myodocopa, 414
pp. [Dana-Report No. 57.]

18

U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3571

SUPPLEMENTARY DESCRIPTIONS
OF TWO MYODOCOPID OSTRACODS
FROM THE RED SEA

By Louts S. KornickER
Associate Curator, Division of Crustacea

This study is based on ostracods collected on the Pola Expedition
to the Red Sea in 1896 and described by Dr. Herbert Graf (1931). In
the original description of Philomedes polae (=Euphilomedes polae),
Graf (1931) did not designate a holotype and omitted some details of
carapace and appendage morphology. Graf’s description may have
been based on more than one species. The species, therefore, is

_ redescribed herein and a lectotype selected from the available type-
series. Specimens designated as Philomedes species by Graf (1931)
_ have been identified as females of E. polae and are described as such.
_ A single specimen included by Graf in the type-series of Philomedes
polae has been identified as Euphilomedes arostrata Kornicker
(1967); this identification is documented with a description and
illustrations of the Red Sea specimen.

The following material was obtained through the courtesy of Dr.
Gerhard Pretzmann from the Naturhistorisches Museum, Vienna,
Austria: (1) One bottle, containing one vial with 3 males, with red
label marked: “‘Philomedes polae, n. sp. &; Rotes Meer: Gulf von Suez;

if!

a PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Coll: ‘Pola’ 1895/8; Dr. Graf det.” (2) One bottle with white label
marked: “Philomedes sp. 9; Rotes Meer: Stat. 90; Coll. ‘Pola’ 1895/8;
Dr. Graf det.” This bottle contained one vial with 2 females con-
taining eggs that had not yet been extruded into the brood chamber
and with a label that had been marked ‘“‘Philomedes sp. 9 (A).”

Since Dr. Graf had not designated a holotype of Philomedes polae
Graf, Dr. Pretzmann suggested that I select a lectotype from the type
material. Examining the material, I found 2 specimens in the first
bottle that I could equate with Philomedes polae Graf, using primarily
as a basis for identification the distinctive secondary appendage
described and figured for this species by Graf (1931, p. 37, fig. 5i).
The third specimen in the first bottle I identify as Huphalomedes
arostratus Kornicker, recently described for the first time from the
Maldive Islands, Indian Ocean.

The 2 specimens in the second bottle I equate to specimens
described by Graf (1931) as Philomedes species on his page 38 and
illustrated in his figure 6. The similarity of the distribution of
primary and secondary claws on the caudal rami of the specimens at
hand with the published description of Philomedes species makes it
certain that this identification is correct. After studying these speci-
mens, I have concluded that they are females of Philomedes polae Graf.

I acknowledge with thanks the assistance of Dr. Gerhard Pretzmann
in obtaining specimens for study from the Naturhistorisches Museum,
Vienna, Austria, and of Miss Caroline Bartlett, who inked the final
illustrations. I would like also to thank Mr. I. G. Sohn and Doctors
Raymond B. Manning and W. Duane Hope for reviewing the manu-
script.

Family Cypridinidae Baird, 1850
Subfamily Philomedinae G. W. Miiller, 1912

Genus Euphilomedes Poulsen, 1962

Typr-sprctus.—Huphilomedes nodosus Poulsen, 1962, by subsequent
designation, Kornicker (1967).

Euphilomedes polae (Graf, 1931)
Fiaures 1-5

Philomedes polae Graf, 1931, p. 37, fig. 5.
Philomedes species.—Graf, 1931, p. 38, fig. 6.
LrecroryPE.—Specimen designated as specimen B on slide. Natur-
historisches Museum, Zoologische Abteilung, Vienna, Austria.
Spx.—Male.

NO. 3571 RED SEA OSTRACODS—KORNICKER 3

Ficure 1.—Euphilomedes polae, male, specimen C (except a and h, spec. B): a, lateral view
left valve showing some muscle scars, surface reticulation shown only near posteroventral
margin of valve; b, lateral view anterior end of left valve showing surface and marginal
hairs; c, lateral view anterior end of left valve showing flap over antennal sinus, and linear
sclerotized area visible in transmitted light; d, lateral view posterior end of left valve
showing surface hairs and some reticulations; ¢, medial view of part of shell posterior
showing reticulations, normal pore canals and surface hairs; f, lateral view anterior end
of right valve showing marginal denticulations around rostrum, flap over antennal sinus
and linear sclerotized area; g, medial view anterior end of right valve; h, medial view
anterior end of left valve; i, medial view of middle section of right valve showing adductor
muscle scars below and dorsal muscle scars above; 7, medial view of dorsal muscle scars
below anterior hinge element and oval lucid area below muscle scars; k, medial view right
valve showing minute reticulations within large reticulations on surface of valve. (Same
scale, in microns: b,c,d,f,i, e,g,h,j.)

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

PARALECTOTYPE.—1 male designated as specimen C, Naturhisto-
risches Museum.

ADDITIONAL SPECIMENS.—2 females from the type-locality desig-
nated as specimens 1 and 2 on slides, Naturhistorisches Museum.

TypE-LocALiTy.—Graf (1931) reported 4 specimens of Philomedes
polae from Pola Expedition Station 90 and 1 specimen from Station 89;
both stations are in the Gulf of Suez; all specimens are males. The
vial containing specimens of P. polae did not bear a station number;
however, as 3 specimens were in the vial, it is probable that all, and
at least 2 of them, are from Station 90 rather than 89. Station 90,
therefore, is considered here to be the type-locality; since Stations 89
and 90 are close to each other, however, whether the lectotype was
collected at one or the other is not of major significance. Females of
P. polae, designated by Graf as Philomedes species, were collected at
Station 90. Data concerning the stations are as follows:

station latitude longitude collecting depth date collected
89 28°40’N 32°57'E surface March 31, 1896
90 28°00’N 33°36/E surface April 1, 1896

DESCRIPTION OF MALE.—Shell (figs. 1, 2a-e): oval, elongate with
ereatest height near middle, prominent rostrum and shallow rostral
incisure (fig. la); anterior margin of rostrum and anteroventral
margin of shell with scalloped outline formed by cresent-like marginal
denticulations; alate shield projecting laterally and anteroventrally
from behind rostral incisure, partly covering incisure, flap strongly
sclerotized anteriorly (figs. 1¢,f). Surface of valves with irregular
polygons; polygons with pebbly texture appearing as minute polygons
under high magnification (figs. 1e,/). Posterior dorsal margin with
linear hinge depressed below outline of shell (figs. 2b,c); hinge not
visible in lateral view except by transmitted light. Posterior hinge
clement of each valve consisting of angular sclerotized process (figs.
2c,d); medial hinge element straight; anterior hinge element not
prominent. Left valve broadly overlapping right valve along antero-
dorsal margin. About 20 individual muscle scars (partly obscured
by muscles in specimen examined) clustered behind and below middle
of valve; small scar and large oval hyaline spot situated some distance
above others; 2 large and 4 smaller scars situated below anterior hinge
element (figs. 1a,i,j). Inner lamella broad with vestibule; no parallel
striations or line of concrescence observed on inner lamella. Selvage
with wide, corrugated, lamellar prolongation having fringe of slender
spines along outer margin. Six long hairs bearing secondary spines,
forming row on inner lamella behind rostrum (figs. 1g,h); inner lamella
ventral to rostral incisure with small plumose hair, followed by wide
space and then about 5 plumose hairs (fig. 2a); about 30 hairs on
posteroventral part of inner lamella (figs. 2d,e).

NO. 3571 RED SEA OSTRACODS—KORNICKER a

Figure 2.—Euphilomedes polae, male, specimen B (except c, spec. C): a, medial view of
anteroventral shell margin of left valve showing medial hairs; b, medial view of left valve
hinge area (anterior on right); c, medial view of posterior hinge element on right valve;
pustules along middle hinge element are dotted because they could not be seen with
certainty on specimen C and were not observed on specimen B; d, medial view marginal
area at posterior end of left valve showing medial hairs; ¢, medial view posteroventral and
posterodorsal margin showing medial hairs; f, lateral view of right Ist antenna (only
proximal part of c- and f-bristles shown); g, exopodite of 2nd antenna; h, distal end 3rd
joint of endopodite of 2nd antenna; 7, protopodite and endopodite of 2nd antenna; 7, medial
view Ist endopodite joint of left mandible, distal bristles are on ventral margin (joint
foreshortened by oblique position of joint in mounting medium); k, medial view 2nd
endopodite joint of left mandible, dorsal margin to right; /, lateral view end joint of right
mandible, all bristles not shown. (Same scale, in microns: a,b,e,f,g,i; c,d,j,k; h,l.)

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Hairs with either pointed or blunt ends distributed on rostral surface
(fig. 1b), some forming row near margin of shell; slender hairs with
pointed ends on posterior surface of shell, some forming row near pos-
terior margin (fig. 1d); a long tapered hair projecting from base
immediately behind posterior hinge element of each valve (figs.
2c-e); all hairs coming through normal pore canals; radial pore
canals not observed.

Dimensions: Lectotype (specimen B) greatest length 1.12 mm,
greatest height 0.63 mm; paralectotype (specimen C) greatest length
1.12 mm, greatest height 0.63 mm. Graf (1931, p. 38) gives length
of this species as 1.05 mm and height 0.60 mm.

First antenna (fig. 2f): First joint with faint clusters of short
hairs on medial surface. Second joint with clusters of long hairs on
medial surface, and distally, a dorsal, ventral, and lateral bristle;
all bristles bare or with short hairs, none with wreaths of long hairs.
Third joint with a few clusters of short hairs on medial surface, and 1
ventral and 2 dorsal bristles, the longer dorsal bristle provided with
wreaths of long hairs near middle and short hairs distally;
other dorsal bristle and ventral bristle with short hairs. Fourth
joint with 2 dorsal bristles, each with wreaths of long hairs and the
more proximal bristle with short hairs distally, and 4 long ventral
bristles subterminally, latter bristles bare or with short hairs distally.
Fifth joint, inferred to be inserted ventrally between 4th and 6th
joints, bearing sensory bristle with broad base and provided with
numerous filaments. Sixth joint with single bristle on 1 limb
and without bristle on other limb (bristle may have been broken off
during dissection). End joints with 5 slender bristles and 2 long
stout c- and f-bristles.

Second antenna (figs. 2g-i): Protopodite triangular with trans-
parent rim (fig. 27). Exopodite (fig. 2g): 1st joint elongate; 2nd
joint about ¥% length of 1st; 3rd joint about same length as 1st; distal
margins of joints 2-8 with comb of short spines (not shown on fig. 2g);
basal spines not observed. Bristle of 2nd joint about % length of
3rd joint, bearing short marginal spines. Bristles of joints 3-8
bearing natatory hairs, without marginal spines. End joint with 4
stout bristles with natatory hairs and 2 short slender bristles. Endop-
odite 3-jointed (figs. 24,7); 1st joint with 5 bare short bristles and
1 long distal bristle with wreaths of long hairs near middle and short
spines distally; 2nd joint elongate, with 2 subterminal bristles pro-
vided with short spines; 3rd joint elongate, broad proximally, with
1 short proximal bristle and 2 longer subterminal bristles; tip with
short spine and about 5 serrate ridges.

Mandible (figs. 2j-1, 3a,b): No coxale endite observed. Basale
(fig. 3b): ventral margin with 3 short slender spinous bristles, and 2

aaa ae

NO. 3571 RED SEA OSTRACODS—KORNICKER

\\ WW \y ( <
ccf Way Qt
WL, \ \ NSP nase

Ww Ye v Yo Im,

Wenn

b, f, 1, m; ¢, g3 d, 7, k.)

Ficure 3.—Euphilomedes polae, male, specimen B: a, medial view of exopodite and Ist
endopodite joint without distal bristles of left mandible; 5, lateral view of basale and
exopodite of right mandible; c, maxilla, all bristles not shown; d, 5th limb, hairs not
shown on all epipodial bristles; ¢, endites and exopodite of 5th limb, all bristles not shown;
f, 6th limb; g, 7th limb (1 bristle in distal group broken); h, distal end of 7th limb; 7, frontal
organ and median eye; j, lateral view of right furcal lamella; k, copulatory organ; /, detail
of distal tip of copulatory organ; m, lateral eye. (Same scale, in microns: a, e, h,1;

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

long bristles with wreaths of long hairs and short spines distally.
Dorsal margin with 3 bristles, 1 near middle and 2 subterminally.
Medial surface with about 5 short bristles proximally near ventral
corner and 1 longer bristle near ventral margin at middle of joint.
Exopodite reaching middle of 1st endopodite joint, with 2 subequal
terminal bristles with short marginal spines (fig. 3a); tip of joint with
blunt hirsute process bearing short spine. Endopodite: First joint
(fig. 27) with 1 short spinous bristle and 3 long hirsute posterior
bristles distally. Second joint (fig. 24) anterior margin with proximal
eroup of 3 bristles and distal group of 5 bristles, all bristles bare or
with short spines; posterior margin with 2 spinous bristles distal and 1
short annulated bristle and 2 clawlike bristles subterminally. End
joint (fig. 27) with 2 large subequal claws, 1 short anterior claw, and 3
annulated bristles. Medial surface of basale and 1st and 2nd joints
of endopodite provided with groups of hairs.

Maxilla (fig. 3c): Very small. It seems to have a structure typical
of the genus, but because of its size I have not attempted to describe
it fully.

Fifth limb (figs. 3d, e): Epipodial appendage with about 47 plumose
bristles. Outer lobe of 3rd exopodite joint with 2 stout plumose
bristles.

Sixth limb (fig. 3f): First endite with 3 bristles, 2nd endite with
4 bristles, 3rd endite with 6 bristles, 4th endite with 6 bristles, 2nd
joint of exopodite with 14 bristles, joint not produced posteriorly;
epipodial appendage represented by 2 short bare bristles. Surface of
second joint of exopodite with clusters of short hairs.

Seventh limb (figs. 3g, 4): Cleaning bristles: 4 in distal group, each
with 4 or 5 bells; 3 in proximal group, each with 1 to 3 bells; surface
hairs not observed on bristles. Terminal comb with about 7 marginal
teeth, some with marginal spines. Two long curved pegs opposing
comb.

Copulatory organ (figs. 3k, 2): Long, slender, divided into 3 lobes,
each with 2 annulated bristles; 1 lobe with large curved sclerotized tip.

Furea (fig. 37): Each lamella with 12 to 13 claws: primary claws
number 1, 2, 6, 10; secondary claws 3, 4, 5, 7, 8, 9, 11, 12, 13 (13th claw
not always present). Primary claws 1 and 2 separated from lamella,
6 and 10 joined to lamella; secondary claws all separated from lamella;
all claws with double row of spines. Primary claws decrease in length
proximally on lamella. Lamella near claws 1 and 2 with clusters of
long hairs. (One specimen has 13 claws on each lamella, another
specimen has 13 claws on the right lamella and only 12 on the left
lamella.)

NO. 8571 RED SEA OSTRACODS—KORNICKER 9

Ficure 4.—Euphilomedes polae, female, specimen 2 (except a-d, j-l, spec. 1): a, lateral
view left valve, dashed area near valve middle contains adductor muscle scars, not all
surface polygons shown; 5, medial view of anterior end of left valve, sclerotized areas
stippled; c, medial view anterior of right valve, sclerotized areas stippled; d, lateral view
of anterior end of left valve showing surface and marginal hairs; e, medial view of antero-
ventral margin of right valve showing distribution of hairs on inner lamella; f, medial
view of valve near posterior showing large and small reticulations; g, medial view hinge
element of right valve (anterior end on right); h, medial view of muscle scars on right
valves, adductor muscle scars clustered at lower left of figure, anterior of valve to left;
7, medial view of posterior and ventral marginal area of right valve showing distribution
of hairs on inner lamella; 7, lateral view of left Ist antenna; &, lateral view of endopodite,
first 3 joints of exopodite and distal margin of protopodite of left 2nd antenna; /, medial
view of right 2nd antenna, ends of some bristles of exopodite not shown. (Same scale,
in microns: b—d, f, k; e, g-J, l.)

221—535—66——2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Eyes: Lateral eyes large; about 19 ommatophores visible in side
view, all except 2 divided by a suture into 2 parts (fig. 3m). Medial
eye large, pigmented (fig. 37).

Frontal organ (fig. 37): Elongate, 2-jointed with short spines on
distal half of 2nd joint and subterminally on basal joint.

DESCRIPTION OF FEMALE.—Shell (figs. 4a-7): lateral outline more
ovoid than male (fig. 4a); hinge structure (fig. 4g) selvage similar to
male; adductor muscle scars (figs. 4a,h) more anteriorly and ventrally
located on shell than on male; rostrum narrower and incisure shallower
than on male; irregular polygons on surface of shell subdivided by
minute polygons larger than on male shell and visible under low
magnification (fig. 4f). Inner lamella with parallel striations on
anteroventral part; 9 long hairs with secondary spines forming row
on inner lamella behind rostrum (figs. 4b,c); inner lamella below rostral
incisure with small hair followed by wide space and then 5 hairs (fig.
4e); posteroventral part of inner lamella with at least 21 hairs (fig. 47);
lateral surface of rostrum and anterior part of shell with numerous
hairs with blunt ends (fig. 4d).

Dimensions: Specimen no. 1 greatest length 1.06 mm, greatest
height 0.86 mm; specimen no. 2 greatest length 1.03 mm, greatest
height 0.73 mm. Graf (1931, p. 38) gives length as 0.90 mm and
1.00 mm and height as 0.70 mm.

First antenna (fig. 47): First and 2nd joints with surface hairs.
Second joint with 3 bristles, 1 dorsal, 1 ventral, and 1 lateral. Third
joint with 1 ventral bristle distally and 1 short bare bristle and 1 long
hirsute bristle on dorsal margin; ventral margin with short spines.
Fourth joint with 2 dorsal subterminal bristles and 1 short and 3 long
ventral subterminal bristles; ventral margin with short spines. Fifth
joint with stout ventral terminal bristle with filaments. Sixth joint
with medial bristle distally. End joints with 1 short bristle and 2
long bristles without filaments and 4 long bristles with filaments.

Second antenna (figs. 44,2): Protopodite subtriangular without
transparent rim. Exopodite (fig.4/) with 9 joints; 1st joint elongate
with small medial spine; 2nd to 9th joints trapezoidal without basal
spines; distal margins of 2nd to 5th joints with comb of slender spines;
2nd and 3rd joints with bristles without natatory hairs or marginal
denticulations; 4th to 8th joints with bristles having natatory hairs
and marginal denticulations; 9th joint with 2 short bare bristles and
2 subequal long bristles with natatory hairs and marginal denticula-
tions. Endopodite (figs. 4/,/) with 2 joints; basal joint with long stout
bristle with wreath of long hairs near middle, and 3 to 5 short bare
bristles; distal joint elongate with small spine at tip and long stout
terminal bristle with wreaths of long hairs near middle and short
distal spines.

NO. 3571 RED SEA OSTRACODS—KORNICKER 11

Figure 5.—Euphilomedes polae, female, specimen 1: a, medial view right mandible (surface
hairs not shown); d, lateral view left maxilla, all bristles not shown; ¢, posterior view left
5th limb; d, medial view right 6th limb; e, 7th limb; f, distal end of 7th limb; g, distal end
frontal organ; h, frontal organ and medial eye; 7, medial view right lamella of furca;
J, lateral view left lamella of furca. (Same scale, in microns: b,d,e,h-j; c,f,g.)

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Mandible (fig. 5a): Coxale endite large, bifurcate, with rows of
spines. Basale: ventral margin with 4 short bristles followed by 1
longer bristle near distal end; dorsal margin with 3 bristles, 1 near
middle and 2 at distal end; medial surface with 1 bristle at middle
and 5 shorter bristles proximally, all near ventral margin. Exopodite
short with 2 bristles and triangular process at tip, mner bristle with
wreath of long hairs near middle. Endopodite: distal end of 1st joint
with 1 short and 3 long bristles ventrally; anterior margin of 2nd joint
with proximal group of 4 and distal group of 5 bristles; posterior margin
with 2 subterminal bristles and terminally with 1 short annulate
bristle and 2 clawlike bristles; distal joint with 2 long stout claws of
subequal length and 3 bristles. Surface of basale and 2nd joint of
exopodite hirsute.

Maxilla (fig. 5b): Precoxa and coxa with fringe of fine hair along
dorsal margin; coxa with short bare bristle on dorsal margin; basale
with 3 distal bristles; Ist endopodite joint with marginal hairs, a
hirsute bristle terminal on anterior margin and about 5 bristles on
distal margin; distal end of endopodite joint provided with numerous
bristles; exopodite with 2 long and 1 short bristle. Three endites: 1st
endite with about 8 bristles; 2nd endite with about 4 bristles; 3rd
endite with about 6 distal and 1 proximal bristle (bristles not shown
in illustration).

Fifth limb (fig. 5c): Epipodial appendage with about 49 bristles.
Triangular tooth anterior to main tooth of Ist exopodite joint about
same size as distal tip of 2nd exopodite joint. Distal anterior margin
of Ist joint with 2 centrally located bristles. Large curved tooth
of 2nd exopodite joint without bristle at distal lateral corner.

Sixth limb (fig. 5d): First endite with 3 bristles; 2nd endite with
4 bristles; 3rd endite with 7 bristles; 4th endite with 6 bristles;
epipodial appendage represented by 2 short bristles. Second joint
of exopodite not produced posteriorly, distal margin with 13 to 14
bristles, medial and lateral surfaces hirsute.

Seventh limb (figs. 5e,f): Five cleaning bristles in distal group,
3 in proximal group, each bristle with 1 to 5 terminal bells. Terminal
comb with about 9 teeth, secondary teeth at base (all teeth not shown
in illustration); 2 elongate pegs opposing comb.

Furea (figs. 57,7): Each lamella with 4 primary claws and 10 to 12
secondary claws. Primary claws 1 and 2 separated from lamella,
7 and 12 (or 11) joined to lamella. Primary claws decrease in length
proximally on lamella. Secondary claws 3 to 6 about same length,
shorter than claw 7, longer than claw 8; claws 8 to 12 (or 11) decrease
in size proximally. Primary claw 12 (or 11) about same size or longer
than preceding secondary claw and longer than following secondary

NO. 3571 RED SEA OSTRACODS—KORNICKER 13

claw. Lamella near basis of claws with long hairs. Primary claws
1 and 2 with double row of teeth, claws 7 and 12 without teeth;
secondary claws with marginal spines. Distribution of primary and
secondary furcal claws on 2 specimens is as follows:

claw number

DS AH! OD OB PEGG.” GN ah Seatelge te Sip tie
specimen 1
right Dap Be Py tSise SS Ais eepey Assia ns
left Dp 8 Di ACSale 8) Sis pay Suis rs
specimen 2
right Dep Bie TS Se <P eh. a8 ess 8) ele Aa eae ec
left Deep B28) 8) 0 De) Sk 8h) 28-5), Spies

Kyes: Large medial eye (fig. 5h); lateral eyes not observed.

Frontal organ (figs. 5g,h): 2-jointed with distal joint 3 times length
of proximal joint; tip rounded; surface of 2nd joint and distal end of
1st joint with fine spines.

Remarxs.—Extreme sexual dimorphism among some species of
ostracods creates inherent difficulties in relating males and females
with certainty. Carapace morphology in the region of the rostrum
and rostral incisure between males and females of Euphilomedes polae
identified herein is sufficiently similar to warrant considering them as
conspecific in spite of some differences in appendage morphology.
Both males and females were collected in the same area. The male
and female of E. arostratus Kornicker, a closely related species, have
similar dimorphism.

Some differences in morphology between males and females of EF.
polae and E. arostratus are compared as follows:

E. polae E. arostratus
male female male female
number of secondary furcal claws 8-9 10-12 7 9
cleaning bristles in distal group of 7th limb 4 5 4 5
number of hairs in row on inner lamella
behind rostrum 6 9 9-11* 11

*The male of E. arostratus described from the Maldive Islands (Kornicker,
1967) has 9 to 10 hairs, whereas the Red Sea form described in this paper has 11.

Comparisons.—Euphilomedes polae is closely related to E. arostratus
Kornicker but may be differentiated from that species by the shallower
rostral incisure with a lateral shield, the subterminal location of
bristles on the 2nd joint of the endopodite of the 2nd antenna of
the male, and the lower number of bristles on the anteroventral part
of the inner lamella of the carapace: 5 compared to 16-21 on LE.
arostratus.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121
Euphilomedes arostratus Kornicker, 1967
FIGURE 6

Euphilomedes arostrata Kornicker, 1967, p. 2.

Hotoryer.—USNM 112658. Sex: female. Male Atoll, Maldive
Islands, Indian Ocean.

Locauiry.—Gulf of Suez, Red Sea. Specimen probably from Sta-
tion 90 but could be from Station 89. Data concerning the stations
is given under “Type-locality” of E. polae. This specimen was in
vial containing specimens identified as Philomedes polae by Dr.
Herbert Graf.

DESCRIPTION OF MALE.—Shell (figs. 6a-e, h): oval, elongate with
ereatest height near middle, prominent rostrum and antennal sinus
(figs. 6a, 6, e); anterior margin of rostrum and anteroventral margin of
shell with scalloped outline formed by crescent-like marginal denticu-
lations. Surface of valves with distinct irregular polygons (fig. 61);
polygons with pebbly texture appearing as minute polygons under
high magnification. Posterior dorsal margin with linear hinge de-
pressed below outline of shell; posterior hinge element of each valve
consisting of angular sclerotized process; medial hinge element
straight; anterior hinge element not prominent. Left valve broadly
overlapping right valve along anterodorsal margin. Numerous
adductor muscle scars (partly obscured by muscles in specimen exam-
ined) clustered behind and below middle of valve; 2 small scars and
large oval hyaline spot situated some distance above other; additional
scars below anterior hinge element (fig. 64). Inner lamella broad with
vestibule; selvage with wide, corrugated lamellar prolongation having
fringe of slender spines along outer margin. Eleven long hairs with
a few secondary spines forming row on inner lamella behind rostrum
(fig. 6b); inner lamella below rostrum with small hair followed by wide
space and then 20 hairs on right valve and 21 on left valve (figs. 6c, d);
about 34 hairs on posteroventral part of inner lamella. Hair with
either pointed or blunt ends distributed on rostral surface, some form-
ing row near margin of shell; slender hairs with pointed ends on
posterior surface of shell, some forming row near posterior margin; a
long tapered hair projecting from base immediately behind posterior
hinge element of each valve.

Dimensions: Greatest length 1.23 mm, greatest height 0.73 mm.
Specimen is larger than male reported by Kornicker (1967) from
the Maldives, length 1.00 mm, height 0.60 mm.

First antenna: First joint with medial hairs. Second joint with
medial hairs and, distally, a dorsal, ventral, and lateral bristle, all
bristles bare or with short hairs distally. Third joint with few medial
hairs and 1 ventral and 2 dorsal bristles, all with short hairs, the

NO. 3571 RED SEA OSTRACODS—KORNICKER 15

Ficure 6.—Euphilomedes arostratus, male, specimen A: a, lateral view of complete carapace
showing position of lateral eye and some reticulations; b, medial view rostrum left valve;
¢, medial view of anteroventral margin of left valve; d, medial view of anteroventral of
margin of right valve; ¢, medial view of antennal sinus area of left valve; f, endopodite of
left 2nd antenna; g, lateral view protopodite, endopodite, and Ist 3 joints of exopodite of
2nd antenna; h, medial view of reticulations, normal pores, and muscle scars (cross-
hatched) located dorsal to adductor muscle scars on left valve, anterior to right, upper
sears are located below anterior hinge element; 7, medial view right 6th limb (surface
hairs not shown); 7, distal part of 7th limb; &, distal tip of 7th limb; /, detail of distal end
of bristle of 7th limb; m, lateral view left caudal lamella. (Same scale, in microns:
b,e,f,h-z; ¢,d,g,m; k,l.)

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

longer dorsal bristle also provided with wreaths of long hairs near
middle. Fourth joint with 2 dorsal bristles, each with wreaths of
long hairs, and 4 long ventral bristles subdistally ; bristles bare or with
short hairs distally. Fifth joint, inferred to be inserted ventrally
between 4th and 6th joints, bearing sensory bristle with broad base
and numerous filaments. Sixth joint with single dorsal bristle
distally having short spines. End joint with 1 short dorsal bristle
with short spines, 2 slender bristles with filaments, 2 bristles with
filaments, and usual long, stout c- and f-bristles.

Second antenna (figs. 6f,9): Protopodite triangular without trans-
parent rim (fig. 6g). Exopodite: first joint elongate; 2nd joint
about % length of 1st; 3rd joint about % length of 1st; distal margins
of joints 2-8 with comb of short spines; basal spines not observed.
First joint without bristle; bristle of 2nd joint about % length of 3rd
joint (fig. 6g), bearing short marginal spines. Bristles of joints 3-8
bearing natatory hairs, without marginal spines. End joint with 4
stout bristles with natatory hairs and 2 short slender bristles without
hairs. Endopodite (fig. 6f) 3-jointed; Ist joint with 5 bare short
bristles and 1 long distal bristle with wreaths of long hairs near
middle and short spines distally ; 2nd joint elongate, with 2 bristles near
middle, bristles provided with short spines; 3rd elongate with 1
proximal and 1 distal short bristle, tip with serrated ridges (tip of
appendage partly obscured with the result that presence or absence
of spine at tip could not be ascertained).

Mandible: No coxale endite observed. Basale: ventral margin with
4 short slender bristles and 2 long bristles with wreaths of long hairs
and short spines distally. Dorsal margin with 3 bristles. Medial
surface with about 5 short bristles proximally near ventral corner and
1 longer bristle near ventral margin at middle of joint. Exopodite
reaching middle of 1st endopodite joint, with 2 subequal terminal
bristles; tip of joint with blunt hirsute process. Endopodite: first
joint with 1 short and 3 long ventral bristles distally. Second joint of
anterior margin with proximal group of 3 and distal group of 5 bristles;
posterior margin with 2 distal spinous bristles and 1 short annulated
bristle and 2 clawlike subterminal bristles. End joint with 2 large
subequal claws, 1 short posterior claw, and 3 annulated bristles. Medial
surface of basale and 1st and 2nd joints of endopodite provided with
groups of hairs.

Maxilla: Very small. It seems to have a structure typical of the
genus, but because of its small size I have not attempted to describe
it fully.

NO. 3571 RED SEA OSTRACODS—KORNICKER 17

Fifth limb: Epipodial appendage with 45 plumose bristles. Outer
lobe of 3rd exopodite joint with 2 stout plumose bristles.

Sixth limb (fig. 67): First endite with 3 bristles, 2nd endite with 4
bristles, 3rd endite with 7 bristles; 2nd joint of exopodite with 14 or
15 bristles, joint not produced posteriorly; epipodial appendage repre-
sented by 2 short bare bristles. Surface of 2nd joint of exopodite with
clusters of short hairs (not shown on illustration).

Seventh limb (figs. 67-/): Cleaning bristles: 4 in distal group, each
with 2 to 4 bells; 3 proximal bristles, each with 1 to 3 bells; some
bristles with short marginal spines. Terminal comb with about 7
marginal teeth, some with spines. Two long curved pegs opposing
comb.

Copulatory organ: Long slender, divided into 3 lobes with bristles.

Furca (fig. 6m): Each lamella with 11 claws: primary claws number
1, 2, 6, 10; secondary claws 3, 4, 5, 7, 8,9, 11. Primary claws 1 and 2
and all secondary claws separated from lamella; all claws with double
row of spines. Primary claws decrease in length proximally on lamella.
Lamella near claws 1 and 2 with clusters of long hairs.

Frontal organ: Elongate, 2-jointed with short spines on surface of
end joint and distal end of basal joint.

Remarks.—Although I have identified the Red Sea specimen as
Euphilomedes arostratus, some differences between this form and that
described from the Maldive Islands (Kornicker, 1967) creates
doubt concerning their conspecificity. A difference of possible
significance is that the Red Sea form has 2 bristles in place of the
epipodial appendage on the 6th limb as contrasted to 3 in this position
on the male of /. arostratus described from the Maldives. The female
of E. arostratus from the Maldives, however, has only 2 bristles,
suggesting that the species may have either 2 or 3 epipodial bristles.
Also, whereas the male of E. arostratus from the Maldives has 9 to 10
bristles on the inner lamella behind the rostrum, the Red Sea form
has 11, the same number as the female from the Maldives. This
suggests that the number of bristles in this position might vary from
9 to 11 for the species. ‘The Red Sea specimen is also larger than the
Maldive form. In most characters, the Red Sea and Maldive Island
forms are so similar that it seems best at this time to include them in
the same species.

Literature Cited
Barrp, W.
1850. The natural history of the British Entomostraca, 364 pp., 36 pls.
[Ostracoda: pp. 138-182, pls. 18—23.]

Grar, H.
1931. Expedition §.M.S. “Pola” in das Rote Meer: Die Cypridinidae des
Roten Meeres. Denkschr. Akad. Wiss. Wiener Math. Naturw.
Klasse, vol. 102, pp. 32-46, figs. 1-10.
Kornickeir, L. S.
1967. Euphilomedes arostrata, 2 new myodocopid ostracod from the Maldive
Islands, Indian Ocean. Proc. U.S. Nat. Mus., vol. 120, no. 3563,
pp. 1-21.
Miuier, G. W.
1912. Ostracoda. Pt. 31 in Das Tierreich, 434 pp., 92 figs.
PouusEn, E. M.
1962. Cypridiniformes-cypridinidae. Pt. 1 in Ostracoda-Myodocopa, 414
pp. [Dana-Report No. 57.]

18

U.S. GOVERNMENT PRINTING OFFICE: 1967

————

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3572

CALIGOID COPEPODS PARASITIC
ON SHARKS OF THE INDIAN OCEAN

By Rocer F. Cressey
Associate Curator, Division of Crustacea

During the International Indian Ocean Expedition, 35 species of
caligoid copepods were collected from 29 species of sharks. The
author collected during Cruise 5 of the R.V. Anton Brwun and also at
Nosy Bé, Madagascar (Malagasy Republic). In addition, several
other participants collected copepods and donated the material to the
author for inclusion in this study.

Of the 35 species of copepods collected, 8 of them were new. Four
of these new species, all members of the Kudactylinidae, are described
here. Two of the others represented new genera (Pagina and Bariaka)
and each has been described separately elsewhere. The remaining 2
new species are members of the Pandaridae and are being described
in a paper currently in press revising the entire family.

A map of the Indian Ocean showing points of collection is included
(fig. 54). All station numbers refer to points on the various cruise
tracks of the R.V. Anton Bruun. In text tabulations, Roman numerals
refer to spines, Arabic to setae.

I would like to thank the following persons for their efforts on my
behalf: Dr. Richard Gooding, University of Singapore; Miss Sherril

al

2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Kite, Woods Hole Oceanographic Institution; Dr. Alan Lewis, Uni-
versity of British Columbia; Mr. Richard Shomura, Bureau of Com-
mercial Fisheries; Dr. Marta Vannucci, University of Sao Paulo; and
Miss Leonie Joubert, University of Natal, who sent a number of
collections from sharks caught near Durban, South Africa, that I have
included in this paper.

I would like to acknowledge the support of the National Science
Foundation-U.S. Program in Biology for the field portion of this work.

All material has been deposited in the U.S. National Museum.

Family Pandaridae Milne-Edwards, 1840

Pandarus cranchii Leach, 1819

Collections: From 22 specimens of Carcharinus longimanus (Poey)
at stations 108, 109, 113, 118, 139, 141, 144, 147, 289, 291, 293, 295,
297, 298, 323, 326, 411, 415, 417, and 418; from 1 specimen of the same
host off Durban, South Africa; from 4 specimens of Carcharinus
floridanus (Bigelow and Schroeder) at stations 111, 289, 293, and 326;
from 1 specimen of Alopias vulpinus at station 115; from 1 specimen of
Carcharodon carcharvas from Durban.

Habitat: Body surface and fins.

Pandarus smithii Rathbun, 1886

Collections: From 2 specimens of Alopias vulpinus (Bonneterre)
at stations 288 and 289; from 1 specimen of Jsurus oxyrinchus Rafines-
que at station 282; from 1 specimen of Carcharinus floridanus at
station 144.

Habitat: Body surface and fins.

Pandarus satyrus Dana, 1852

Collections: From 6 specimens of Prionace glauca (Linnaeus) at
stations 124, 292, 295, 306, 311, and 324.
Habitat: Body surface and fins.

Pandarus carcharini Ho, 1963

Collections: From 3 specimens of Carcharinus maculipinnis (Poey)
from Nosy Bé, Madagascar; from 2 specimens of Carcharinus leucas
(Valenciennes) from Nosy Bé and 1 specimen of the same host from
Durban.

Habitat: Body surface and fins.

Pandarus niger Kirtesinghe, 1950
Collections: From 10 specimens of Carcharinus species at stations
408 and 409; from 1 specimen of Carcharinus spallanzani (Boneparte)
at station 407.
Habitat: Body surface and pectoral fins.

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NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY 3

Pseudopandarus gracilis Kirtesinghe, 1950

Collections: From 1 specimen of Scoliodon palasorrah (Bleeker)
from Nosy Bé; from 1 specimen of Scoliodon species at station 412.
Habitat: Body surface and fins.

Pseudopandarus longus (Gnanamuthu, 1951)

Collections: From 7 specimens of Rhizoprionodon species at station
402 (Beira Harbor); from 2 specimens of Carcharinus tyutjot at the
same location; from 1 specimen of Carcharinus obesus and Rhizopri-
onodon acutus both from Durban.

Habitat: Body surface.

This copepod was originally assigned to the genus Pandarus by
Gnanamuthu, but I have placed it in the genus Pseudopandarus. A
discussion of this species together with my reasons for the change can
be found in my paper (1966b) revising the Pandaridae.

Gangliopus pyriformis Gerstaecker, 1854

Collections: From 2 specimens of Prionace glauca at stations 292
and 295; from 3 specimens of Alopias vulpinus at stations 106, 109, and
115.

Habitat: Gill filaments.

Phyllothereus cornutus (Milne-Edwards, 1840)

Collections: From 2 specimens of Prionace glauca at stations 292
and 306.
Habitat: Gill filaments.

Perissopus dentatus Steenstrup and Liitken, 1861

Collections: From 1 specimen of Carcharinus tjutjot at station 402;
from 1 specimen of Mustelus species, from 1 specimen of Carcharinus
obscurus, from 1 specimen of Carcharinus leucas, all from Durban.

Habitat: Caudal fin and right clasper.

Echthrogaleus coleoptratus (Guerin, 1837)

Collections: From 5 specimens of Prionace glauca at stations 131,
132, 306, and 309; from 2 specimens of Lamna species (mackerel
shark) at stations 132 and 309.

Habitat: Body surface.

Echthrogaleus denticulatus Smith, 1874

Collections: From 3 specimens of Alopias vulpinus at stations 106,
109, 110, 115, 288, and 289; from 2 specimens of [surus oryrinchus
Rafinesque at stations 106, 108, 123, and 282.

Habitat: Body surface, sometimes in large clusters around vent of
female.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Dinemoura latifolia (Steenstrup and Liitken, 1861)

Collections: From 2 specimens of Jsurus oryrinchus at stations
123, 139, 296, and 310; from 1 specimen of Jsurus glaucus from Durban.
Habitat: Body surface.

Dinemoura species

Collections: From 2 specimens of Alopias vulpinus at stations 289
and 295; from 1 specimen of Alopias superciliosus (Lowe) at Nosy Bé
and station 119.

Habitat: Ventral body surface.

This new species is described in detail in my paper (1967) re-
vising the Pandaridae.

Pagina tunica Cressey, 1964

Collections: From 2 specimens of Alopias superciliosus from Nosy
Bé and Majunga, Madagascar, and station 119.
Habitat: Body surface (usually ventral).

Nesippus orientalis Heller, 1868

Collections: From 6 specimens of Carcharinus maculipinnis, 4
specimens of Sphyrna lewint (Griffith), 3 specimens of Carcharinus
leucas, 1 specimen of Galeocerdo cuvier (Peron and LeSueur), 1 speci-
men of Rhizoprionodon acutus, 1 specimen of Scoliodon palasorrah, all
from Nosy Bé; from 1 specimen of Sphyrna mokarran (Riippell) at
station 409; from 1 specimen of Sphyrna zygaena (Linnaeus) from
Durban; from 2 specimens of Carcharodon carcharias from Durban.

Habitat: Roof of mouth and gill arches.

Nesippus crypturus Heller, 1865

Collections: From 13 specimens of Carcharinis maculipinnis, 5
specimens of Sphyrna lewinr, 5 specimens of Carcharinus leucas, 1
specimen of Galeocerdo cuvier, all from Nosy Bé; from 1 specimen of
Carcharinus longimanus at station 291; from 1 specimen of Carcharinus
leucas at station 409; from 1 specimen of Sphyrna mokarran at station
409; from 1 specimen of Carcharinus species ( floridanus?) at station
412.

Habitat: Roof of mouth and gill arches.

Nesippus species

Collections: From 1 specimen of Galeocerdo cuvier from Nosy Bé;
from 1 specimen of the same host caught off Comores Island (Zaudsi).

Habitat: Nasal cavities.

This new species is described in detail i my paper (1967) revising
the Pandaridae.

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY a

Family Euryphoridae Wilson, 1905

Alebion gracilis Wilson, 1905

Collections: From 1 specimen of Carcharinus albimarginatus at
station 400; from 1 specimen of Carcharinus leucas at station 409; from
1 specimen of Carcharinus species at station 381; from 1 specimen of
Carcharinus species at station 412; from 1 specimen of Galeocerdo cuvier
from Comores Island; from 1 specimen of Carcharinus longimanus at
station 291; from 1 specimen of Carcharinus obscurus from Durban.

Habitat: Body surface and fins (dorsal surface usually).

Alebion alatus Gnanamuthu, 1951
Collection: From 1 specimen of Carcharinus limbatus from Nosy Bé.
Habitat: Body surface.

This copepod was originally described from a carcharinid shark of
India and this new record is only the second record of this species.

Alebion elegans Capart, 1953
Collection: From 1 specimen of Sphyrna mokarran at station 409.
Habitat: Caudal fin.

This copepod has previously been reported from the west coast of
Africa (Capart, 1953, and Vaissiére, 1959). This record extends the
known range to the east coast of Africa.

Paralebion elongatus Wilson, 1911

Collections: From 7 specimens of Carcharinus leucas from Nosy Bé;
from 1 specimen of the same host from Durban.

Habitat: Mouth and gill arches.

This parasite is probably worldwide in its distribution. I have
collections from the Atlantic and Pacific as well as the Indian Ocean.
It is the only euryphorid that is routinely found in the mouth of the
host.

Family Anthosomatidae Dana, 1853

Anthosoma crassum (Abildgaard, 1794)

Collection: From 1 specimen of Jsurus species caught off East
London, South Africa; from 1 specimen of Jsurus oxyrinchus at station
139.

Habitat: Near base of teeth of the host.

This copepod has been reported many times and apparently is
worldwide in distribution.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Family Eudactylinidae Yamaguti, 1963

Nemesis lamna Risso, 1826

Collections: From 2 specimens of Jsurus oxryrinchus at stations 296
and 310; from 2 specimens of Jsurus species and 1 specimen of Car-
charodon carcharias from Durban.

Habitat: Gill filaments.

Nemesis versicolor Wilson, 1913
Collections: From 12 specimens of Carcharinus maculipinnis from
Nosy Bé.
Habitat: Gill filaments.

Nemesis robusta (Beneden, 1851)

Collections: From 1 specimen of Sphyrna mokarran at station 409;
from 2 specimens of Sphyrna lewini and 4 specimens of Carcharinus
leucas from Nosy Bé.

Habitat: Gill filaments.

Nemesis aggregatus, new species
Fiaures 1, 2, 6-20

Collections: From 5 specimens of Alopias vulpinus at stations 109,
110, 115, 288, and 289. Material from station 288 designated as types.
Holotype, female, USNM 113291. Allotype, male, USNM 113292.
Paratypes female and male USNM 113293.

FrmaLe.— Body form as in figure 1. Total length 4.5 mm. Greatest
width 1.5 mm. Cephalon slightly longer than wide (1.3 by 1.2 mm),
comprising nearly one-third of body length. First pedigerous segment
(2nd thoracic segment) fused with head. Thoracic segments 3-5 free,
about equal in length (0.85 mm). Thoracic segment bearing leg 5 free,
not combined to form genital segment (fig. 6). Genital segment wider
than long (0.7 by 0.4 mm), often with spermatophores attached at
posterior corners (fig. 6). Abdomen 3-segmented, segments measuring
140u, 103u, and 140u respectively. Caudal rami (fig. 7) about twice
as long as wide (85u by 40y), each with 6 posterior setae. All setae of
about equal length, 2 broad and blunt as in figure.

First antenna (fig. 8) of 12 segments, bearing short setae. Second
antenna (fig. 9) in form of claw, terminal segment with 2 short setae.
Second segment with patch of 10-12 short, broad, spinules on inner
surface. Mouth tube and mandible as in other members of genus.
First maxilla (fig. 10) biramose, each ramus with 2 terminal setae.
Second maxilla, in general, as in other members of genus. Tip armed
as in figure 11. Maxilliped (fig. 12) in form of stout claw.

Figures 1-2.—Nemests aggregatus,

new species, dorsal view: 1, female; 2, male.

Figures 3-5.—Eudactylina pusilla, new species: 3, dorsal view, female. Eudactylina pollex,

new species: 4, dorsal view, female. Kroyeria gemursa, new species: 5, dorsal view,
female.

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY C

Legs 1-4 (figs. 13-16) biramose, all rami 2-segmented. Spine and
setal formula as follows:

leg 1 leg 2 leg 3 leg 4
; exo. end. exo. end. exo. end. exo. end.
seg. 1 1:0 0:1 It 0:1 leealt 0:1 I:I 0:1
seg. 2 eta Il VII V VII IV VIII IV

Leg 1 with endopod turned so that inner edge faces exopod. First
segment of exopod curved toward endopod forming chela-like struc-

Ficures 6-15.—Nemesis aggregatus, new species, female: 6, abdomen, dorsal; 7, caudal
ramus; 8, first antenna; 9, second antenna; 10, first maxilla; 11, tip of second maxilla; 12,
maxilliped; 13, first leg; 14, second leg; 15, third leg.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

ture. Legs 2-4 with rows of spinules as indicated in figures. Leg
5 (see fig. 6) a lobe on posterior corner of thoracic segment 5 bearing
3 short setae. Leg 6 absent.

Egg strings uniseriate, long.

Ma.e.—Body form as in figure 2. Total length4.0mm. Greatest
width 1.0 mm. Genital segment as long as wide (0.8 mm). Con-
figuration of body otherwise as in female. Caudal rami as in female.

Oral area and cephalic appendages as in female.

Legs 1-4 biramose. All rami 2-segmented. Spine and setal for-
mula as follows:

leg 1 leg 2 leg 3 leg 4
exo. end. exo. end. exo. end. exo. end.
seg. 1 1:0 0:1 eh 0:1 tt 0:1 et O:1
seg. 2 nel II 1Ves- 56 1:4 5 T5375

Legs 2-4 (figs 17,18, and 19) armed as in figures. Terminal endopod
segment of leg 3 bearing a modified seta as shown in figure 18. Ter-
minal endopod segment of leg 4 also with modified seta but differing
from that of leg 3 (compare figs. 18 and 19). Leg 5 a lobe with 3
setae along distal border and 1 seta near base of lobe. Segment
bearing leg 5 small and inconspicuous. Leg 6 (fig. 20) a broad lobe on
ventrodistal corner of genital segment bearing 3 setae, innermost
twice as long as outer 2.

Remarks.—This species differs from N. pallida Wilson, N. vermi
Scott, N. robusta (van Beneden), N. atlantica Wilson, N. pilosa Pearse,
and N. macrocephalus Shimo by the nature of the spinules on segment
2 of the second antenna. In N. aggregatus there is a patch of 10-12
prominent spinules on this segment whereas in the above-mentioned
species this patch is composed of 25-40 small spimules. Nemesis
tiburo and N. versicolor differ from N. aggregatus in the presence of a
row of fine spinules on segment 3 of second antenna in addition to
those on segment 2.

Habitat: The parasites were found attached to the free end of
each gill filament nearest to the gill slit. The tissue at the site of
attachment was swollen and pale in color (normally red). Each
gill filament could carry as many as 8 copepods on its tip.

The name aggregatus from Latin, meaning “clustered or united,”
refers to the appearance of the species.

Eudactylina aspera Heller, 1865
Collections: From 8 specimens of Carcharinus maculipinnis; from
2 specimens of Rhizoprionodon acutus; from 1 specimen of Sphyrna
leuwini; all from Nosy Bé.
Habitat: Gill filaments.

NO. 8572 INDIAN OCEAN COPEPODS—CRESSEY 9

Eudactylina pusilla, new species
Fiaurss 3, 21-30

Collection: 3 females from 1 specimen of Galeocerdo cuvier from
Nosy Bé. Holotype USNM 113299. Paratype USNM_ 113300.
One specimen dissected for study.

FrmMaLe.—Body form as in figure 3. Dorsal surface of body
covered with broad spinules. Total length 4.6 mm. Greatest width
1.3mm. Cephalon wider than long (1.3 by 0.8 mm), widest postero-
laterally. First pedigerous thoracic segment fused with head.

Ficures 16-22.—Nemesis aggregatus, new species, male: 16, fourth leg; female: 17, second
leg; 18, third leg; 19, fourth leg; 20, sixth leg. Eudactylina pusilla, new species, female:
21, abdomen and genital segment, lateral; 22, caudal ramus.

221-536—67——__2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Thoracic segments 3-5 free. Thoracic segment bearing leg 5 free,
not combined to form genital segment. Genital segment (fig. 21)
somewhat wider than long (0.8 by 0.7 mm). Abdomen 2-segmented.
Caudal ramus (fig. 22) terminating in blunt process with accessory
spine and 2 subterminal setae.

Ficures 23-30.—Eudactylina pusilla, new species, female: 23, first antenna; 24, second
antenna; 25, mandible and first maxilla; 26, second maxilla; 27, maxilliped; 28, first leg;

29, second leg; 30, fourth leg.

First antenna (fig. 23) 5-segmented, second segment with stout
clawlike spine on outer distal corner. Other segments armed as
in figure. Second antenna (fig. 24) 3-segmented, second segment

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY it

with sclerotized process on inner border. Terminal segment with
clawlike spine at tip. Mandible and first maxilla (fig. 25) as in other
members of genus. Mandible with 5 terminal teeth. Second maxilla
(fig. 26) with terminal claw, segment bearing claw with patch of
prominent spinules. Maxilliped (fig. 27) chelate, tip of movable
process bearing broad scalelike spine opposed by process on preceding
segment.

Legs 1-4 biramose, covered with stout spinules. Spine and setal
formula as follows:

leg 1 leg 2 leg 8 leg 4
end. exo. end. exo. end. exo. end.
seg. 1 1:0 0:0 1:0 0:0 T:0 0:0 1:0 0:0
seg. 2 1:0 0:0 1:0 0:0 1:0 0:0 10 0:0
seg. 3 3 2 II i Til 1 III 1

Leg 1 (fig. 28) with 1 long and 1 short seta at tip. Outer edges of
segments on exopod and endopod with rows of pointed spines as in
figure. Leg 2 (fig. 29) exopod elongate as in figure, terminal exopod
segment bearing 2 short inner spines. Second exopod segment re-
duced. First exopod segment with prominent lobe on inner distal
corner. Endopod without spines and setae except for very small
spine at tip. Leg 3 (fig. 30) exopod with well-developed spines on
outer corners of segments. Endopod unarmed except for terminal
seta. Leg 4sameasleg 3. Leg 5 (see fig. 21) broad lobe with 3 distal
setae and 1 basal seta. Lobe covered with spinules. Leg 6 (see
fig. 21) reduced to single short spine on genital segment at area of egg
string attachment.

Egg strings uniseriate.

Mare.—Unknown.

Remarks.—This species can be distinguished from all other mem-
bers of the genus by the prominent lobe on the inner distal corner of
the first exopod segment of leg 2. It is further characterized by the
conspicuous spinules on the rami of legs 1-5.

Habitat: Gill filaments.

The name pusilla, from Latin, meaning “small,” refers to the size of

the species.
Eudactylina pollex, new species

Figures 4, 31-42

Collection: 23 females from 1 specimen of Sphyrna mokarran at
station 409. Holotype female USNM 113297. Paratypes (21
females) USNM 113298. One specimen dissected for study.

FremaLte.—Body form as in figure 4. Dorsal surface of body
smooth. Total length 3.6mm. Greatest width 0.8mm. Cephalon
slightly longer than wide (0.72 by 0.66 mm). First pedigerous
thoracic segment fused with head. Thoracic segments 3-5 free,

12 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 121

measuring 360u, 600u, and 960u, respectively. Thoracic segment
bearing leg 5 free, not combined to form genital segment. Genital
segment somewhat wider than long (3004 by 340u). Abdomen 2-
segmented, segments measuring 141y and 118y, respectively. Caudal
rami (fig. 31) about twice as long as wide (132y by 75), bearing 3
terminal spines and 3 subterminal setae. Rami covered with spinules
on dorsal surface.

Ficures 31-40.—Eudactylina pollex, new species, female: 31, caudal ramus; 32, first an-
tenna; 33, second antenna; 34, mandible; 35, first maxilla; 36, second maxilla; 37, maxil-
liped; 38, first leg; 39, second leg; 40, endopod of third leg.

Oral area as in other members of genus. First antenna (fig. 32)
5-segmented, each segment armed with short, naked setae as in
figure. Second antenna (fig. 33) 4-segmented. Terminal segment in

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY 13

form of short claw bearing 3 spines. Second segment with 2 short
setae on inner margin. Mandible (fig. 34) styliform process project-
ing with mouth tube, armed with 4 teeth at tip. Second maxilla
(fig. 36) with patches of stout spinules as in figure. Maxilliped (fig.
37) chelate. Movable process of maxilliped opposed by immovable
dactylate process on preceding segment. Immovable process with
distal depression into which tip of movable process fits.
Legs 1-4 biramose with spine and setal formula as follows:

leg 1 leg 2 leg 3 leg 4
exo. end. exo. end. 0. end. exo. end.
seg. 1 1:0 0:0 0 0:0 TO 0:0 1:0 0:0
seg. 2 1:0 1 1:0 0:0 1:0 0:0 1:0 1
seg. 3 III III 4 Iil 1 Iil

Leg 1 (fig. 38) bearing stout spinules on both rami. Exopod 3-
segmented. Endopod 2-segmented. Leg 2 (fig. 39) with both rami
3-segmented, small spinules on both rami. Leg 3 similar to leg 4
except endopod last segment incompletely divided (fig. 40). Leg
4 (fig. 41) endopod 2-segmented. Exopod 3-segmented. Exopods of
legs 2-4 with few spinules on first segment only. Leg 5 (fig. 42) a
lateral lobe on thoracic segment 6, bearing 2 terminal setae, 1 sub-
terminal seta. Leg 6 represented by short spine incorporated into
area of ege string attachment.

Egg strings uniseriate.

Mae.— Unknown.

Remarks.—This new species can be separated from all known
species of the genus (except EL. acanthii Scott, EL. turgipes Bere, and
E. corrugata Bere) by the exopod of leg 2. In all other species the
exopod is unusually long and somewhat recurved (compare figs. 29
and 39). . pollex can be separated from the other 3 species by the
segmentation of the legs. In E. acanthii and E. corrugata the exopods
of legs 1-4 are 2-segmented. In E. pollex the exopods of legs 2 and
3 are 3-segmented. The endopod of leg 4 of EL. turgipes is 3-segmented.
It is 2-segmented in LZ. pollez.

Habitat: Gill filaments.

The name pollex, from Latin, meaning ‘‘thumb,” refers to the shape
of the endopods of legs 2-4.

Kroyeria gracilis Wilson, 1932

Collections: From 3 specimens of Carcharinus leucas from Nosy Bé;
from 1 specimen of the same host from station 409; from 1 specimen
of Carcharinus longimanus from station 291.

Habitat: Gill filaments.

This copepod has been reported from a wide range of hosts and is
probably cosmopolitan in distribution.

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Kroyeria dispar Wilson, 1935

Collections: From 3 specimens of Galeocerdo cuvier from Nosy Bé.
Habitat: Gill filaments.

The only previous report of this copepod is the original description
from Puerto Rico from Squalus.

A2

Ficures 41-48.—Eudactylina pollex, new species, female: 41, fourth leg; 42, fifth leg.
Kroyeria gemursa, new species, female: 43, distal end of abdomen and caudal rami; 44,
first antenna; 45, second antenna; 46, labrum; 47, second maxilla; 48, maxilliped.

Kroyeria spatulata Pearse, 1948

Collections: From 5 specimens of Carcharinus maculipinnis, 2
specimens of Carcharinus sorrah, and 2 specimens of Rhizoprionodon
acutus all from Nosy Bé.

Habitat: Gill filaments.

This copepod has been reported previously from off the south-
east coast of the United States. The previously reported hosts

NO. 3572 INDIAN OCEAN COPEPODS—-CRESSEY 15

plus the new records here indicate that this species may be restricted
to sharks generally found only in shallow coastal waters.

een

50

SS

= \

—
fF

52 so)
ir}

Wy

Figures 49-53.—Kroyeria gemursa, new species, female: 49, first leg; 50, second leg; 51,
third leg; 52, fourth leg. Male: 53, genital segment, abdomen and caudal rami.

Kroyeria gemursa, new species
FicureEs 5, 43-53
Collections: 4 females and 4 males from 1 specimen of Sphyrna
mokarran at station 409. Holotype, female, USNM 113294. Allo-
type, male, USNM 113295. Paratypes (2 females, 3 males) USNM
113296. One female specimen dissected for study.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

FrmMa.LE.—Body form as in figure 5. Total length 9.7mm. Great-
est width 1.5 mm. Cephalon as wide as long (1.3 mm), widest near
posterior border. Thoracic segments bearing legs 1-4 free. Segment
bearing leg 1 with well-developed aciculum, 1.3 mm in length, bifid
at tip. Thoracic segments 6 and 7 fused to form genital segment.
Genital segment long, comprising slightly more than one-half body
length. Abdomen 2-segmented. Terminal segment with posterior
sclerotized ring at point of attachment of caudal rami (see fig. 43).
Caudal rami (fig. 43) 3 times as long as wide (264u by 94u). Each
ramus fringed along inner margin, bearing 4 terminal and 2 sub-
terminal setae. Outer 2 terminal setae stout, plumose on inner
margins only.

Oral area as in other members of genus. First antenna (fig. 44)
7-segmented, each segment with short, naked setae as in the figure.
Second antenna (fig. 45) chelate. Movable process with short spine
on inner margin, tips of immovable and movable processes pointed.
Mouth tube short. Labrum with lateral spinose lobes. Mandible in
form of stylet. First maxilla biramose. Second maxilla (fig. 47)
with short, blunt claw at tip. Manxilliped (fig. 48) with long re-
curved claw. Maxilliped most prominent appendage of cephalon.

Legs 1-4 biramose; each ramus 3-segmented. Spine and setal
formula as follows:

leg 1 leg 2 eg 3 leg 4
exo. end. exo. end. exo. end. exo. end.
seg. 1 en 0:1 Ta 0:1 T:1 0:1 Werk O:1
seg. 2 0:1 0:0 Tien 0:0 Ted 0:0 Teal O:1
seg. 3 I1:4 6 TE5 6 Il:4 4 II:4 3

Leg 1 (fig. 49) exopod with fringe on outer margin of all 3 segments.
Outer margin of Jast 2 endopod segments with row of many short
spines. Legs 2 (fig. 50) and 3 (fig. 51) rami with outer margins armed
as leg 1. Leg 4 (fig. 52) exopod with fringe on first segment only.
Endopod outer margin as in preceding legs. Each corner of intercoxal
plate of legs 2-4 with well-developed, posteriorly directed spine.
Corners of intercoxal plate of leg 1 with knobs. Leg 5 represented by
4 setae located on margin near middle of genital segment. Setae
arranged in 2 groups of 2 setae each. Leg 6 a single seta at posterior
corner of genital segment.

Egg strings uniseriate.

Maue.—Body form and appendages like those of female. Total
length 4.4 mm. Greatest width 1.0 mm. Male differs from female
in form of genital segment, abdomen, and caudal rami (fig. 53).
Abdomen 38-segmented, end of abdomen not sclerotized as in female.
Caudal rami nearly 6 times as long as wide (352y by 61y).

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY ws

Figure 54.—Map of Indian Ocean showing various stations of the R/V Anton Bruun cited
in this paper.

Legs 1-4 armed as in female except spines on outer edges of endopod
segments fewer in male. Leg 5 as in female. Leg 6 consisting of 2
setae on posterior corner of genital segment.

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Spermatophores easily visible within genital segment.

Remarks.— Kroyeria gemursa can be separated from all known
members of the genus by the sclerotized area at the end of the abdomen.
It can be further distinguished from all species except K. dispar by
the lateral swellings on the labrum. It can be separated from K.
dispar by the presence of the prominent spines on the intercoxal plates
of legs 2-4.

Habitat: Gill filaments.

The name gemursa, from Latin, meaning “swelling between toes,”
refers to sclerotization on abdomen.

Bariaka alopiae Cressey

Collection: From 1 specimen of Alopias superciliosus at Nosy Bé
and station 119.

Habitat: Guill filaments.

Copepods Parasitic on Sharks Examined

Lamnidae

TIsurus oxyrinchus
Pandarus smithii
Echthrogaleus denticulatus
Dinemoura latifolia
Nemesis lamna
Anthosoma crassum

Isurus glaucus
Dinemoura latifolia

TIsurus species
Anthosoma crassum
Nemesis lamna

Lamna species
Echthrogaleus coleoptratus

Carcharodon carcharias
Pandarus cranchii
Nesippus orientalis
Nemesis lamna

Alopidae

Alopias vulpinus
Pandarus cranchii
Pandarus smithii
Gangliopus pyriformis
Echthrogaleus denticulatus
Dinemoura sp.
Nemesis aggregatus

Alopias superciliosus
Dinemoura species
Pagina tunica

Bariaka alopiae
Triakidae

Mustelus species
Perissopus dentatus
Carcharinidae

Carcharinus longimanus
Pandarus cranchii
Nesippus crypturus
Alebion gracilis
Kroyeria gracilis

Carcharinus floridanus
Pandarus cranchii
Pandarus smithii
Nesippus crypturus

Prionace glauca
Pandarus satyrus
Phyllothereus cornutus
Gangliopus pyriformis
Echthrogaleus coleoptratus
Galeocerdo cuvier
Nesippus orientalis
Nesippus crypturus
Nesippus sp.
Alebion gracilis
Kroyeria dispar
Eudactylina pusilla
Carcharinus obscurus
Perissopus dentatus
Alebion gracilis

NO. 3572 INDIAN OCEAN COPEPODS—CRESSEY 19

Carcharinus obesus?
Pseudopandarus longus
Carcharinus leucas
Pandarus carcharini
Perissopus dentatus
Nesippus orientalis
Nesippus crypturus
Alebion gracilis
Paralebion elongatus
Nemesis robusta
Kroyeria gracilis
Carcharinus maculipinnis
Pandarus carcharini
Nesippus orientalis
Nesippus crypturus
Nemesis versicolor
Eudactylina aspera
Kroyeria spatulata
Carcharinus limbatus
Alebion alatus
Carcharinus albimarginatus
Alebion gracilis
Carcharinus spallanzani
Pandarus niger
Carcharinus tjutjot
Pseudopandarus longus
Perissopus dentatus
Carcharinus sorrah
Kroyeria spatulata

ABILDGAARD, P. C.

Scoliodon palasorrah
Pseudopandarus gracilis
Nesippus orientalis

Rhizoprionodon acutus
Pseudopandarus longus
Nesippus orientalis
Eudactylina aspera
Kroyeria spatulata

Rhizoprionodon species
Pseudopandarus longus

Scoliodon species
Pseudopandarus gracilis

Carcharinus species
Pandarus niger
Alebion gracilis

Sphyrnidae

Sphyrna mokarran
Nesippus orientalis
Nesippus crypturus
Alebion elegans
Nemesis robusta
Eudactylina pollex
Kroyeria gemursa

Sphyrna lewini
Nesippus orientalis
Nesippus crypturus
Nemesis robusta
Eudactylina aspera

Sphyrna zygaena
Nesippus orientalis

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20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

CressEy, R. F.
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1854. Beschreibung zweier neuer Siphonostomen-Gattungen. Arch. Naturg.,
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1963. On five species of Formosan parasitic copepods belonging to the sub-

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NO. 3572 INDIAN OCEAN COPEPODS—-CRESSEY 21

Scort, T.

1901. Notes on some parasites of fishes. 19th Ann. Rep. Fish. Bd. Scotland,

pt. 3, pp. 120-153.
Suino, 8. M.

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VAISSIERE, R.

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Witson, C. B.

1905. New species of parasitic copepods from the Massachusetts Coast.
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Nat. Mus. Bull. 158, 635 pp.

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YAMAGUTI, S.
1963. Parasitic Copepoda and Branchiura of fishes, 1104 pp.

1

=

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3573

NEW CYCLOPOID COPEPODS
ASSOCIATED WITH THE ALCYONARIAN CORAL
TUBIPORA MUSICA (LINNAEUS)

IN MADAGASCAR

By Artuur G. Humes ann Ju-SHey Ho!

At Nosy Bé, in the northwestern part of Madagascar, several species
of copepods (Humes and Frost, 1964) are known to be associated
with members of the aleyonarian order Alcyonacea, but until now
none have been reported living with members of the order Stolonifera.

While participating in the work of the U.S. Program in Biology of
the International Indian Ocean Expedition at Nosy Bé in 1963-64,
the first author collected 4 species of cyclopoid copepods from wash-
ings of Tubipora musica (Linnaeus), the common organ-pipe coral.
These species are described below.

All figures have been drawn with the aid of a camera lucida.

The study of the material has been aided by a grant (GB-1809)
from the National Science Foundation. We wish to thank Dr.
Michel Pichon for verification of the identity of the coral and to
acknowledge the many courtesies received from the staff of the Centre
d’Océanographie et des Péches at Nosy Bé.

1 Both authors: Department of Biology, Boston University, Boston, Mass.
1

2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121

Family Lichomolgidae Kossmann, 1877
Genus Lichomolgus Thorell, 1860

Lichomolgus oerganicus, new species

Figures 1-30

TYPE MATERIAL.—207 99, 98 oo’, and 15 copepodids from a
colony of Tubipora musica (Linnaeus), in 1m, Pte. Ambarionaomby,
Nosy Komba, near Nosy Bé, Madagascar, collected June 8, 1964:
holotype 2 (USNM 113387), allotype (USNM 113388), and 144
paratypes (USNM 113389) (100 99 and 44 @o@) deposited in the
United States National Museum, 80 paratypes (50 99 and 30 oo)
in the Museum of Comparative Zoology, Harvard University, and
the remaining paratypes in the collection of A. G. Humes.

OTHER spEcIMENS.—AI from Tubipora musica collected in 1964:
8 99 and 8 o'’, in 1 m, Pte. 4 la Fiévre, Nosy Bé, Jan. 5; 1 9, in
15 cm, Tany Kely, a small island south of Nosy Bé, Mar. 29; 4 99
and 8 oc’, in 1 m, Pte. Ambarionaomby, Mar. 31; 15 99 and 1
co, in 1 m, Pte. a la Fiévre, Apr. 15; and 14 99, in 2 m, Pte.
Ambarionaomby, Sept. 27.

FrmaLe.—Body (fig. 1) with moderately broad prosome. Body
length (not including setae on caudal rami) 0.99 mm (0.92-1.05 mm)
and greatest width (in middle of cephalosome) 0.49 mm (0.44-0.53
mm), based on 10 specimens. Ratio of length to width of prosome
1.5:1. Segment of first leg separated dorsally and laterally from
head by a transverse furrow; lateral areas of this segment not ex-
panded. Lateral areas of segments bearing legs 2 and 3 expanded
and rounded posteriorly; those of segment of leg 4 smaller and
somewhat truncated.

Seement of leg 5 (fig. 2) wider than long, 61y x 133y, with fifth
legs borne laterally. Ventrally between this segment and the genital
segment no apparent intersegmental sclerite. Genital segment (fig. 2)
about as long as wide, 140u x 142u, in dorsal view expanded laterally
in its midregion, posterior to which the segment is constricted so
that its width at the posterior end is only 80u. Areas of attachment of
ege sacs located dorsolaterally on posterior part of expansions. Each
area (fig. 3) with 2 naked setae, 12 and 13y in length; posterior to the
setae a small rounded sclerotized protuberance. Three postgenital
segments, each without ornamentation except for a few surficial
hairs (sensilla), 837u x 67y, 264 x 55y, and 28u x 51yn, respectively, from
anterior to posterior.

Caudal ramus (fig. 4) only a little elongated, 36u x 24u in greatest
dimensions, or 1.5 times longer than wide. Length along inner edge

NO. 3573 COPEPODS—HUMES AND HO 3

to innermost distal seta 23u, along outer edge to base of lateral seta
23u, and to base of outermost distal seta 33u. Of the usual 6 setae,
outer lateral seta naked and 66y long; remaining 5 feathered: pedi-
cellate dorsal seta 47u, innermost distal seta 115y, outermost distal
seta 91yu, and the 2 long median terminal setae, both “pegged,”
169u (outer) and 200 (inner), and inserted between dorsal (unorna-
mented) and ventral (with marginal row of spinules) flaps. In
addition, a minute hyaline setule dorsally near insertion of outermost
distal seta. Two hairs on dorsal surface of ramus.

Dorsal surface of prosome and dorsal and ventral surfaces of
urosome with a few minute hairs. Ratio of length of prosome to that
of urosome about 2.1:1.

All ovigerous females observed with broken ege sacs (one shown
in fig. 5).

Rostral area (fig. 6) moderately well formed. At level of bases of
second antennae a small longitudinal sclerotized ridge between rostrum
and anterior part of labrum.

First antenna (fig. 7) 7-segmented, with third segment showing
ventrally a proximal sclerotized area suggesting an intercalary seg-
ment. Lengths of the segments, measured along their posterior
nonsetiferous margins: 28y (56u along anterior margin), 95y, 25, 50p,
50u, 37p, and 22u, respectively. Formula for armature: 4, 13, 6, 3,
4+1 aesthete, 2+1 aesthete, and 7+1 aesthete. All setae naked.

Second antenna (fig. 8) 4-segmented, with last segment moderately
elongated, 65u along its outer edge, 39u along its inner edge, and 27
wide. Each of first 2 segments with a small inner seta, third segment
with 3 setae (1 of them jointed), and last segment with 7 elements:
5 setae and 2 unequal recurved claws 45u and 34y long (measured along
ereatest axis). All setae naked.

Labrum (fig. 9) with 2 relatively short posteroventral lobes, free
border of each lobe showing 2 small hyaline lamellae.

Mandible (fig. 10) with its basal region bearing on its posterior
surface a small sclerotized area and separated by a constriction from
its distal region, whose inner margin bears a prominent sclerotized
pointed toothlike process followed by a striated fringe, whose outer
margin has a row of slender spinules and which terminates in a long
flagellum with lateral spinules. Paragnath (see fig. 9) a small lobe
bearing inner hairs. First maxilla (fig. 11) a single segment bearing
2 terminal setae. Second maxilla (fig. 12) 2-seemented, first segment
unarmed, second segment produced distally and bearing row of 4-5
teeth (compare figs. 12 and 13) and slender naked terminal process.
(If 4 such teeth, distalmost often slightly bifurcated; if 5, teeth all
entire.) On postero-inner surface of second segment 2 setae, one
finely pectinate along one margin, other bearing long bluntly tipped

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

setules along one side. A minute setule on proximal outer edge of
this segment. Maxilliped (fig. 14) 3-segmented. Two naked setae on
second segment. Third segment (fig. 15) bearing a naked seta, a
setiform element with bifurcated tip, and a terminal process about
22u long and clawlike, with a small hyaline lobe on each side of dis-
tal point.

Postoral area (fig. 16) only slightly produced ventrally; an incom-
plete line of sclerotization between bases of maxillipeds.

Legs 1-4 (figs. 17-20) with trimerous rami except for 2-segmented
endopod of leg 4. Armature of legs as follows (Roman numerals=
spines, Arabic numerals=setae):

Pa protopod O:1 1:0 exp 1:0 Mea III,1,4
end 0:1 0:1 1,5

iP protopod 0:1 1:0 exp 1:0 I:1 TitT 5
end Oe 0:2 I,1I,3

P 3 protopod 0:1 1:0 exp 1:0 Tet III, 1,5
end O:1 0:2 ELT

PA protopod 0:1 1:0 exp 1:0 Let IDL
end O:1 II

Inner seta on coxa of legs 1-3 long and plumose, but in leg 4 short
(17u) and naked. In first 3 pairs of legs inner margin of basis with
short row of hairs, but such hairs absent in leg 4. In all 4 legs setae
on inner side of last segment of exopod with lateral hairs shorter prox-
imally than distally. Spine on last segment of endopod of leg 1
24u inlength. Endopod of leg 4 shorter than exopod. First segment
33u X 25u with inner distal plumose seta 41y long. Second segment
61u long (including 2 terminal spinous processes) and 23 wide in
basal half and 19u wide in distal half, the 2 halves demarcated by
very slight indentation. Along outer margin of both segments rows
of minute spinules. Terminally second segment with a row of minute
spinules near insertion of 2 unequal spines, both with spinulose
fringes, outer spine 21y, inner 39u long, ratio about 1:1.86.

Leg 5 (fig. 21) with free segment elongated and slightly arcuate in
outline, 664 x 23u, with short stout spines along outer edge and extend-
ing around to distal ventral surface; bearing terminally 2 naked setae
35u and 37u in length. Seta on body near free segment slightly
feathered with group of spines near its insertion.

Leg 6 probably represented by 2 setae near areas of attachment of
each ege sac (see fig. 3).

Color in life in transmitted light rather translucid, eye red.

Maur.—Body (fig. 22) resembling in general form that of female.
Length (excluding ramal setae) 0.89 mm (0.85-0.92 mm) and greatest
width 0.32 mm (0.30-0.34 mm), based on 10 specimens. Ratio of
length of prosome to that of urosome 1.6:1.

NO. 3573 COPEPODS—-HUMES AND HO 5

Segment of leg 5 (fig. 23) 29u x 67u. Genital segment longer than
wide, 164u x 122u in greatest dimensions, and in dorsal view rather
bottle shaped, its lateral borders gently rounded. No intersegmental
sclerite ventrally between these 2 segments. Four postgenital seg-
ments 18u x 45u, 194 x 43y, 15u x 40u, and 17y x 43yn, respectively,
from anterior to posterior.

Caudal ramus (fig. 24) resembling that of female, but slightly less
elongated, its greatest dimensions 3lu x 21y, or 1.48 times longer
than wide.

Surfaces of prosome and urosome with minute hairs as in female.
Ratio of length of prosome to that of urosome 1.56:1.

Rostral area like that in female.

First antenna resembling that of female, but an aesthete added on
segments 2 and 4 (at exact points indicated by arrows in fig. 7), so
that formula is 4, 13+1 aesthete, 6, 3-+1 aesthete, 4+1 aesthete,
2+1 aesthete, and 7+1 aesthete. Second antenna (fig. 25) more
slender than in female, and last segment elongated, 68u along outer
edge, 52u along inner edge, and 15y wide. First segment with inner
crescentic row of small spinules. Second segment with numerous
small spinules along inner surface, arranged in 2 groups proximal and
distal to seta. Last segment with row of very small spinules along
inner surface. Armature like that of female, but 2 claws more nearly
equal in length (34u and 35y), though posterior claw more slender
than other.

Labrum, mandible, paragnath, first maxilla, and second maxilla like
those in female. Maxilliped (fig. 26) 4-segmented (assuming that
proximal part of claw represents fourth segment). First segment
unornamented. Second segment bearing on medial surface 2 setae
and 2 rows of long spinules. Small third segment unarmed. Slender
slightly recurved terminal claw 178y in length (measured along axis),
with narrow terminal lamella, smooth lamellar fringe along its concave
surface, and 2 setae near its base, one on posterior surface 44u long
and finely barbed, other on anterior surface 17y long, naked, and
hyaline. Suggestion of division about midway along claw.

Postoral area resembling that of female.

Legs 1-4 resembling those in female and with same spine and setal
formula, except for endopod of leg 1 (fig. 27), which has arrangement
of 0:1;0:1;1,1,4. Onlast segment of this endopod 2 spines, outermost
of setae in female being here replaced by a spine; outer spine 29y long,
inner spine 43y long and recurved distally, and both spines with strong
lateral spinules. In alcoholized specimens last segment of endopod
flexed outwardly. Terminal spines on endopod of leg 4 in one male
19» (outer) and 51y (inner), in another 16u and 37y, with an average
of 17.5y and 44uy, ratio of 1:2.5.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Leg 5 (fig. 28) with free segment elongated and slender, 36yu x Qu,
with nearly parallel sides in dorsal view, and with 2 terminal setae
unequal, outer 31y and inner 19, long. Few minute spinules on outer
distal surface of segment. No spinules near insertion of seta on body
near free segment.

Leg 6 (fig. 29) a posterolateral flap on ventral surface of genital
segment bearing 2 naked setae 23u and 25y long.

Spermatophore (fig. 30) inside body of male about 148y x 68y plus
neck of 14u.

Color in life resembling that of female.

Erymoutocy.—The specific name organicus, from Latin, relating
to musical instruments, refers to the association of this species with
the organ-pipe coral Tubipora musica.

CoMPARISON WITH RELATED SPECIES.—Among those species in the
large genus Lichomolgus, wherein the mandible has been described,
there are 3 that, like LZ. organicus, show a prominent toothlike
process on the proximal inner margin of this appendage. From these,
L. organicus may be readily distinguished. In L. actinophorus Humes
and Frost (1964) there is a single claw on the last segment of the
second antenna, the free segment of leg 5 in the female is somewhat
irregular and has a ratio of 2.1:1, the caudal ramus of the female has
a ratio of 3.5:1, and there is a setiferous sphere on the second segment
of the second maxilla. In ZL. decorus Humes and Frost (1964) there
is one claw and one clawlike spine on the last segment of the second
antenna, the last segment of the exopod of leg 4 has the formula ITI,I,5,
and the free segment of leg 5 in the female is 2.2:1 with a basal ex-
pansion. In ZL. protulae Stock (1959) the shape of the body is trans-
formed, there are 3 strong claws and 2 setiform claws on the last
segment of the second antenna, the caudal ramus of the female has a
ratio of about 8.3:1, the free segment of leg 5 in the female is about
2.1:1, and the toothlike process on the mandible is directed distally
rather than proximally as in L. organicus. In addition, the females
of L. actinophorus and L. protulae are distinctly larger.

There are 8 species of Lichomolgus in which the form of the mandible
is unknown. These may be separated from L. organicus, however, on
the basis of other characters (in each case using the female, except in
the last mentioned species where the female is unknown). In L.
dentipes Thompson and A. Scott (1903) the formula for the last
segment of the exopod of leg 4 is III,I,5 and the free segment of leg 5
has a prominent toothlike inner process. In L. elegans Thompson and
A. Scott (1903) there is 1 claw on the second antenna and the body
is larger (1.5 mm) and more slender. In ZL. gigas Thompson and
A. Scott (1903) there is one claw on the second antenna and the body
islarger (2mm). In L. longipes (Sewell, 1949) leg 5 is very long (about

NO. 3573 COPEPODS—HUMES AND HO 7

8:1) and the body is larger (1.63 mm) and more slender. In ZL.
rigidus Ummerkutty (1962) the formula for the last segment of the
exopod of leg 4 is III,I,5 and the body is larger (1.25 mm). In ZL.
rotundus Sewell (1949) the free segment of leg 5 is short, swollen, and
somewhat produced on the outer margin, with a ratio of about 2:1.
In L. tenuicornis Brady (1910) the caudal ramus is about 7:1, the free
segment of leg 5 has a small basal expansion, and the body is larger
(1.7mm). In L. vagans Gurney (1927) (based on the male), the body
is larger (1.07 mm) and the last segment of the second antenna
(Gurney, 1927, p. 466, fig. C) has a ratio of length of the outer margin
to width of about 5.6:1, thus being slightly more slender than in L.
organicus, where the ratio is about 4.5:1.

Since we still do not know the taxonomic importance of the vari-
ation in many characters observed in Lichomolgus (such as the termi-
nal armature of the second antenna, the armature of the exopod of leg
4, the exact form of the mandible and leg 5, etc.), we find it impossible
in Lichomolgus to say with which species L. organicus is most closely
related. Such a conjecture would seem to be premature in view of the
apparent very incomplete knowledge of the species in this genus.

Lichomolgus conjunctus, new species

Figures 31-36

TYPE MATERIAL.—163 99 and 25 oo’ from a colony of Tubipora
musica (Linnaeus), in 1 m, Pte. Ambarionaomby, Nosy Komba, near
Nosy Bé, Madagascar, collected June 8, 1964: holotype 9 (USNM 113391),
allotype (USNM 113392), and 89 paratypes (USNM 113393) (79 99
and 10 oo") deposited in the United States National Museum, 45
paratypes (40 99 and 5 oc”) in the Museum of Comparative Zoology,
Harvard University, and the remaining paratypes in the collection
of A. G. Humes.

OrHeR specIMENS.—AII from Tubipora musica collected in 1964:
121 99 and 53 77, in1 m, Pte. a la Fiévre, Nosy Bé, Jan. 5; 9 99, in
15 cm, Tany Kely, a small island south of Nosy Bé, Mar. 29; 8 99, in
1 m, Pte. Ambarionaomby, Nosy Komba, Mar. 31;14 99 and 3 oc,
in 1 m, Pte. 4 la Fiévre, Apr. 15; and 43 99, in 2 m, Pte. Ambari-
onaomby, Sept. 27.

In the description that follows, morphological features not specifi-
cally mentioned may be assumed to be essentially like those in L.
organicus.

FremaLe.—Body (fig. 31) with prosome more pointed anteriorly
than in LZ. organicus. Length of body (without ramal setae) 0.80 mm
(0.72-0.87 mm) and greatest width 0.38 mm (0.34-0.40 mm), based on
10 specimens. Ratio of length to width of prosome 1.53:1.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Segment of leg 5 (fig. 32) 464 x 107u. Genital segment slightly
wider than long, 110 x 115, in dorsal view expanded laterally just
behind its midregion, posterior to which segment is constricted, with
width at posterior end of segment 60. Each area of attachment of
egg sacs, located dorsolaterally on expanded part of segment, bearing 2
naked setae and rounded protuberance (more prominent than in pre-
vious species). Three postgenital segments 25 x 53y, 21p x 48y, and
23u x 46u, respectively, from anterior to posterior.

Caudal ramus (fig. 33) only slightly longer than wide, 28 & 23y in
greatest dimensions, ratio of length to width 1.22:1. Length along
inner edge to innermost distal seta 19, along outer edge to base of
lateral seta 19 and to base of outermost distal seta 25y. Pedicellate
dorsal seta apparently naked. Relative lengths of 6 setae approxi-
mately those of L. organicus.

Ratio of length of prosome to that of urosome 2.6: 1.

Egg sacs on all ovigerous females broken.

Rostral area (fig. 34) slightly more elongated posteriorly than in L.
organicus. Segments of first antenna 23y (44u), 97u, 21, 46, 45y,
31u, and 20y in length (measured as before) and arrangement of
naked setae and aesthetes as in previous species. Second antenna
resembling that of L. organicus, but last segment slightly more slender
(63u along its outer edge, 39 along its inner edge, and 21» wide).
Larger claw 37u long, and more slender claw 32y in length.

Labrum, mandible, paragnath, and first maxilla like those in L.
organicus. Second maxilla closely resembling that species also, but
pectinate seta on second segment slightly longer, reaching nearly
to end of first tooth; in females studied 4-5 entire teeth, or 4 teeth
with distalmost slightly bifurcated. Maxilliped resembling that of
L. organicus, but subterminal setiform element apparently not as
distinctly bifurcated at its tip.

Postoral area similar to that in previous species.

Legs 1-4 with same segmentation and spine and setal formula as
in L. organicus. Spine on last segment of endopod of leg 1 27y long.
Endopod of leg 4 (fig. 35) with the following dimensions: first segment
33 x 25u with inner seta 40u long, second segment 60p x 21 (greatest
width) and 18y (least width) with 2 terminal spines 25y (outer)
and 54, (inner) in length, the ratio about 2.16:1.

Leg 5 similar to that of Z. organicus, dimensions of free segment
about 63 x 24u.

Leg 6 (represented by 2 setae near attachment of egg sacs) like
that in previous species.

Color in life resembling that of Z. organicus.

No. 3573 COPEPODS—HUMES AND HO 9
Maux.—Body (fig. 36) with prosome more pointed than in previous
species. Body length (excluding setae on caudal rami) 0.71 mm
(0.65-0.76 mm) and greatest width 0.27 mm (0.23-0.30 mm), based
on 10 specimens. Ratio of length of prosome to its width 1.5:1.
Segment of leg 5 25u x 66y. Genital segment 151u x 117». Four
postgenital segments 14y x 45, 16u x 42y, 134 x 40u,and 17p x 42, respec-

tively, from anterior to posterior.

Caudal ramus 25y x 21y, or 1.19:1.

Ratio of length of prosome to that of urosome 1.5721.

TaBLE 1.—Comparison of certain features of Lichomolgus organicus and

Characters

Female

body size

anterior border of
prosome

ratio of prosome to
urosome

caudal ramus

genital segment

second antenna

spine on last segment of
endopod of leg 1

spines on last segment of
endopod of leg 4

Male

body size

anterior border of
prosome

caudal ramus

spines on last segment of
endopod of leg 4

free segment of leg 5

Rostral area like that of female.

L. conjunctus

LI. organicus

0.99 x 0.49 mm
rounded

Ze

36u x 24y (1.5:1)

lateral indentations near
middle (fig. 2)

last segment 274 wide,
claws 45y and 34y

24u

21y and 39y (1:1.85)

0.89 x 0.32 mm
rounded

oly x 21p (1.48:1)
17.54 and 44 (1:2.5)

36u x 9p (4:1)

L. conjunctus

7—_eeeeeee——————

0.80 x 0.88 mm
more pointed

2.61

28 x 23y (1.22:1)
lateral indentations
further back (fig. 32)
last segment 21p wide,
claws 37p and 32y
27 pu

25p and 54y (1:2.16)

0.71 x 0.27 mm
more pointed

254 x 21y (1.19:1)
184 and 50p (1:2.8)

28u x 8u (3.5:1)

First antenna resembling that
of L. organicus, with same formula for setae and aesthetes. Second
antenna also resembling that species; last segment 73y along outer
edge, 54y along inner edge, and 14y wide. Two claws (34 and 35y)
similar to those in L. organicus.

Labrum, mandible, paragnath, first maxilla, and second maxilla
like those in female; in males observed 4 teeth on distal end of second

271-537-672

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

maxilla, last tooth slightly bifurcated or entire. Maxilliped and
postoral area much like those in L. organicus.

Legs 1-4 resembling those of female with same spine and setal
formula, except for sexual dimorphism in leg 1 (as in L. organicus).
Two spines on last segment of endopod of leg 1 like those in previous
species, outer 27n and inner 40u long. Two terminal spines on
endopod of leg 4 18 (outer) and 50y (inner), ratio of 1:2.8.

Leg 5 resembling that of LZ. organicus, but free segment relatively
slightly shorter, 28 x 8.

Leg 6 similar to that in previous species, but 2 naked setae slightly
longer (28y).

Spermatophores, seen only inside body of male, resembling those of
L. organicus.

Color in life like that of female.

Erymo.tocy.—The specific name conjunctus, from Latin, meaning
“closely related or bordering on,’ refers to the close similarity be-
tween this and the preceding species.

CoMPARISON WITH RELATED SPECIES.—L. conjunctus closely re-
sembles ZL. organicus in general appearance, but a careful study shows
certain constant and characteristic differences. These are sum-
marized in table 1.

ASSOCIATION OF THE TWO LICHOMOLGIDS WITH THE ALCYONARIAN.—
Both LZ. organicus and L. conjunctus were recovered after washing
unbroken colonies of T'ubipora musica in sea water to which about 5
per cent ethyl alcohol had been added. The exact habitat of the
copepods is unknown, but there seems to be little doubt of their
association with the alcyonarian in view of the large numbers of
individuals recovered.

Family Clausidiidae Embleton, 1901

Genus Hippomolgus Sars, 1917
Hippomolgus latipes, new species
Figures 37-58

TYPE MATERIAL.—8 99 from a colony of Tubipora musica (Linnaeus),
in 1m, Pte. Ambarionaomby, Nosy Komba, near Nosy Bé, Madagas-
car, collected Mar. 31, 1964: holotypic @ (USNM 113395) and 4
paratypic 92, (USNM 113396) deposited in the United States National
Museum; remaining 3 99 (dissected) in the collection of A. G. Humes.

OTHER MATERIAL.—Also from T'ubipora musica: 1 9, in 1 m, Pte.
Ambarionaomby, Nosy Komba, June 8, 1964.

FrmaLe.—Body (fig. 37) with a rather harpacticoid appearance.
Length (excluding setae on caudal rami) 1.13 mm (0.97-1.19 mm)

No. 3573 COPEPODS—-HUMES AND HO a

and greatest width 0.32 mm (0.27-0.37 mm), based on 8 specimens.
Ratio of length to width of prosome 1.65:1. Segment bearing leg 1
completely fused with head. Lateral areas of segments of legs 1-3
not produced; those of segment of leg 4 slightly angular posteriorly.

Segment of leg 5 (fig. 38) broad, 94u x 242u, with fifth legs borne
posterolaterally. Between this segment and genital segment on
ventral side a weak intersegmental sclerite. Genital segment
(fig. 38) wider than long, 130u x 198u. In ventral view its lateral
border (fig. 39) with a small notch near middle of segment and a
more pronounced indentation posteriorly. Areas of attachment of
egg sacs situated dorsolaterally in midregion of segment. Each
area (fig. 40) bearing outwardly a rather hyaline flange and inwardly
a slender seta 11y long, a spiniform seta 8u long, and 2 small spinous
processes. On ventral surface of posterior half of genital segment a
transverse row of prominent spinules. Three postgenital segments
78u X 1835p, 60u x 118y, and 91u x 91y, respectively, from anterior to
posterior. Anal segment (fig. 41) bearing anteriorly 4 transverse rows
of spinules, 2 rows ventrolateral and 2 ventral ; along its posterior
border near insertion of each caudal ramus a row of small spinules,
this row continued on dorsal side.

Caudal ramus (fig. 42) 104u x 34y, or 3 times longer than wide.
Minute basal outer spinule 3u long. Outer lateral seta 22u long and
naked. Outermost terminal seta 41, long, naked, and composed of
2 parts: a sclerotized proximal half and a more slender distal portion.
Innermost terminal seta 57 long and haired along inner side. Two
long terminal setae 209u (outer) and 462u (inner), both basally
“pegged” and bearing lateral spinules. Pedicellate dorsal seta
394 long and naked. Dorsal surface of ramus with small hairs and
refractile points as indicated in figure.

Dorsal surface of prosome and dorsal and ventral surfaces of
urosome with scattered refractile points and hairs. Prosome slightly
longer than urosome, ratio 1.1:1.

In single ovigerous female collected egg sac (fig. 43) 319u x 107p,
extending a little beyond ends of caudal rami, with each ege about 40u
in diameter.

Rostral area (fig. 44) small and well sclerotized. Posterior to it,
between bases of second antennae, a small circular sclerotization.

First antenna (fig. 45) 6-segmented and relatively short and robust,
its length about 143. Lengths of segments measured along their
posterior nonsetiferous margins: 15u (39u along anterior margin),
19y, 28u, 18u, 134, and 24u, respectively. First segment with 5 setae,
proximal one lightly haired, distal one spiniform, with a row of
scalelike spinules along one edge. On second segment 2 similar
spiniform setae. All remaining setae naked. Each of last 3 seg-

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

ments with long prominent aesthete. Formula for armature: 5, 14,
9,4+1 aesthete, 2+1 aesthete, and 7+1 aesthete. Only ornamenta-
tion on first antenna consisting of crescentic row of spinules on first
segment.

Second antenna (fig. 46) 4-segmented, last segment 32 x 19» in
ereatest dimensions. Each of first 2 segments with a seta, third
segment with a spine and 2 slender setae on its inner distal expansion,
and last segment bearing 7 elements (a spine and 6 setae). Orna-
mentation indicated in figure, inner margins of third and fourth
segments having rows of scalelike spinules.

Labrum (figs. 47, 48) ornamented with spinules as indicated in
figures.

Mandible (fig. 49) bearing 3 terminal elements: a clawlike ventral
(outer) spine with 1-2 spinous processes on its concave edge, a spine
with lateral spinous projections on its basal half, and a slender barbed
seta. No paragnath. First maxilla (fig. 50) bilobed, with 8 setae
disposed in groups of 3, 4, and 1. Second maxilla (fig. 51) 2-seg-
mented; first segment unarmed, second with outer haired seta and 2
inner barbed spinulose setae and terminating in a short clawlike
process with at either side a setiform barbed process (these 2 processes
perhaps setae but their articulation not evident). Maxilliped (fig. 52)
4-segmented, first segment with inner barbed seta, second with 2
naked setae, third unarmed, and fourth (fig. 53) subterminally with
small naked seta and terminally with minute spiniform process and
fringe of spiniform ornamentations resembling a small fan.

Area behind mouth and between mouthparts (fig. 47) showing
anteriorly a transversely elongated median lobe (lingua) with a few
very minute lateral spinules, followed immediately by a transverse row
of hairs in 2 groups. Posterior to this a row of prominent spinules,
with at either end a cluster of hairs. Bases of maxillipeds joined by
sclerotized lines. Postoral area between maxillipeds and leg 1
broadly protuberant, especially noticeable in lateral view.

Legs 1-4 (figs. 54-57) with 3-segmented rami. Armature of legs
as follows (Roman numerals=spines, Arabic numerals=setae):

Pal protopod 0:1 de exp 1:0 ped III,1,4
end 0:1 0:1 13
pe protopod 0:1 1:0 exp 1:0 ee TIl,L5
end O:1 0:2 IES
P3 protopod O:1 1:0 exp 1:0 Gl LES
end O:1 0:2 II, 11,2
P4 protopod Os 1:0 exp 1:0 Tel TES
end O:1 0:2 Lie

Free edges of all 4 intercoxal plates with rows of long slender
setules. Inner margin of basis of leg 1 (fig. 54) bearing spinulose

NO. 3573 COPEPODS—HUMES AND HO 13

spine 25u long, with near its insertion an outer row of spinules and an
inner spiniform process with slightly bifurcated tip. In legs 2-4 this
margin more or less rounded and carrying only a row of hairlike
setules. Terminal segment of endopod of leg 1 with 1 spine and 3
setae. Spinous processes at each side of insertion of spine either
entire or bifurcated at tip. Ornamentation of legs indicated in figures.

Leg 5 (fig. 58) having an expanded free segment 83u x 53y in greatest
dimensions. Four elements borne on this segment consisting (from
dorsal to ventral) of a finely barbed spine 39u long, a naked seta 66u
long, a strong spinulose spine 44y long, and a short weakly barbed
spine 23 in length. Along dorsal edge of segment a row of spinules.
Seta borne on body dorsal to insertion of segment 40u long and naked.
(In dorsal or ventral view of undissected animal, segment of leg 5
appearing much less expanded than actually.)

Leg 6 probably represented by seta and spine near attachment of
each egg sac (see fig. 40).

Color in life unknown.

Matz.—Unknown.

Erymotoey.—The specific name latipes, from Latin, meaning
“having broad feet,’ refers to the wide free segment of leg 5 in this
species.

CoMPARISON WITH RELATED SPECIES.—Only a single species of the
genus Hippomolgus, H. furcifer Sars (1917), has been known until now.
This species was described on the basis of 3 females found free in 60
fathoms, muddy bottom, on the coast of Norway. Males of Hip-
pomolgus are unknown, although Nicholls (1944, p. 46) has expressed
the view that the male of Hersiliodes dubia Thompson and A. Scott
(1903) is in reality a Hippomolgus. This opinion was followed by
Krishnaswamy (1953), who reported males of Hippomolgus dubia from
the plankton of Madras. It seems improbable to us, however, that
Hersiliodes dubia is a Hippomolgus. The mandible, described by
Thompson and Scott as having ‘2 horizontal plumose projections
and 2 plumose setae,” is very different. Bocquet and Stock (1957)
think it probable that a new genus should be created for Hersiliodes
dubia.

Like Hippomolgus furcifer, H. latipes has a body form that is less
cyclopoid and more harpacticoid, has a relatively short 6-segmented
first antenna with a prominent aesthete on each of the last 3 segments,
lacks paragnaths, and has a generally similar structure of the second
antennae, mouthparts, and legs 1-5.

There are, however, important differences between these 2 species
(based of necessity on the female only). In H. latipes the first segment
of the first antenna does not bear a spine such as described in H.
Jurcifer. The mandible bears 3 terminal elements (2 spines and

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

1 seta) instead of 4, as in Sars’ figure (pl. 82, m). The maxilliped
is 4-segmented, the last segment ornamented with a small fanlike
fringe, instead of being 3-segmented as in Sars’ figure (pl. 82, mp’).
The armature of the last segment of the endopod of leg 1 is I,3
instead of I,II,3 as in Sars’ species. The free segment of leg 5 is
expanded instead of being narrow as in H. furcifer. The caudal
ramus is only 3 times longer than wide instead of about 13:1 as in
the Norwegian species. The length of the female is 1.13 mm,
instead of 1.40 mm as in H. furcifer.

Hippomolgus cognatus, new species
Figures 59-69

TYPE MATERIAL.—Holotype 2 (USNM 113397) from a colony of
Tubipora musica (Linnaeus), in 1 m, Pte. Ambarionaomby, Nosy
Komba, near Nosy Bé, Madagascar, collected Mar. 31, 1964,
deposited in the United States National Museum.

OTHER MATERIAL.—2 99 from Tubipora musica, in 1 m, Pte. a la
Fiévre, Nosy Bé, Jan. 5, 1964, 1 9 (USNM 113398) in the United
States National Museum, the other (dissected) in the private collection
of A. G. Humes.

FEMALE.—Body (fig. 59) resembling that of H. latipes. Length
(not including setae on caudal rami) 0.90 mm (0.82-0.95 mm) and
greatest width 0.29 mm (0.28-0.31 mm), based on 3 specimens.
Ratio of length to width of prosome 1.56:1.

Segment of leg 5 broad, 78u x 188u. Genital segment (fig. 60)
only slightly wider than long, 120u in length, 1324 wide in its an-
terior half, and 110 wide in its posterior half, where sides in dorsal
view are nearly parallel. Areas of attachment of egg sacs located
dorsolaterally on midregion of segment. Each area (fig. 61) with
a slender naked seta 22y long, a spine 8y long, and 2 spinous proc-
esses. Three postgenital segments 66y x 101, 60u x 91p, and 80u x 82y,
respectively, from anterior to posterior.

Caudal ramus (fig. 62) 64u x 31y in greatest dimensions, or 3 times
longer than wide. Outer lateral seta 25u, outermost terminal seta
39u, innermost terminal seta 88u, 2 long terminal setae 220y (outer)
and 450u (inner), dorsal seta 33y long.

Ratio of prosome to urosome 1.17:1.

In single ovigerous female collected egg sac (fig. 63) 3524 x 138u,
reaching nearly to ends of caudal rami, with each egg about 47u
in diameter.

Rostral area (fig. 64) bearing 2 rows of minute denticles.

First and second antennae like those in H. latipes. Labrum (figs.
65, 66) ornamented as indicated in figures. Mandible (fig. 67) bearing
3 terminal elements as in H. latipes, but these with different orna-

NO. 3573 COPEPODS—-HUMES AND HO 15

mentation, i.e., with spinous processes on both sides of 2 spines and
with seta naked. Paragnath absent. First and second maxillae
and maxilliped like those in H. latipes.

Area behind mouth and between mouthparts with median lobe
(lingua) weakly developed. Postoral area between maxillipeds and
leg 1 (fig. 65) less protuberant than in H. latipes.

Legs 1-4 with same spine and setal formula as in H. latipes, except
for endopod of leg 1 being 0:1; 0:1; 1,4. Leg 1 (fig. 68) with inner
spine on basis 22 long and adjacent spiniform process asymmetrically
bifurcated; terminal segment of endopod with 4 setae and with 3
small and rather obtuse processes between spine and first seta.
Legs 2-4 resembling those of H. latipes.

Leg 5 (fig. 69) with relatively slender free segment, 61 along inner
and ventral margin, 51y along outer and dorsal margin, and 22y
wide. Armature from dorsal to ventral consisting of spine 21y,
seta 50u, spine 25y, and spine 40 in length, all ornamented as in
figure.

Leg 6 probably represented by spine and seta near attachment of
each egg sac (see fig. 61).

In other respects similar to H. latipes.

Color in life unknown.

Matze.—Unknown.

Erymotocy.—The specific name cognatus, from Latin, meaning
“related or closely allied,” refers to apparent close relationship be-
tween this species and H. latipes.

CoMPARISON WITH RELATED SPECIES.—H. cognatus differs from
HI. furcifer Sars (1917) in lacking a spine such as described on the
first segment of the first antenna in the Norwegian species, in having
3 terminal elements instead of 4 on the mandible, in having a 4-seg-
mented maxilliped with a terminal fanlike fringe, in having the last
segment of the endopod of leg 1 armed as I,4 instead of I,IJ,3 as in
Sars’ species, and in having a relatively much shorter caudal ramus,
3:1 instead of 13:1 as in H. furcifer. The length of the female is
0.90 mm instead of 1.40 mm as in the species described by Sars.

H. cognatus apparently is closely related to H. latipes but differs
from it chiefly in the form of the genital segment, the presence of
denticles on the rostral area, the form and ornamentation of the
labrum, the ornamentation of the 3 elements on the mandible, the
armature of the last segment of the endopod of leg 1, and the form of
the free segment of leg 5.

ASSOCIATION OF THE TWO CLAUSIDIIDS WITH THE ALCYONARIAN.—
Whether Hippomolgus latipes and H. cognatus live in a close association
with Tubipora musica or occur only sporadically in the coral colony

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

is difficult to determine on the basis of the relatively few specimens
collected. (No Hippomolgus were found in 4 other colonies of
Tubipora musica.)

Literature Cited

BocaueEt, C., and Srocr, J. H.

1957. Copépodes parasites d’invertébrés des cétes de France, 1: Sur deux
genres de la famille des Clausidiidae, commensaux de mollusques:
Hersiliodes Canu et Conchyliurus nov. gen. Proe. Koninkl.
Nederlandsche Akad. Wetensch. Amsterdam, ser. C, vol. 60,
no. 2, pp. 212-222.

Brapy, G. 8.

1910. Die marinen Copepoden der Deutschen Siidpolar-Expedition 1901-
1903, 1: Uber die Copepoden der Stimme Harpacticoida, Cyclo-
poida, Notodelphyoida und Caligoida. No. 5 of vol. 11 (Zool.
vol. 3) in Deutsche Siidpolar-Expedition 1901-1903, pp. 497-593.

GURNEY, R.

1927. Zoological results of the Cambridge expedition to the Suez Canal,
1924, XX XIII: Report on the Crustacea: Copepoda (Littoral and
semiparasitic). Trans. Zool. Soc. London, vol. 22, no. 4, pp.
451-577.

Humes, A. G., and Frost, B. W.

1964. New lichomolgid copepods (Cyclopoida) associated with aleyonarians
and madreporariansin Madagascar. Cah. O.R.S.T.O.M. Océanogr.,
Sér. Nosy-Bé, vol. 2, no. 6, pp. 131-212.

KRISHNASWAMY, S.

1953. Pelagic Copepoda of the Madras coast. Journ. Madras Univ., vol.

23 B, no. 1, pp. 61-75.
NicHo.is, A. G.

1944. Littoral Copepoda from South Australia, II: Calanoida, Cyclopoida,
Notodelphyoida, Monstrilloida and Caligoida. Rec. South Aus-
tralian Mus., vol. 8, no. 1, pp. 1-62.

Sars, G. O.

1917. Clausidiidae, Lichomolgidae (part). Pts. 11 and 12 of vol. 6 (Co-
pepoda, Cyclopoida) zr An account of the Crustacea of Norway
with short descriptions and figures of all the species, pp. 141-172.

SEWELL, R. B. 8.

1949. The littoral and semi-parasitic Cyclopoida, the Monstrilloida and
Notodelphyoida. No. 2 of vol. 9 in John Murray Expedition
1933-34 Scientific Reports, pp. 17-199.

Stock, J. H.

1959. Copepoda associated with Neapolitan invertebrates. Pubbl. Staz.

Zool. Napoli, vol. 31, no. 1, pp. 59-75.
Tuompeson, I. C., and Scort, A.

1903. Report on the Copepoda collected by Professor Herdman, at Ceylon,
in 1902. Rep. Gov. Ceylon Pearl Oyster Fish. Gulf of Manaar,
suppl. rep., no. 7, pp. 227-307.

UmmerxutTry, A. N. P.

1962. Studies on Indian copepods, 5: On eleven new species of marine
cyclopoid copepods from the south-east coast of India. Journ.
Mar. Biol. Assoc. India, 1961, vol. 3, nos. 1 and 2, pp. 19-69.

ue

Figures 1-8.—Lichomolgus organicus, new species, female: 1, dorsal (A); 2, urosome, dorsal
(B); 3, area of attachment of egg sac, dorsal (C); 4, caudal ramus, dorsal (D); 5, egg
sac, dorsal (E); 6, rostral area, ventral (B); 7, first antenna, wi
Positions of aesthetes added in male, dorsal (F);
(Capital letters=scale at which drawn; a/=
tral area.)

th two arrows indicating
8, second antenna, posterior (F).
first_antenna, a’’=second antenna, r=ros-

iT

——
<FO

)
WING

Ficures 9-18.—Lichomolgus organicus, new species, female: 9, posterior part of labrum, with
paragnaths indicated by dashed lines, ventral (G); 10, mandible, posterior (G); 11, first
maxilla, posterior (G); 12, second maxilla, postero-inner (G); 13, tip of second maxilla,
postero-inner (D); 14, maxilliped, posterior (G); 15, terminal segment of maxilliped,
antero-inner (H); 16, postoral area between maxillipeds and leg 1, ventral (F); 17, leg 1
and intercoxal plate, anterior (F); 18, leg 2, anterior (F). (Capital letters=scale at which
drawn; mxpd= mazxilliped, Py=leg 1.)

18

Uh
(He

3

Ficures 19-27.—Lichomolgus organicus, new species, female: 19, leg 3, anterior (F); 20,
leg 4 and intercoxal plate, anterior (F); 21, leg 5, dorsal (D). Male: 22, dorsal (A); 23,
urosome, dorsal (B); 24, caudal ramus, dorsal (C); 25, second antenna, anterior (F); 26,
maxilliped, anterior (F); 27, endopod of leg 1, anterior (G). (Capital letters=scale at

which drawn.)
19

/

Oe
\
ee
Ficures 28-36.—Lichomolgus organicus, new species, male: 28, leg 5, dorsal (D); 29, leg 6,
ventral (G); 30, spermatophore, inside male, ventral (F). Lichomolgus conjunctus, new
species, female: 31, dorsal (A); 32, urosome, dorsal (B); 33, caudal ramus, dorsal (C); 34,

rostral area, ventral (B); 35, endopod of leg 4, anterior (F). Male: 36, dorsal (I). (Capital

letters=scale at which drawn; a’= first antenna, a’/=second antenna, r=rostral area.)
20

4 \< "iTpponrenRee \ I
May Wu

+ MOgqyHe DAN

-

Ficures 37-44.—Hippomolgus latipes, new species, female: 37, dorsal (A); 38, urosome,
dorsal (E); 39, right side of genital segment, ventral (F); 40, area of attachment of egg
sac, dorsal (G); 41, anal segment, ventral (F); 42, caudal ramus, dorsal (G); 43, egg sac,
lateral (1); 44, rostral area, ventral (F). (Capital letters=scale at which drawn Siae——fITSE
antenna, a’’=second antenna, Im= labrum.)

21

“iy fact Ye

Aw a
AMARASNND NAMA gg, 97” a

Ficures 45-53.—Hippomolgus latipes, new species, female: 45, first antenna, dorsal (G);
46, second antenna, posterior (D); 47, oral and postoral areas, with labrum erected,
ventral (G); 48, labrum, ventral (G); 49, mandible, posterior (C); 50, first maxilla, anterior
(D); 51, second maxilla, anterior (D); 52, maxilliped, anterior (D); 53, last segment of
maxilliped, ventral (H). (Capital letters=scale at which drawn; md= mandible, mx’=
first maxilla, mx’’=second maxilla, mxpd=maxilliped, pi=leg 1.)

22

Py
LPG GOOPS
SEER,

Seer

Mirna verarnianngw

Ficures 54-60.—Hippomolgus latipes, new species, female: 54, leg 1 and intercoxal plate,
anterior (F); 55, leg 2, anterior (F); 56, leg 3, anterior (F); 57, leg 4 and intercoxal plate,
anterior (F); 58, leg 5, lateral and outer (G). H. cognatus, new species, female: 59, dorsal
(I); 60, genital segment, ventral (F). (Capital letters=scale at which drawn.)

23

mi tn

Ficures 61-69.—Hippomolgus cognatus, new species, female: 61, area of attachment of egg
sac, dorsal (G); 62, caudal ramus, dorsal (D); 63, egg sac, lateral (E); 64, rostral area,
ventral (G); 65, oral and postoral areas, ventral (G); 66, free edge of labrum, ventral
(D); 67, mandible, posterior (C); 68, leg 1, anterior (G); 69, leg 5, ventrolateral (D).
(Capital letters=scale at which drawn; md= mandible, mx’= first maxilla, mx”’= second
maxilla, mxpd=mazxilliped; Pi=leg 1, a’ =first antenna, a’’=second antenna, In=
labrum.)

24

tl ¢ GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3574

COPEPOD CRUSTACEANS
PARASITIC ON TELEOST FISHES
OF THE HAWAIIAN ISLANDS!

By Avan G. Lewis?

Introduction

This is the third and last of a series of papers covering the copepod
parasites of Hawaiian fishes. The first (Lewis, 1964a) deals with the
caligoid copepod parasites of Hawaiian acanthurid fishes, the second
(Lewis, 1966) covers the copepod crustaceans parasitic on elasmo-
branch fishes, and the present paper deals with copepod crustaceans
parasitic on teleost fishes. An additional paper (Lewis, 1964b), deal-
ing with the caligoid genus Dentigryps, includes descriptions of two spe-
cies found on Hawaiian fishes.

Other than the papers mentioned above, the references to copepods
taken from Hawaiian teleosts are those of Nordmann (1864), de-
scribing Norion expansus and Peniculus calamus from unknown
hosts, and Edmondson (1946), who figures a species of Lernaeenicus
from dolphins. Bonnet (1948) lists some Hawaiian copepods taken
primarily from pelagic fishes, and Shiino (1963) describes Midias
lobodes, Caligus coryphaenae, and Brachiella thynni from fishes ex-
amined in the Honolulu Fish Market. In addition to these, Randall

1 This study was supported by grants (G-24956 and GB-2464) from the National
Science Foundation.

2 Associate Professor of Oceanography, Institute of Oceanography, University
of British Columbia, Vancouver, Canada.

1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

(1958) lists, by family, the copepods taken from the stomachs of
some parasite-picking fishes of the genus Labroides and (1961) the
parasitic copepods taken from the Manini (Acanthurus triostegus
sandvicensis).

The study here reported has benefited from collections made by
the Honolulu division of the U.S. Fish and Wildlife Service and
from the assistance given to the author by the Division of Marine
Invertebrates of the Smithsonian Institution, especially by Drs.
Bowman, Manning, and Cressey, whose assistance with the syn-
onymies is deeply appreciated. The author is also grateful to the
officers and crew of the U.S. Coast Guard vessel Buttonwood and to
Lester Zukeran and Samuel Kaolulo for the collection of both host
and copepod material.

Mernops.—The external surface, gill cavities, buccal cavity, and
nasal cavities of the teleost hosts were examined for parasitic copepods.
Copepods collected were killed in either 95 percent ethyl alcohol or
10 percent formalin and later transferred to 95 percent ethyl alcohol.
Specimens to be drawn or dissected were placed in 85 percent lactic
acid to clear and soften them, stained with Chlorazol Black E dis-
solved in 85 percent lactic acid, and then placed in benzyl alcohol for
final clearing and for dissection and drawing.

Drawings of the entire animal were made from specimens placed
in benzyl alcohol and covered with a cover slip, supported so that
the shape of the organism was not distorted. Both a camera lucida
and a Bausch and Lomb Tri-Simplex Micro-Projector were used in
making the drawings. ‘The appendages and processes were drawn in
situ. or were removed and mounted in either Hoyer’s mounting
medium or a 1:1 mixture of Turtox’s CMC-10 and CMC-S. Meas-
urements were made with an ocular micrometer on specimens softened
in lactic acid and held loosely in place by a cover slip supported
by spacers.

In the following figures the 9 and @ signs are used separately
under each drawing to indicate a difference between the appendage
or body part of the female and that of the male. The symbols are
used together (2 co’) to indicate the similarity of the appendage or
body part in both sexes. In the latter case the sex of the specimen
from which the drawing is made is indicated by a line under the
appropriate symbol. If only one sex is represented in the collection,
the symbols are not used.

TpRMINoLocy.—The term “cephalothorax” is used to indicate a
condition in which one or more of the thoracic segments are fused with
the cephalon. The maxilliped-bearing segment is considered as the
first thoracic segment. The term “pedigerous segment’ is used to
indicate a leg-bearing thoracic segment, while the terms “free thoracic

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 3

segments” or “free pedigerous segments” are used to designate those
thoracic segments that are not fused with the cephalon. The term
“genital segment” is used, although with some question, to designate
the fused sixth and seventh thoracic segments (=fifth and sixth pe-
digerous segments) in the Caligidea (Caligoidea of Yamaguti, 1963).
In the lernaeocerids, pennellids, and lernaeopodoids, the terms “ceph-
alothorax” and “trunk” are used. The term “cephalothorax”’ has
the same meaning as that given above, while the term “trunk” desig-
nates the region immediately posterior to the cephalothorax. Theterm
“ovigerous” is used to indicate a female with egg strings, the term
“nonovigerous” to indicate a female without egg strings even though
the genital segment (or trunk) may contain eggs.

With three exceptions, the terminology applied to the appendages
and processes is the same as that in Lewis (19640). Thus, the term
“antennules” refers to the first antennae and the term “antennae”
refers to the second. The term “mandibles” refers to the pair of ap-
pendages immediately adjacent to the mouth and normally projecting
into a “mouth cone” if it is present. The term “mazxillipeds” refers
to the modified pair of appendages on the first thoracic segment, while
the “sternal furca”’ is a bifurcate, posteroventrally directed projection
between and slightly posterior to the maxuliped bases of trebiids,
euryphorids, and caligids. The term “thoracic legs” designates the
pair of semifoliaceous or foliaceous appendages arising from some of
the thoracic segments posterior to the first or maxilliped-bearing seg-
ment, while the “caudal rami” are the pair of appendages at the
posterior end of the body.

The three exceptions to the original terminology have arisen from
a recently completed study made by the author on the nature of the
maxillae in the Caligidea (ms in preparation). It appears that the
setule- or seta-bearing node lateral to the mouth cone of this group
of caligoids is the maxillule, but it seems that the spine or plate imme-
diately behind this structure in the trebiids, eirgids, euryphorids, and
caligids is an accessory formation and not part of the maxillule. The
name “maxillule” is now given to the setule-bearing node (maxillary
palp of Yamaguti, 1963), while the term “postoral process” refers
only to the spine or plate immediately behind the maxillule. The
term “maxillae” applies to the second pair of maxillae with its asso-
ciated pair of openings to the maxillary glands. The term “post-
antennal process” is used to designate the process-bearing nodules and
the spinelike projection, if present, situated lateral and slightly
posterior to the base of the antenna.

To facilitate the use of the thoracic leg tables, a hypothetical tho-
racic leg is shown in figure 1, giving the various thoracic leg armature

4 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

elements used in the tables. Further, an analysis of the hypothetical
thoracic leg is given in table 1.

In the section entitled “Material” for each of the species, the
abbreviation USFWS means U.S. Fish and Wildlife Service, HMS
means the USFWS vessel Hugh M. Smith, CHG the USFWS vessel

pH Gh

1 mh mH
Ww

fe

S

Lit
= ulus
Trey stagih dre hye

Ficure 1.—Hypothetical thoracic leg showing various armament components:

a: adhesion surface IP: interpodal plate

C: heavy fringing plumosities m: membrane

c: light fringing plumosities P: plumose seta

D: large denticulations p: plumose setule

d: small denticulations P’: naked seta

dH: large denticulated spine p’: naked setule

dm: denticulated membrane pH: large plumose spine

dmH: large spine with denticulated mem- Q: seta plumose on one side and mem- |
brane branous on other

fm: frilled membrane gH: large spine plumose on one side and

fmH: large spine with frilled membrane membranous on other

H: large spine rh: spinule

h: small spine s: solitary hairlike process

T,pie 1—Armature of hypothetical thoracic leg shown in figure 1

Inter- | Protopodite Exopodite | Endopcdite
Leg | Surface podal |
Plate
1 2 1 2 | 3 1 2 3
Outer fm,p rh,dm, {mi H,h,mH, da,D, e id. pipiens
m dmii mh,dH Cc 12
Inner se) | mja;s) |e c,P ¢,Q,qH,pH | P c,2P epee!

col Ne ee

NO. 3574 COPEPOD CRUSTACEANS—LEWIS Oo

Charles H. Gilbert, JRM the USFWS vessel J.R. Manning, and LL
means a USFWS longline cruise on board a chartered vessel.

The names of the Hawaiian fishes that served as hosts are taken
from the “Handbook of Hawaiian Fishes” by Gosline and Brock
(1960). The names of previously reported hosts are as they appear
in the references giving these hosts.

A list of the Hawaiian fishes examined is given following the de-
scriptions. Included in this list are the species of copepods found
on each of the hosts.

Order Cyclopoida
Family Bomolochidae
Pseudotaeniacanthus Yamaguti and Yamasu, 1959

DraGcnosis.—Body cyclopoid, cephalon fused with first 2 thoracic
segments, slightly broader than rest of prosome; second to fifth pe-
digerous segments gradually narrowed posteriorly, projecting laterally
over respective protopodites; fifth pedigerous segment not appre-
ciably expanded laterally. Abdomen 4-segmented. Antennule 5-
or 6-segmented; antenna uniramous, with 4 clawlike terminal spines.
Postantennal processes present, fused, Y-shaped, with rows of spi-
nules giving brushlike appearance, with or without large, posterior
hooks at apex of Y. Mazxillule setiferous; maxillae as in other bom-
olochids or reduced; maxilliped clawlike in male, reduced in female.
Thoracic legs 1-4 biramous, rami 3-segmented; endopodite of first
leg longer than exopodite, with few marginal setae; fifth and sixth
legs uniramous.

Pseudotaeniacanthus puhi, new species
Figures 2, 3

MarTeErRIAL.—One adult female (holotype, USNM 112862), 1 adult
male (allotype, USNM 112863), 12 adult females, and 8 adult males
(paratypes, USNM 112864) from the gill cavity of a ‘Brown Moray”?
from the Honolulu Aquarium. One adult female and 1 adult male
(paratypes, USNM 112865) from the gill cavity of a “Green Moray”
from the Honolulu Aquarium.

MEASUREMENTS.—(In mm) 14 females and 10 males:

female male

Greatest length, excluding mean range mean range

caudal setae 1.15 0.92-1.27 1.05 0. 92-1.18
Length of prosome 0.78 0. 68-0.88 0.66 0. 54-0. 74
Width of prosome 0.39 0.35-0.42 0.37 0. 32-0. 42
Length of cephalothorax 0.30 0.27-0.32 0.32 0. 28-0. 37
Length of genital segment 0.10 0.09-0.12 0.11 0. 08-0. 14
Width of genital segment 0.14 0.13-0.15 0.14 0.13-0.16

Length of egg sac (3 sacs) 0. 16, 0. 18, 0. 48

6 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

Descrietion.—Female cephalothorax (fig. 2a) bell shaped, consist-
ing of cephalon, maxilliped-bearing, and first pedigerous segments.
Second pedigerous segment distinct from and slightly narrower than
cephalothorax; third pedigerous segment distinct from and slightly
narrower than second; fourth pedigerous segment distinct from and
slightly narrower than third, broadly rounded posteriorly. Fifth
pedigerous segment approximately half the width of fourth, anterior
end overhung by posterior end of fourth pedigerous segment.
Genital segment (fig. 2c) distinct from fifth pedigerous segment, lateral
margins convex; oviducal opening surrounded by series of small
sclerites. Remaining urosome segments 4 in number, each tapered
slightly, narrower and shorter than preceding segment. Caudal ramus
(fig. 2f) subrectangular, short, with 2 long and 3 short terminal setae.

Male prosome (fig. 2b) more ovoid than that of female, general
makeup otherwise similar. Urosome tapered, genital segment (fig.
2d) similar in shape to remaining 4 urosome segments although
larger.

Female and male antennule (fig. 27) 5- or 6-segmented, proximal
segment expanded on proximal posterior surface. Female and male
antenna (figs. 21, m) uniramous, 3-segmented, situated posterior and
slightly lateral to antennule base; first segment approximately one
and one-half times the length of second, both segments without arma-
ture. Third segment elongate, length slightly less than that of first
segment, proximal portion with folded, flaplike extension with dentic-
ulations, naked seta and large, spinelike projection distally (fig. 2m)
Distal end of third segment with 1 naked seta and 4 clawlike spines.

Female and male mandible (fig. 27) rodlike, 2-parted. First part
tapered slightly from proximal to distal end, second part tapered to
pointed distal end, curved inward slightly distally. Female and
male with Y-shaped postantennal process on anterior ventral surface
(fig. 27), arms of Y with 9-10 transverse rows of spinules giving
brushlike appearance. Female and male maxillule (fig. 27) situated
immediately posterior to mandible base, consisting of pair of setae
attached to platelike area of heavy sclerotization contiguous with
posterior edge of platelike area of heavy sclerotization forming attach-
ment surface for mandible. Maxilla (fig. 24) represented by large,

Ficure 2.—Pseudotaeniacanthus puhi, new species, dorsal view: a, female; b, male. Fifth
pedigerous segment, fifth leg, genital segment, sixth leg, and first abdominal segment: c, fe-
male; d, male, same, also includes anterior end of first abdominal segment. Right fifth
leg: e, posterior view. Ventral view: f, caudal ramus; g, female cephalothorax (mdbl=
mandible, mxpd=maxilliped) ; h, male cephalothorax (mxpd=maxilliped); i, right antennule
and heavily sclerotized postantennal process; j, right mandible (mdbl), maxillule (ma-l),
and paragnath (pg); k, maxilla; 7, antenna; m, projection on proximal portion of third
segment of antenna.

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

Or
“ol
E
E
wo
Q
Oo
y™!

0.03 mm.2c

8 PROCEEDINGS OF THE NATIONAL MUSEUM vou. 121

heavily sclerotized, knoblike projection posterior and lateral to oral
region. Knob with broad base, tapered to narrow distal region with

a
ye ee 005mm. #¢
sean .

oO

oO

ol

5

2
+O
10,
NALA)

SS Stith ah
== pS

a: J p ie
7 irr AT Fr . = )
gM" VT #2 ge

ps /

+
005mm. 3c
¢

Ficure 3.—Pseudotaeniacanthus puhi, new species: 4a, male maxilliped, ventral view.
Thoracic legs: J, left first, anterior view; ¢, left, second, posterior view; d, right third
posterior view; é, right fourth, posterior view.

medially facing cuplike indentation, indentation bearing single, seta-
like process. Female and male paragnath (fig. 27) minute, situated

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 9

immediately posterior to maxillule, consisting of single, flabby,
subconical projection bearing pair of setules distally.

Female maxilliped (fig. 2g) knoblike, broad based, distally concave,
without armature. Male maxilliped (fig. 3a) 2-segmented, first seg-
ment large, strongly developed, narrow proximally, broad distally,
with 3 small, lappet-like projections along inner surface and single,
large, distally recurved projection from distal inner surface. Second
segment fused with clawlike terminal process, bearing single, acces-
sory seta proximally.

For nature of legs and armature, see figures 2c-e, 3b-e, and table 2.

TABLE 2.—Armature of thoracic legs 1—4 of the female and male of
Pseudotaeniacanthus puhi, new species

Tee Nsaeracs Bee Protopodite Exopodite Endopodite
Plate
ree |e 1 2 3 1 2 3
1 | Outer Dp iE 9P c c ¢,3P

Inner 1 P Dp p 2P’

a | Outer a foe D,pH*| D,pH*| D,pH*,D,2pH| 2D | 2D | 2D,P,2D,2P
Inner Py c iP 5P iP 2P 2P

“qt | Outer Z oral ea D, pH | D,pH | D,pH,D,2pH| C,D | ©,D | ©, D,P,2D,
Inner rh c iP 5P iP 2P 22

“Iv | Outer D |P |D,pH | D,pH | D,pH,D,2pH|2C0 | C,D | C,2D,2pH
Inner ” rh 4 c 12 5P iP le 12

*Present on female, male=H.

Discussion.—Pseudotaeniacanthus puhi differs from P. congeri
Yamaguti and Yamasu, 1959, primarily in not having the large ‘‘pos-
terior hooks” of the latter species, in the armature of the thoracic legs,
and in the smaller thoracic pleura. The species name is derived
from ‘‘Puhi,’”’ the Hawaiian name for moray eels, the host of the
species.

Anchistrotos Brian, 1906

Draanosis.—Body cyclopoid, cephalothorax consisting of cepha-
lon and first 2 thoracic segments. Cephalothorax the broadest
part of prosome, 3 free prosomal segments tapering posteriorly.
Small frontal region present, with rostrum. Abdomen 8- or 4-seg-
mented; caudal rami short. Antennule 6- or 7-segmented, proximal
segments not fused; antenna 3-segmented, distal segment with pecti-
nate margin, bearing terminal claws and setae. Mandible 2-seg-
mented, second segment with terminal blade and subterminal palp.
Postantennal process well developed, clawlike. Maxillule nodular or
conical, typically with 5 associated setae or setules; maxilla tipped

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

with pectinate blade, with 1-2 subterminal setalike or palplike proc-
esses. Maxilliped with or without claw. Thoracic legs 1-4 bira-
mous, first with 2- or 3-segmented rami; second to fourth with 3-
segmented rami. Fifth legs uniramous, 2-segmented; sixth legs
setiform or lacking.

Remarks.—Anchistrotos is quite similar to Taeniacanthus; the only
consistent difference appears to be that the length of the caudal rami
of Anchistrotos is shorter than that of Taeniacanthus. The variation in
this characteristic, however, is such that no distinct relationship can
be stated. Yamaguti (1963) uses the presence of a “rostral projec-
tion at center of frontal margin’ as a key characteristic to separate
Anchistrotos from the remaining taeniacanthins, but this projection
is not present in A. occidentalis Wilson, 1924, and is not apparent from
the figures of several other species of the genus. Additionally, the
frontal region of members of Anchistrotos and Taeniacanthus is similar.
Wilson (1911a, p. 387) states that “the three basal joints [of the anten-
nule are] thoroughly fused . . .” in the diagnosis of the genus
Taeniacanthus. If this characteristic had been used in later descrip-
tions (e.g., Taeniacanthus sebastichthydis Yamaguti, 1939a), it would
have provided a better means of separating the two genera.

The following species is included in the genus Anchistrotos because
it does have a slight rostral projection, the caudal rami are “short,”
the proximal 3 segments of the antennule are not fused, and, in
general, the other characteristics most closely approximate those of
the species previously described for the genus.

Anchistrotos moa, new species
Fiaures 4, 5

MateriaL.—Two adult, nonovigerous females (holotype, USNM
112866; paratype, USNM 112868) from a specimen of Ostracion
lentiginosus captured at Rabbit Island, adjacent to Oahu, Hawaii.
Two adult nonovigerous females (paratypes, USNM 112867) col-
lected from a second specimen of O. lentiginosus captured at Rabbit
Island. All 4 specimens were given to the author without information
concerning their location on the host.

MEASUREMENT.—(In mm) 4 females:

mean range

Greatest length, excluding caudal setae 0. 76 0. 70-0. 85
Length of prosome 0. 57 0. 52-0. 62
Width of prosome 0. 40 0. 34-0. 43
Length of cephalothorax 0. 33 0. 30-0. 36
Length of genital segment 0. 07 0. 06-0. 07
Width of genital segment 0.138 0. 12-0. 14
Length of egg sac (1 sac) 0. 55

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 11

DESCRIPTION OF FEMALE.—Body (figs. 4a, 6) of general taenia-
canthiform nature. Cephalothorax consisting of cephalon, maxilliped-
bearing, and first pedigerous segments; second through fourth
pedigerous segments distinct. Frontal region (fig. 4e) narrow, un-
armed except for small, slightly projecting rostrum. Cephalothorax
projecting ventrally well below ventral surface, extending posteriorly
over lateral anterior surface of second pedigerous segment, with
irregular, heavily sclerotized band across dorsal surface of posterior
region. Second, third, and fourth pedigerous segments decreasing in
width, with ventrally and posteriorly projecting extensions as on
cephalothorax except not as well developed. Fourth pedigerous
segment rounded posteriorly, extending over dorsal surface of fifth
pedigerous or first urosomal segment. Genital segment (fig. 4c)
indistinctly swollen, slightly larger than fifth pedigerous segment;
sixth legs present as pair of very lightly plumose setae and spinule-
like projection immediately posterior to oviducal openings. Abdomen
3-segmented, segments diminishing in length and width, from dorsal
viewpoint. Ventral surface of third abdominal segment with 2 rows
of minute spinules anteriorly, separated by pair of larger spinules
medially, and incomplete row of spinules in region of attachment of
caudal rami (fig. 4c). Caudal rami short, slightly longer than wide,
with 2 long, lightly plumose median setae, 2 short outer setae, and 1
inner setule on distal surface.

Antennule (fig. 4f) 7-segmented although segmentation not com-
pletely distinct, first and second segments and third and fourth
segments appearing partially fused. First segment with 5 lightly
plumose setae along anterior surface; second with 11 on anterior, 3
on dorsal surface; third segment with 4 or 5 plumose setae anteriorly;
fourth with 3; fifth with 4; sixth with 3 naked setules; seventh with
one naked setule from distal posterior surface, 3 small, naked setules
from posterior distal surface, 3 long, naked setules and one aes-
thete(?) from anterior distal surface. Antenna (figs. 4g, h) uniramous,
3-segmented; first segment tapered slightly toward distal end, with
slight knob on outer distal corner. Second segment slightly more
than half the length of first, with small spine on distal inner surface;
third segment 1% times the length of second, with large, elongate
projection on inner surface, extending from distal end of segment
proximally, past proximal end to form cup-shaped projection over-
lapping distal half of second segment. Additional irregularly awl-
shaped projection present on distal ventral surface, projecting past
distal end of segment. Both projections with complete marginal
denticulations. Additional armature of third segment consisting of
3 naked setules and 4 clawlike spines distally.

PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

12

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 13

Mandible (fig. 5a) 2-segmented, first segment slightly longer than sec-
ond, second bearing 2 short, flattened, pectinate processes distally,
ventralmostlongest. Postantennal process (fig. 5b) well developed, con-
sisting of spinelike falciform projection slightly posterior to antennule
base and directed posteroventrally; base heavily sclerotized and en-
larged, tip sharply pointed. Maxillule (fig. 5a) nodular, situated im-
mediately posterior to mandible base, bearing 2 long, very lightly plumose
setae, 1 shorter naked seta and 2 minute setules. Maxilla (figs. 5a, ¢)
with one distinct segment although indication of second segment fused
with cephalothorax; armature of distinct segment consisting of 2spikelike
projections, one subterminal and very lightly plumose, second terminal,
finely denticulated along inner margin. Paragnath (fig. 5a) lobate,
curved inward sharply, with slightly fuzzy distal end.

Maxilliped (fig. 5d) 2-seemented although indication of third seg-
ment suggested by heavily sclerotized, platelike process intimately
associated with cephalothorax but forming articulation surface for max-
illiped. First segment elongate, with 1 or 2 naked setae arising from
slight depression of median posterior surface. Second segment with
long, indistinctly separable falciform process, distal end of segment
with nodular swelling bearing 1 or 2 naked, setule-like accessory
processes.

For nature of legs and armature see figures 4c, d, 5e-h, and table 3.
TABLE 3.—Armature of thoracic legs I-V of the female of Anchistrotos moa, new

species
Inter- |Protopodite Exopodite Endopodite
Leg | Surface || podal
Plate} 1 | 2 1 2 3 Tesleee 3
' Outer a ce 2H 2H c
Inner 2P,d 6P IP oe
II | Outer Fp Che dm,H dm,H | 2dm,3H dm,3H
Inner C 1 4P PACs SEAC aro:
III | Outer E C,P dm,H dm,H | 2dm,3H dm,3H
Inner cic iP 4P PAC | Se @) Ze
IV | Outer CAE. dm,H dm,H | 3H dm | 2dm,3H
C
Inner iB 4P iPeCalees€
V | Outer Cres dm,2H, dm, P, dm,
HCx

*Designation under protopodite and exopodite for convenience; association of segment uncertain.

Ficure 4.—Anchistrotos moa, new species, female: a, dorsal view; BD, lateral view; c, genital
segment, sixth legs, abdomen, and caudal rami, ventral view; d, fifth leg, ventral view;
é, rostrum, frontal region and proximal segment of antennules, dorsal view; f, right anten-
nule, dorsal view (setal plumosities not figured); g, antenna, ventral view; h, third segment
of antenna, dorsal view.

14

PROCEEDINGS OF THE NATIONAL MUSEUM

VOL. 121

NO. 3674 COPEPOD CRUSTACEANS—LEWIS 15

Discussion.—Anchistrotis moa is most closely related to A. ostra-
cionis (Richiardi, 1870). The overhanging tergal regions of the
cephalothorax and prosomal pedigerous segments of A. moa, the
absence of the small basal projection found on the postantennal
process of A. ostracionis, and the nature of the spination on the
ventral surface of the third abdominal segment are the most readily
distinguished characteristics which separate the two species. The
species name is derived from ‘‘Moa,” one of the Hawaiian names for
Ostracion lentiginosus, the host of this copepod.

Order Caligoida

Family Dichelesthiidae
Hatschekia Poche, 1902

DiaGnosis OF FEMALE.—Cephalothorax consisting of cephalon
and first thoracic segment; distinct although incompletely separated
from second thoracic segment. At least second and third thoracic
segments pedigerous, segments either distinct or fused. Fourth
and fifth thoracic segments fused with genital segment. Abdomen,
if distinct, short, 1-segmented; frequently fused with genital segment;
caudal rami minute. Antennule 3- to 7-segmented, antenna 2- or
3-segmented, terminal segment with claw. Mandible rodlike, den-
ticulated distally. One pair of maxillae apparently lacking; maxillules
or maxillae (probably maxillules) 2-parted or biramous (depending on
degree of reduction), each part (if rudimentary) or ramus (if well
developed) typically with 2 setae or spines. Maxilliped slender,
terminal process typically bifurcate. First 2 thoracic leg pairs
biramous, rami 1- or 2-segmented; third and fourth thoracic Jeg
pairs rudimentary or absent; fifth pair absent.

DraGNosis OF MALE.—Cephalothorax distinct from second thoracic
segment, postcephalothoracic segments incompletely or completely
fused. Abdomen, if distinct from genital segment, 1-segmented;
caudal rami larger than in female, armed with spines or stiff setae.
Appendages similar to those of female.

Hatschekia breviramus, new species

Figures 6, 7

Mareriau.—Thirteen adult females from the external surface of
Mulloidichthys auriflamma Forskil collected at Kauai, Hawaii. One

Figure 5.—Anchistrotos moa, new species, female, ventral view: a, oral region, right side
showing labrum (lab), mandible (mdbl), maxillule (ma-1), maxilla, and paragnath (pg);
b, antennule base (a-1) and postantennal process; c, projecting portion of maxilla; d, max-
illiped. Thoracic legs, anterior view: e, right first; f, left second; g, left third; A, right
fourth.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

of these females has been designated as the holotype (USNM
112870), the others as paratypes (USNM 112869).
M£EASUREMENTS.—(In mm) 10 females:

mean range
Greatest length of body 0. 67 0. 61-0. 73
Length of cephalothorax 0.18 0. 16-0. 20
Width of cephalothorax 0. 29 0. 24-0. 33
Length of trunk 0. 56 0. 49-0. 62
Width of trunk 0. 23 0. 20-0. 25
Length of egg strings (5 strings) 0. 44 0. 36-0. 52

DESCRIPTION OF FEMALE.—Body (fig. 6a) divisible into two regions.
Cephalothorax, including cephalon and maxilliped-bearing thoracic
segment, forming first region, pedigerous segments and genital segment
forming second. Cephalothorax of general ovoid shape, margins
slightly irregular. Anterior dorsal region of cephalothorax heavily
sclerotized, with cross-shaped region of sclerotization extending pos-
teriorly on median dorsal surface to middle of cephalothorax although
fine line of heavy sclerotization extending posteriorly to posterior-
median dorsal surface. Second region composed of fused pedigerous
and genital segments, constricted anteriorly into necklike connection
with cephalothorax, constriction composed anteriorly of 2 dorsally
fused pedigerous segments. Posterior two-thirds of second region
tapered in some specimens to posterior end although shape variable,
depending at least partially on number of eggs in genital segment.
Posterior lateral and posterior ventral surfaces of second region irregu-
lar, knobby, knobs forming margin of oviducal openings and more
apparent in ovigerous females. Posterior end rounded, projecting past
oviducal openings in some specimens, bearing knoblike caudal rami on
lateral ventral surface. Caudal rami (fig. 6b) small, each tipped
by 2 small setules.

Antennule (fig. 6c) 6-segmented, strongly compressed. First
sepment with 6 irregular naked setae from ventral margin; second
segment with 3 setae; third with 1 from ventral margin, 1 from distal,
and third from dorsal surface; fourth segment naked; fifth with one
seta from dorsal surface; sixth with 7 setules distally. Antenna (fig.
6d) 2- or 3-segmented, basalmost portion squat and irregular, segment-
like but questionable, with several knoblike articulation surfaces
received by proximal end of penultimate segment. Penultimate

Ficure 6.—Hatschekia breviramus, new species, female: a, dorsal view; J, posterior region,
left side showing caudal ramus (r) and anal indentation (a); ¢, right antennule, anterior
view; d, left antenna, ventral view; e, membrane-like flap at base of antenna, ventral view;
f, oral region, left side showing mouth cone, mandible (mdbl), and maxillule (ma); g, oral
region showing mouth cone, mandible, maxilla?, antenna base (a-2), and maxilliped base
(mxpd); h, maxilliped, ventral view; 7, distal end of maxilliped, ventral view.

17

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

— 0.03rmm.

226-321—67——_2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

segment elongate, broader proximally than distally, posterior and me-
dial surfaces rugose. Distalmost segment short, subspherical, fused
with clawlike terminal process, with very fine, hairlike accessory
process (not shown in figure). Distinct flaplike projection present on
heavily sclerotized cephalothoracic depression associated with an-
tenna base (figs. 6d, e). Mandible (fig. 6f) appearing 3-parted although
divisions indistinct; appendage rodlike, curved inward distally, distal
surface composed of 3 small denticulations. Maxillule? platelike,
projecting posteriorly and slightly ventrally from just lateral and
posterior to mouth cone base, irregularly pointed distally, with single
setule from distal surface.

Maxilliped (figs. 6h,7) 3- or 4-segmented, basalmost portion appear-
ing as segment-like projection of cephalothorax. Projection concave
distally, forming articulation surface for first distinct segment; segment
slightly wider proximally than distally although proximal end tapered
sharply to narrow proximal surface. Penultimate segment slender,
proximal end with knob-shaped articulation surfaces, distal end with
hairlike process. Distalmost segment short, heavily sclerotized, with
bifurcate terminal process. Terminal process with both tines sharply
pointed, innermost slightly less than twice the length of outermost.

First thoracic legs with 2-segmented exopodite, 1-segmented
endopodite; second thoracic legs with 2-segmented exopodite and
endopodite. Interpodal plates heavily sclerotized. For nature and
armature of legs see figures 7a, 6, and table 4.

b

0.03mm. 0.03mm.

Figure 7.—Hatschekia breviramus, new species, female, thoracic legs: a, first; b, second.

Discusston.—Hatschekia breviramus has characteristics which can
be compared most closely to 5 other members of the genus:
1. H. diodontis Yamaguti, 1953: Similar in general body shape,

No. 3574 COPEPOD CRUSTACEANS—LEWIS 19

TaBLe 4.—Armature of thoracic legs I and II of the female of Hatschekia brevira-
mus, new species

Exopodite Endopodite
Leg Surface Interpodal | Protopodite
Plate 1 2 1 9
I | Outer p’ p’ p’
er p’ p’ p’
II | Outer pe Pe Pe
er p p’ 2p’

distinct principally in nature of armature of antennule and thoracic
legs.

2. H. ostracu Yamaguti, 1953: Similar in body shape and in
the presence of the processes at the base of the antennae. JH. brevi-
ramus differs from this species in having 6, not 7, segments in the
antennule, the processes at the base of the antennae are not as large,
the caudal rami are tipped by 2 setules, while there are 5 in H. ostra-
ev. The endopodite of the first thoracic leg is 2-segmented in H.
ostracii and 1-segmented in H. breviramus; H. ostracii also has a
rudimentary third thoracic leg which is not present in H. breviramus.

3. H. cadenati Nunes-Ruivo, 1954: The thoracic leg armature
is similar although the caudal rami of H. cadenati, in addition to
other body parts, differ, having 5 setae and setules instead of 2 as in
H. breviramus.

4. H. bodiani Nunes-Ruivo, 1954a: This species, like H. ostracii
Yamaguti, 1953, has the projections at the base of the antennae that
are present in HZ. breviramus. The maxilliped armature is also similar
but the 2 species differ, most distinguishably in the armature of the
caudal rami and thoracic legs.

5. H. cluthae (Scott, T., 1902): Similar in reduced armature of
first thoracic leg, differs firstly in having 2-sezmented endopodite on
the first thoracic leg, secondly in having a more distinct abdomen,
different armament on maxilliped, and general body shape.

The species name “breviramus’’ is derived from the Latin words
“brevis” (short) and ‘‘ramus”’ (branch) and refers to the short caudal
ramus.

Family Lernaeoceridae
Peniculus Nordmann, 1832

Dracnosis.—See Lewis, 1964a.

Peniculus calamus? Nordmann

Peniculus calamus Nordmann, 1864, p. 5, no figures.
Peniculus calamus?—Lewis, 1964a, p. 233, fig. 24. [For bibliography, see Lewis
1964a.]

20 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

Hosts AND DISTRIBUTION.—S host records:

location hosts reference
Hawaiian Islands Unknown Nordmann, 1864
Acanthurus dussumieri Lewis, 1964a
A. mata

A, olivaceus

A. triostegus sandvicensis
Ctenochaetus strigosus
Naso hexacanthus

N. unicornis

MarTertAu.—One ovigerous female collected by D. Watson from
the dorsal fin of Pervagor spilosoma (Lay and Bennett) from the
Honolulu Aquarium.

MEASUREMENTS.—Length from anterior end of head to posterior
end of abdomen, excluding projecting second antennae, 2.89mm.
Length of head 0.41mm; greatest width 0.28mm. Length of neck
0.22mm; greatest width 0.13mm. Greatest length of combined
fourth pedigerous segment, genital segment, and abdomen 2.15mm.
Length of egg strings 2.81 and 2.96mm.

Derscription.—See Lewis, 1964a.

Family Pennellidae
Pennella Oken, 1816

Dracnosis.—Female: Body heavily sclerotized, separable into
cephalothorax, neck, and trunk (including abdomen). Cephalothorax
suborbicular, consisting of cephalon and at least first thoracic segment;
oral region at or near anterior end, surrounded by one or more types of
papillae. Two to three heavily sclerotized horns present on posterior
region of cephalothorax, at junction of cephalothorax and neck, or on
anterior end of neck; horns of variable length and shape, usually
projecting laterally or posterolaterally. Neck slender, cylindrical,
distinct from cephalothorax due to size difference, usually continuous
with trunk. Trunk elongate, cylindrical or slightly flattened dor-
soventrally, with ventral indentation at genital openings and frequently
with annulations along entire length. Abdomen forming posterior part
of trunk, behind ventral indentation at genital openings, tapered
posteriorly; posterior surface bilobed due to anal indentation, bearing
minute caudal rami. Numerous branched or simple filiform processes
arising from lateral or ventral and ventral lateral surface of abdomen,
giving plumose appearance to posterior part of body. Egg strings
long, frequently more than twice the lengthof body. Antennule2-to5-
segmented, situated on dorsal surface of cephalothorax, behind an-
tennae. Antennae chelate, 2- or 3-segmented (movable portion of
claw designated as segment with terminal process). Mandibles of

No. 3574 COPEPOD CRUSTACEANS—LEWIS pal

unknown nature; maxillules, maxillae, and maxillipeds present in late
stages of development but absent in adult. Four pairs of thoracic
legs present although poorly developed; first 2 pairs in close proximity
on anterior ventral surface of trunk, third and fourth pairs removed
variable distance behind second pair. First 2 pairs biramous, third
pair typically uniramous, fourth pair probably always uniramous.

Remarks.—The third and fourth thoracic legs are described as
uniramous in the genus Pennella (Wilson, 1917; Leigh-Sharpe, 1928;
Yamaguti, 1963). The third leg of one of the species here described is
biramous. The difference may be interspecific in nature although
the rami of any of the thoracic legs of the adult female are brittle and
easily broken.

Pennella histiophori? Thomson, 1889

Fiagurss 8, 9

Pennella histiophori Thomson, 1889, p. 368, pl. 38, fig. 2—Bassett-Smith, 1899, p.
483.— Wilson, 1917, p. 113 (key) — Yamaguti, 1963, p. 208.

Pennella zeylanica? Kirtisinghe, 1932, p. 137, figs. 1-5.

Pennella instructa? Kirtisinghe, 1964, p. 110, fig. 153.

Hosts AND DISTRIBUTION.—2 host records:

location hosts reference
New Zealand Histiophorus herschelli Thomson, 1889
Ceylon H. gladius Kirtisinghe, 1932, 1964

MatertaL.—One complete, ovigerous female, 1 nonovigerous female
without cephalothoracic papillae, and 1 incomplete female (trunk and part
of neck missing), in addition to part of larval exuvium, collected by D.
W. Strasburg from external surface of Makaira audax (Philippi) at
9°34’S, 1386°45’W (USNM 112871).

Remarxs.—Although the host was collected far from the Hawaiian
Islands, the wide distribution of the host and its common occurrence
around Hawaii suggested the inclusion of the copepod in this treatise.

MASUREMENTS.—(In mm) 2 females:
Total length (1 specimen) 172
Length of cephalothorax, excluding papillae le

8, 8
(

Width of cephalothorax
Length of neck

Width of neck 3, 2

Length of trunk 86, 89

Width of trunk 655

Approximate length of abdominal plumosities 25, 28

Length of attachment horns Give 10), 342312)
Length of egg string (1 string) 178

Description.—Female: Cephalothorax (figs. 8a, 6) presumably
consisting of cephalon and first thoracic segment although maxillipeds

22 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

-

i 0.5mm. §

Ficure 8.—Pennella histiophori? Thomson, 1889, female: a, dorsal view; b, cephalothorax,
dorsal view (oc=ocular region); c, posterior end of abdomen, dorsal view showing caudal
rami and anal indentation; d, genital region, ventral view showing egg strings (es); ¢, part
of one filamentous posterior process showing connection with gut (g); f, cephalothorax,
anterior view showing oral region; g, lateral view of some papillae in oral region.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 23

not visible. Cephalothorax with 2 large, heavily sclerotized, dac-
tyliform attachment processes on posterior lateral surface, processes
projecting laterally and slightly posteriorly. Anterior end of cepha-
lothorax with 4 adjacent rows of soft, irregular papillae (figs. 8, g)
running dorsoventrally, oral opening a distinct depression situated in
middle of anterior end of cephalothorax, between inner 2 rows of
papillae. Cephalothorax indistinctly separable from neck by indis-
tinct, incomplete line of division extending between posterior ends
of bases of attachment processes. Neck indistinctly separable into
two parts by slight constriction of neck and alimentary tract; pos-
terior end of neck separable from anterior end of trunk only by
difference in width. Anterior part of neck with distinct constriction
in figured specimen (fig. 8a) but constriction believed due to twisting
of specimen. Trunk elongate, width varying little throughout
length. Oviducal openings situated on indentation in posterior
third of trunk (fig. 8d), at beginning of abdominal portion of trunk.
Abdominal region tapered distally, bearing numerous ramified proc-
esses from lateral, ventral lateral, and ventral surfaces, with exten-
sion of alimentary tract running to blind termination at distal ends
of processes (fig. 8¢). Caudal rami knoblike, on distal lateral surfaces
of abdomen, flanking V-shaped indentation at anal opening (fig. 8c),
without setae; indication of armature present as 3 small irregularities
of heavily sclerotized ramal surface.

Antennules and antennae situated on median dorsal surface, just
posterior to frontal process region. Antennae anterior to antennules,
articulating on heavily sclerotized ridge, ridge expanded medially,
forming convex projection bearing small, indistinct remains of copep-
odite rostrum and small, heavily pigmented ocular region posterior to
rostrum (fig. 9a). Antennule (fig. 9b) 3-segmented although articulat-
ing in doughnut-shaped structure distinct from cephalothoracic
cuticle. First segment with 7 very lightly plumose setae on anterior
surface, second segment more elongate than first, with approximately
11 very lightly plumose setae on anterior and anterior dorsal surface in
addition to 1 elongate, aesthete-like structure on distal anterior sur-
face. Third segment slightly narrower than second, with one aesthete-
like structure from distal posterior surface, 7 naked processes from
distal surface (1 of 7 aesthete-like). Antennae (fig. 9c) 3-segmented,
first 2 segments broad, distalmost segment with clawlike terminal
process. Inner proximal corner of first segment forming knoblike
articulation surface, inner lateral margin of segment longer than
outer margin; distal surface irregularly cup shaped, inner portion
receiving inner proximal surface of second segment. Second segment
with cup-shaped projection in inner distal surface, projection receiving
distal end of terminal process of third segment when segment flexed,

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

with large, irregular, heavily sclerotized projection of outer portion
that forms articulation surface for third segment. Third segment
small, proximal end irregular, heavily sclerotized; division between
terminal process and segment indistinct; setalike accessory process
present, distal to division between segment and terminal process.

First 2 thoracic leg pairs in close proximity at anterior end of neck,
third placed well behind second, and fourth behind third; ratio of
proximity equals 1 (first to second leg) : 5.5 (second to third) : 5.5 (third
to fourth) (fig. 9d). Protopodite of legs 1-segmented although appear-
ing 2-parted, attached to well-developed interpodal plate. First
thoracic leg biramous, rami 2-segmented, second leg appearing
uniramous, both in adult female (fig. 9f) and larval exuvium (fig. 97),
although outline of leg and presence of exopodite on adult leg and
endopodite on larval exuvium suggest loss of endopodite in adult
specimen. Third and fourth thoracic legs without distinct indication
of rami. Protopodite of all legs with dark brown to black pigment
spot in alcohol-preserved specimens. For nature and armature of
thoracic legs, see figures 9e-j and table 5.

TaBLe 5.—Armature of thoracic legs I-IV of the female of Pennella histiophori?
Thomson, 1889

Exopodite Endopodite
Leg Surface Interpodal | Protopodite
Plate
1 2 1 2
I | Outer p 2P 3P
Inner p’ p 3P 12 4P
II | Outer 3P
Inner le 3P
III | Outer p’
Inner
IV | Outer
Inner p’

Discussion.—The identification of the Hawatian material as
Pennella histiophori? Thomson, 1889, as well as the inclusion of
the two names used by Kirtisinghe in the synonymy, is done with
some question. The intraspecific variation present within the mem-
bers of the genus, the wide distribution of many of the hosts of the
genus, and the relative absence of definitive descriptions leave the

Figure 9.—Pennella histiophori? Thomson, 1889, female: a, ocular region, dorsal view
showing rostrum (r), antennule base (a-1), and antenna base (a-2); b, antennule, dorsal
view; c, antenna, dorsal view; d, pedigerous region of neck, ventral view. Right thoracic
legs: e, first, posterior view; f, second (endopodite missing), anterior view; g, third (ramus
or rami missing), posterior view; h, fourth (ramus missing), posterior view; 7, distal end
of protopodite of fourth, posterior view. Pedigerous region of larval exuvium: j, ventral
view (A=anterior).

NO. 3674 COPEPOD CRUSTACEANS—LEWIS 25

~ <7

ee
\)
ae

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

taxonomy of the genus in confusion. Of all of the species, P. instructa
Wilson is by far the most easily recognized, and the characteristic
ramshorn-shaped attachment structures show little variation in the
specimens present in the U.S. National Museum. Yamaguti’s iden-
tification of Japanese pennellids from Xiphias gladius as P. instructa
(1939b) may warrant some question although there is more evidence
for his identification, from his figure (pl. 33, fig. 193) and from the
host, than there is for the inclusion of other material in the species.

The original description of P. histiophori Thomson, 1889, indicates
some differences between the type material and that here described.
P. histiophori is figured as having a larger rounded protuberance on
the ventral surface, between the bases of the attachment structures,
the 4 pairs of thoracic legs appear to be more closely grouped, and the
pedigerous area of the neck appears more swollen and distinct from
the rest of the neck. The size of the minute ventral protuberance
between the bases of the attachment structures is variable in the
3 specimens examined for this description, and there is variation in
the size, shape, and position of numerous small irregularities on the
cephalothorax. The grouping of the thoracic legs may be distinctive
although Thomson’s original figure is not clear enough to offer
grounds for separation. ‘The difference in the size of the pedigerous
region of the neck and the distinction of this region from the rest of
the neck may, based on an examination of the variation in some of
the other members of the genus, be due to intraspecific variation.

Pennella species

MarEriAL.—One complete and 1 incomplete (abdomen missing) non-
ovigerous female (USNM 112872) from the external surface of Remo-
ropsis brachypterus (Solander) collected by the USFWS from the
stomach of a yellow-fin tuna, Neothunnus macropterus (Schlegel), cap-
tured off Hawaii.

MEASUREMENTS.—(In mm) females:

Greatest length of body (N=1) 35.85

Length of cephalothorax (N=2) 4.00, 3.75

Width of cephalothorax (N=2) 2.85, 3.00

Length of neck (N=2) 10.88, 12.75

Width of neck (N=2) 1:65, 1.35

Length of trunk (N=1) 21.00

Width of trunk (N=1) 1.50

Approximate length of abdominal plumosities (N=1) 7.13

Length of attachment structures (N=2) (2.25, 2.25), (3.15, 4.05)

Description.—Cephalothorax (figs. 10a, 6) presumably consisting
of cephalon and first thoracic segment although maxillipeds not dis-
cernible. Pair of long, dactyliform attachment processes present
immediately posterior to cephalothorax. Cephalothorax with 2 types

NO. 3574 COPEPOD CRUSTACEANS—LEWIS Di

eZ Va\\\N\4 Nay
Hy ii Mi

i My
hin

Ficure 10.—Pennella species, female: a, dorsal view (specimen 1); d, anterior trunk region
and cephalothorax, dorsal view (spec. 2); c, cephalothorax, ventral view showing large

peripheral and smaller oral papillae (spec. 2); d, three oral papillae (spec. 2); ¢, f, abdominal
processes.

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

of soft protuberances (figs. 10c, d): first large, irregularly shaped, clus-
tered on anterior end but extending posterolaterally to mouth region,
pair also present on posteroventral surface, somewhat isolated from
other large protuberances; second type also irregular, smaller, clustered
around oral orifice. Neck indistinctly separable from cephalothorax,
width varying little throughout length, separated from trunk by slight
constriction. Trunk width varying little throughout length, with
narrow, V-shaped, heavily sclerotized indentation just posterior to
minute, lappet-shaped spermatophores adjacent to oviducal openings
(fig. 116). Abdominal portion of trunk tapered distally, with numer-
ous rodlike uniramous or biramous processes from lateral surface (figs.
10e, f), each with extension of alimentary tract running to blind termi-
nation at distal end. Distal end of abdomen bilobed, anal opening at
median indentation, each lobe with minute caudal ramus. Caudal
rami (fig. 1l@) consisting of 5 naked setules, 3 on irregular knob, 2
on abdominal surface adjacent to knob.

Dorsal surface of cephalothorax with narrow, U-shaped region
of heavy sclerotization anteriorly, apex of U directed anteriorly,
with irregular suborbicular projections; ends of U with minute, spike-
like projections. Antennal articulation surfaces present on region,
just lateral to apex. Antennules not present although indication
of articulation surface present lateral and slightly posterior to antennal
base. Antennae (fig. 11d) 3-segmented, segments 1 and 2 broad,
segment 3 and associated terminal process forming clawlike structure.
Outer proximal surface of first segment extending as knoblike artic-
ulation surface, inner lateral surface approximately three-fourths the
length of outer, with knoblike heavily sclerotized region distally.
Second segment slightly more than half the length of first, inner
distal and lateral surfaces forming spikelike projection, outer distal
surface irregular, forming articulation surface for third segment.
Third segment short, continuous with terminal process; small, lappet-
shaped accessory process present on proximal surface of fused segment
and terminal process.

First 2 thoracic leg pairs in close proximity at anterior end of
trunk, third somewhat distant from second and fourth from third.
Ratio of proximity of thoracic leg pairs equals 1 (first to second leg):
2 (second to third): 1.5 (third to fourth). Protopodites of all 4
legs 1-segmented although appearing 2-parted, all attached to distinct
interpodal plate. Thoracic legs I-III biramous (exopodite of leg II
lost in one specimen), rami 2-segmented; fourth thoracic leg probably
uniramous, ramus 2-segmented. Protopodite of all 4 legs with
dark brown to black pigment spot. Nature and armature of thoracic
legs given in figures 1le-h and table 6. Some errors are believed
incorporated into both the figures and the table because of the method

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 29

: o pee yt ————_f
be + 4 + Ht es +
ri O.1mm. Oi1mm. © 0.1mm. hb

Ficure 11.—Pennella species, female: a, caudal ramus, dorsal view (a=anus); b, ventral
portion of genital region, lateral view (sp—=spermatophore); c, ocular region of cephalon,
dorsal view showing antenna, heavily sclerotized spine, and ridge; d, right antenna, dor-
sal view. Thoracic legs, anterior view: e, right first; f, left second (exopodite missing); g,
right third; A, right fourth.

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

of collection of the specimens and the brittle nature of the thoracic
legs of adult females of the genus.

TABLE 6.—Armature of thoracic legs I-IV of the female of Pennella species

Exopodite Endopodite
Leg Surface Interpodal | Protopodite
Plate
1 2 1 2
I | Outer S, p’
Inner p’ ? 5P? ? 6P?
II | Outer s
Inner ? 6P? Pe 7P
Iii | Outer s
Inner 5P,p iE 6P,p
IV | Outer
Inner 4P,p

Discusston.—Although much of the body and many of the ap-
pendages and processes of the 2 specimens were apparently undamaged
by the ingestion and partial digestion of the host, there is some evi-
dence of damage (e.g., thoracic legs). For this reason it is felt that
specimens in better condition are essential before a name is given to
this species. The specimens, however, have characteristics similar
to several species of the genus:

1. Pennella selaris Kirtisinghe, 1964: Similar in cephalothorax,
orientation of abdominal plumosities. Differs from Pennella species
in possessing shorter and unbranched abdominal plumosities and in
the arrangement of the thoracic legs.

2. P. biloba Kirtisinghe, 1932: Cephalothorax and general body
shape similar. Abdominal processes unbranched in P. biloba,
branched in P. sp.

3. P. diodontis Oken, 1816: Arrangement of cephalothoracic
papillae and abdominal processes similar. Abdominal processes
unbranched in P. diodontis.

4. P. exocoeti (Holten, 1802): Arrangement and branching of
abdominal processes similar. Cephalothorax not bilobed in P.
exocoeti, nature and arrangement of cephalothoracic papillae appear
different.

5. P. remorae Murray, 1856: Abdominal processes in 2 rows, as
in P. sp. Cephalothorax of P. remorae not lobed, with 3 horns
(characteristic variable), anterior end cuplike (not depressed in P. sp. ),
neck more distinct, abdominal processes more ramified.

Family Anthosomatidae
Norion Nordmann, 1864

Diacnosis.—See Lewis, 1964a.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 31

Norion expansus Nordmann

Norion expansus Nordmann, 1864, p. 489, pl. 6—Lewis, 1964a, p. 226, figs. 22-23.
[For bibliography, see Lewis, 1964a.]

Hosts AND DISTRIBUTION.—3 host records:

location hosts reference
Hawaiian Islands unknown Nordmann, 1864
Naso lituratus
N. hexacanthus Lewis, 1964a

DeEscrIption.—See Lewis, 1964a.

Family Pandaridae
Nesippus Heller, 1865
Diacnosis.—See Lewis, 1964a.

Nesippus costatus? Wilson

Nesippus costatus Wilson, 1924, p. 2138, pl. 20.
Nesippus costatus?—Lewis, 1964a, p. 211, figs. 19-21.

Hosts AND DISTRIBUTION.—7-+ host records:

location hosts reference
Galapagos Islands ‘“‘A nine-foot shark” Wilson, 1924
Hawaiian Islands Cysts on: Lewis, 1964a
Acanthurus triostegus sandvi-
censis
A. nigroris

A. zanthopterus
Ctenochaetus strigosus
Diodon holocanthus
Scarus species
labrids
pomacentrids
zanclids

MAaTERIAL.—One immature male (early encysted stage of Lewis,
1964a) in cyst on dorsal fin of Seriola dumerilli (Risso) from Hono-
lulu Aquarium. Cysts with exuviae (no animal) or partially resorbed
cysts found on specimens of Aulostomus chinensis (Linnaeus), Chae-
todon miliaris Quoy and Gaimard, and Myripristis pralinius Cuvier
and Valenciennes.

MEASuUREMENTS.—Length of body, excluding caudal setae, 1.96mm.
Length of cephalothorax, including frontal region, 1.39mm; width
155mm. Length of genital segment 0.34mm; width 0.47mm.
Length of caudal rami 0.22mm.

DeEscrIPTION.—See Lewis, 1964a.

Family Euryphoridae
Euryphorus Milne-Edwards, 1840

Draenosis.—Cephalothorax suborbicular, consisting of cephalon

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

and first 4 thoracic segments, frontal region well defined. Free fourth
pedigerous segment with alae (larger in female than in male). Female
genital segment large, suborbicular, with pair of posterior lobes; male
genital segment longer than wide, of variable shape but lateral mar-
gins generally parallel, without posterior lobes. Abdomen 2- or 3-
segmented, female first segment elongate, with pair of large, lamellate
processes extending posteriorly past caudal rami, remaining segment
or segments without lamellate processes; male first segment shorter
than in female, lamellate processes thicker and smaller, extending
posteriorly only to region of second abdominal segment, remaining
segment or segments as in female. Caudal rami similar to those of
other euryphorids. Antennule 2-segmented; antenna 3-segmented,
prehensile, male with enlarged secondary spines. Mandible rod
shaped, denticulated distally; maxillule nodular, tipped by 3 setae;
maxillae 2-segmented. Postantennal process consisting of 2 nodules
with hairlike processes; postoral process spinelike. Maxilliped 2-
segmented, prehensile, with 2 subtriangular protuberances on first
segment. Sternal furca with divergent tines in female, almost paral-
lel tines in male; furca-like processes present posterior to interpodal
plate of first and second thoracic legs. Thoracic legs I-IV biramous,
ramal segment count 2-2, 3-3, 3-3, 3-2, fifth legs setiform.

Euryphorus nordmanni Milne-Edwards

Figures 12-15

Euryphorus nordmanni Milne-Edwards, 1840, p. 462, pl. 39, fig. 1.—Kner, 1859,
p. 268, figs. 1-3.—Bassett-Smith, 1899, p. 461.—Kirtisinghe, 1937, p. 445, figs
74-87.— Yamaguti, 1963, p. 98.

Euryphorus nympha Steenstrup and Liitken, 1861, p. 365, pl. 6, fig. 12.—Heller,
1868, p. 198.—Bassett-Smith, 1899, p. 461.—Wilson, 1913, p. 225.—Shiino,
1954b, p. 284, figs. 5-6; 1959a, p. 350; 1959b, p. 20, fig. 9—Ho, 1963, p. 83,
figs. 1-3.— Yamaguti, 1963, p. 99.— Kirtisinghe, 1964, p. 88, fig. 104.

Huryphorus coryphaenae Krgyer, 1863, p. 161, pl. 10, fig. 4a—h.— Wilson, 1913, p.
225.—Yamaguti, 1936b, p. 1, pl. 1, figs. 1-17 Bonnet, 1948, p. 7.—Causey,
19538a, pp. 7, 9; 1953b, p. 11; 1955, p. 6.

REMARKS.—See discussion section, following description, for dis-
cussion of synonymy.
Hosts AND DISTRIBUTION.—10 host records:

locality host reference
‘“‘Des mers d’ Asie” Unknown Milne-Edwards, 1840
Western Pacific Coryphaena hippurus Yamaguti, 1936b
Neothynnus macropterus Shiino, 1954b
Hawaii Coryphaena hippurus Bonnet, 1948
Indian Ocean “‘Seefischen”’ Kner, 1859

Coryphaena hippurus
C. h. equisetis Kirtisinghe, 1937

No. 3574 COPEPOD CRUSTACEANS—LEWIS 33

ocality host reference
Tropical, sub-
tropical
Atlantic Lampugus punctulatus Steenstrup and Litken, 1861
Coryphaena hippurus Krgyer, 1863
Gulf of Mexico Coryphaena hippurus Causey, 1953a

Mareriat.—Two females and 2 males (USNM 112873) from the
gill cavity of Coryphaena hippurus Linnaeus taken by trolling 120
miles south of Oahu, Hawaii (USFWS). Two females and 2 males
(USNM 112874) from the gill cavity of C. hippurus Linnaeus taken
by trolling at 15°N, 115°W (USFWS). One female and one male
(USNM 112875) from the gill cavity and buccal cavity of C. hippu-
rus Linnaeus taken by trolling, 130 miles south of Nihau, Hawaii
(USFWS).

MEASUREMENTsS.—(In mm) 5 females and 5 males:

female male
mean range mean range

Length of body, excluding caudal setae 9.65 7.80-10.50 5.84 5. 33-6. 45
Length of cephalothorax, including frontal

region 2.77 2.63- 2.96 2.46 2.37-2. 52
Width of cephalothorax 2.75 2.63- 2.81 2.35 2. 26-2. 52
Length of genital segment, excluding gen-

ital flap POS) rol 62022) N27 pAb 1 44
Width of genital segment 29D el So 26 Sh. ded sand 04 18
Length of abdomen 4.31 2.70—- 5.10 1.50 1. 43-1. 58
Length of alae 0.78 0.74- 0.83 0.55 0. 45-0. 61
Length of genital flap 0.67 0.61- 0. 72
Length of abdominal projection 4.85 2.70- 6.08 1.10 0. 98-1. 28
Length of caudal rami 0.46 0.43- 0.49 0.34 0. 31-0. 36

Length of egg string (1 string) 9. 23

Description.—Female cephalothorax (fig. 12a) ovoid, consisting
of cephalon, maxilliped-bearing, and first 3 pedigerous segments.
Frontal region broad, approximately one-tenth the length of cepha-
lothorax, distinctly separable from rest of cephalothorax; anterior
edge with fine membrane. Lateral cephalothoracic margins flatly
convex except for sharp median indentation, margin bearing narrow
membranous flange. Posterior lateral cephalothoracic regions sharply
rounded, extending posteriorly slightly past median cephalothoracic
region. Posterior sinuses distinct (fig. 12c), U-shaped, with narrow
but distinct flange extending medially from outer margin and bear-
ing fine membrane. Dorsal cephalothoracic grooves forming irreg-
ular H, posterior longitudinal grooves of H formed by junction of
posterior lateral cephalothoracic region and median cephalothoracic
region; crossbar irregular, in posterior third of body; anterior longi-
tudinal grooves extending anteriorly to just lateral to 3-parted ocular
region. Ocular region consisting of 2 ovoid contiguous pigmented
bodies, each with lens, and smaller pigmented area between posterior

226-321 67—-_8

34

PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 127

2.0mm.

Ficure 12.—Euryphorus nordmanni Milne-Edwards, 1840, dorsal view: a, female; ), male;
c, female posterior cephalothoracic sinus and free fourth pedigerous segment, including

ala.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS a

regions of ovoid bodies. Posterior median cephalothoracic region with
several spinules dorsally.

Female free fourth pedigerous segment distinct from cephalotho-
rax and genital segment, greatest length, excluding alae, slightly
less than one-fifth the length of cephalothorax. Segment broadest
medially, with short, necklike region anteriorly. Alae small, sub-
rectangular although shape variable, arising from broad medial
portion of segment, extending posteriorly over part of anterior dorsal
surface of genital segment. Alae not connected as indicated by
Shiino (1954b). Median dorsal surface of segment with several
spinules.

Female genital segment (fig. 13a) broad, lateral margins irregular,
lateral surface projecting laterally past ventrally swollen medial
region. Anterior end of segment narrow, bearing pair of heavily
sclerotized, leglike plates (fig. 13¢) ventrally, each with fimbriated
edge. Plates connected by ridge of heavy sclerotization similar
to interpodal plates of fourth thoracic legs. Homologies of struc-
ture unknown, no comparable structure on male or on members of
genus Elytrophora, which appears closely related to HEuryphorus.
Posterior surface of genital segment irregular, with pair of knobby
projections ventrally, in region of oviducal openings and with pair
of small, lappet-shaped projections dorsally, overlapping egg strings.
Female fifth legs (fig. 13c) situated ventral and lateral to base of
lappet-shaped projections, consisting of 2 nodules, first bearing
single plumose setule, second bearing 3 plumose setules.

Female abdomen (fig. 12@) elongate, distinctly separable from
genital segment ventrally, indistinctly separable dorsally. Segmen-
tation of adult female obscured although suggestion of 3 segments.
First segment approximately 65 percent of total length, slender,
bearing pair of large, curled, or uncurled flaps projecting posteriorly
past caudal rami. Division between first and second segments
suggested only by end of base of flaps although division complete
in male. Second segment elongate, forming 25 percent of abdomen
length. Division between second and third “segments” indicated
by cuticular constriction, third “segment” short, approximately 9
percent of abdomen length, median posterior surface projecting
slightly in biconvex anal region. Caudal rami (fig. 13f) ovate, mar-
gins irregular, distal inner surface plumose; distal surface with 2
large plumose setae from tubercular projection, 2 additional large
setae, one on either side of projection, and 2 plumose setules.

Male cephalothorax (fig. 12) similar to that of female although
indentation of lateral margin less distinct and posterior median
region with fewer dorsal spinules. Free fourth pedigerous segment
similar, alae smaller, of general ovoid shape. Genital segment (fig.

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

13b) distinct from fourth pedigerous segment, shape irregularly
ovoid. Segment appearing 2-parted dorsally, with indication of
break ventrally; anterior fourth of segment (in region of ventrally
projecting plates in female) with distinct line of division on lateral
dorsal surface. Fifth legs (fig.13d) situated on lateral ventral surface
in middle of segment, consisting of 2 nodules, anteriormost with
single plumose setule, posteriormost with 3 plumose setules. Abdo-
men distinct from genital segment, elongate, segmentation more
distinct than in female. First segment approximately 53 percent of
abdomen length, with pair of large flaps (smaller than those of female
although thicker) extending posteriorly to end of second segment.
Second segment approximately 27 percent of abdomen length, cylin-
drical; third ‘‘segment’’ approximately 20 percent of abdomen length,
similar in shape to that of female. Caudal rami as in female.

Female antennule (fig. 13g) 2-segmented, attached to lateral-ante-
rior ventral surface of cephalothorax and posterior-lateral ventral
surface of frontal region. First segment slightly more than twice
the length of second, with approximately 15 naked or lightly plumose
setules. Second segment cylindrical, with single naked setule from
median dorsal surface, second from median posterior surface, ap-
proximately 11 from distal region. Male antennule similar to that of
female although sclerotization of first segment irregular, giving
pseudosegmented appearance (Shiino’s ‘‘2 false joints,” 1954b, p. 288).
Female antenna (fig. 13h) 3-segmented, attached posterior and
medial to antennule base. First segment short, irregular, with
posteriorly projecting spike from posterior surface. Second segment
strongly developed, without any major irregularities. Third segment
and terminal process clawlike, segment separable from terminal
process only by indistinct break in sclerotization, with setalike acces-
sory process from proximal posterior surface, second from distal posterior
surface. Male antenna (fig. 132) similar to that of female except
spikelike projection of first segment smaller, third segment distinctly
separable from terminal process, terminal process with spikelike
projection from proximal posterior surface.

Female and male mandible (figs. 13h, 7) 3-parted, rodlike. First
part broad proximally, tapered distally; second part short, slightly
less than half the length of first, tapered. Third part elongate, slightly

Figure 13.—Euryphorus nordmanni Milne-Edwards, 1840; ventral view: a@, female free
fourth pedigerous segment, genital segment (with projecting plates at anterior end), and
anterior end of abdomen; b, male, same; c, female fifth leg; d, male fifth leg; ¢, anterior
end of female genital segment showing projecting plates; f, caudal ramus; g, left anten-
nule; h, female oral region, left side showing antenna, postantennal process, mouth cone,
mandible (mdbl), maxillule, postoral process, maxilla base (ma-2), and postoral V-shaped
ridge; 7, male left antenna; j, mandible.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS a7

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

less than twice the length of first, distal region curved inward
slightly, with 12 denticulations on inner surface. Female and male
postantennal process (fig. 13h) evidenced only by pair of nodules
lateral to antenna base, each with several hairlike processes. Female
and male postoral process (fig. 13) spikelike, with broad base, tapered
to sharp distal end, outer margin sharply indented medially. Female
and male maxillule (fig. 13h) nodular, with 3 setule-like processes
distally. V-shaped heavily sclerotized ridge present posterior to

0.3 mm.d

Ficure 14.—Euryphorus nordmanni Milne-Edwards, 1840, ventral view: a, left maxilla;
b, female left maxilliped; c, male, same; d, female sternal furca, interpodal plates of first and
second thoracic legs (ip-1, 2), and furcal modification of platelike projections behind inter-
podal plates; ¢, male sternal furca.

mouth cone (fig. 13h), im same region as adhesion pads of pandarids,
some euryphorids and some ealigids. Female and male maxilla (fig.
14a) 2-segmented, situated lateral and slightly posterior to postoral
process, arising from padlike projection. First segment approximately
three-fourths the length of second; second segment elongate, broadest
medially, with membrane extending along median posterior surface.
Distal end of second segment with 2 setalike processes, innermost
slightly less than twice the length of outermost, with fine membrane

NO. 3574

COPEPOD CRUSTACEANS—LEWIS 39

along inner margin; outermost process with frilled membrane along
outer margin.

Female maxilliped (fig. 140) 2-segmented, situated posterior and
medial to maxilla base, on ovoid, padlike projection. First seg-
ment strongly developed, irregular, narrow proximally, broad medially,
with 2 subtriangular projections from inner surface. Second segment
and terminal process clawlike, segment short, distinct from terminal
process, with setule-like accessory process from distal inner surface.
Male maxilliped (fig. 14) similar to that of female except with addi-
tional, knoblike projection on proximal inner surface of first segment.
Female sternal furca (fig.14d) V-shaped, tines bluntly pomted. Male
sternal furea (fig. 14e) with V-shaped sinus but lateral margins almost
parallel, tines not diverging as in female.

For nature of thoracic legs and armature, see figure 15 and table 7.
Flabby, bifurcate projection present just posterior to interpodal plate
of first thoracic leg, similar though less distinct projection present
posterior to second thoracic legs.

TABLE 7.—Armature of thoracic legs I-IV of the female and male of
Euryphorus nordmanni Milne-Edwards, 1840

Inter- Protopodite Exopodite Endopodite
Leg | Surface || podal
Plate
1 2 1 2 3 1 2 3
I | Outer h d,dH,2dH, Cle
12
Inner c 3P 2P
II | Outer m,p | m,d,fmH | fmH 2imHe@E2P kc Ee 3P
m
Inner See TMSea Cee ee oe. Pae2P 3P
Ili* | Outer m,P d,H d,h 3h,2P Cialae 3P
m
Inner 2s,rt,P,m,3s Cue Cre 3P JEW OpAe iP
IV* | Outer p d,fmH d,fmH d,fmH,d, Cu a2
2fmH
Inner fm ie 4P Paar e

*Numerous hairlike projections (s) in middle of segment, not included in table.
tRoughened area (in this case by denticulations).

Discusston.— The synonymyzing of Euryphorus coryphaenae Kr¢yer
with EL. nympha Steenstrup and Liitken was first done by Bassett-
Smith (1899) although no reasons were given. Wilson (1913) sug-
gested that the two were synonymous but it was not until Shiino
(1954b) that Bassett-Smith’s belief was reiterated. Shiino did not
have Krgyer’s original description of 7. coryphaenae so did not give
an analysis of the two species.

In a comparison of EL. nympha and FE. coryphaenae, the major differ-
ences appear, from the publications, to be in the sternal furea (that of
EF. nympha having more widely spaced tines) and in the armature and
its position on the first thoracic leg (7. coryphaenae figured as having

AO PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Ficure 15.—Euryphorus nordmanni Milne-Edwards,1840, right thoracic legs, anterior view:
a, first; b, distal end of second segment of exopodite of first; c, second; d, third; ¢, exopodite
of third (posterior view); f, fourth.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 4]

the terminal elements more broadly spaced on the exopodite and
having 3 short, spinelike projections in addition to the 3 setae figured
for both species). The variation present in the angle of the furcal
tines, the rather confusing picture given by the terminal elements on
the second segment of the exopodite of the first thoracic leg (fig. 158),
the similarity of the general body shape and particularly the first
abdominal segment (which is considered, by this author, to be of
taxonomic significance in this genus and in the genus Elytrophora),
in addition to the similarity of other body parts described and figured
in the original descriptions suggest that the two species are conspecific.

The reasons given by Steenstrup and Liitken (1861) for separating
Euryphorus nympha from £. nordmanni Milne-Edwards are:

1. EH. nordmanni has a better developed pair of antennules than
E. nympha.

2. The specimens in the Paris museum (£. nordmanni) were
from the “‘Asiatic’’ ocean while #. nympha was described from material
collected in the subtropical Atlantic.

Kner (1859) described H. nordmanni from fish captured off Zanzibar.
Steenstrup and Liitken, however, did not feel that Kner had sufficient
material or sufficient experience (based at least partially on his
misinterpretation of the first thoracic legs) to identify his specimens
properly and felt that his material was 7. nympha.

Specimens identified as either HL. nympha or E. coryphaenae have
been reported from most of the tropical and subtropical regions of the
world (Japan, Indian Ocean, Atlantic, West Indies, Gulf of Mexico).
The distribution of #. nympha (recognizing EF. coryphaenae as a syno-
nym of this species) appears to parallel that of the principal host,
Coryphaena hippurus Linnaeus. The host that Milne-Edward’s and
Kner’s specimens came from is unknown and £. nordmanni has been
reported only once, from Coryphaenae hippurus, since Kner’s publi-
cation (Kirtisinghe, 1937). Even this report was later believed
erroneous, Kirtisinghe (1964 and personal communication) feeling
that his specimens should have been identified as H. nympha.

The validity of EL. nympha appears to be dependent upon the
absence of the host name in the original description of #. nordmanni,
the different locality from which EL. nordmanni was collected, and
the belief that EL. nordmanni has a better developed pair of anten-
nules. There even appears to have been some question, by Steen-
strup and Liitken (1861), as to whether their species, H. nympha,
was distinct. The following is a translation of the second footnote
on page 366 of their 1861 publication:

There is, unfortunately, an almost complete lack of precedent as to whether

or not we can refuse to equate an animal type to one already described in the
literature just because one is from the Atlantic and one from the Indian Ocean.

42 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Are the larger pelagic fish forms actually the same for these two oceans, or does
each have their own characteristic species? So little is at present known about
this that one assumption appears to be valid as the other. We are inclined
to prefer the latter, and do therefore not wish to accept that the same species
of parasitic crustaceans occur in both oceans as long as a possible comparison
is lacking.

The wide distribution and the similar and unique morphology
of the E. nordmanni-E. nympha-E. coryphaenae complex indicates,
to this author, that these 3 species are synonymous.

Elytrophora Gerstiecker, 1853

Diagnosis.—Cephalothorax consisting of cephalon and first 4 tho-
racic segments, frontal region distinct; lateral margins with slight
indentation medially. Free fourth pedigerous segment with alae
extending posteriorly over anterior region of genital segment; fe-
male alae with sharp indentation medially, male alae with or without
slight indentation. Female genital segment variable in size and
shape, longer in egg-producing females than in immature females,
with pair of posteriorly projecting, lobe-shaped plates, plates straight
or angled inward slightly. Male genital segment without plates.
Abdomen 2-segmented, first segment of female with small, posteri-
orly projecting lateral lobes, lobes of variable size and shape; first
segment of male without lobes. Second segment of abdomen with
small, bilobed anal projection. Antennule 2-segmented; antenna 3-
segmented, with clawlike terminal process (male with small second-
ary spine at base of terminal process). Mandible rodlike, with 12
denticulations on inwardly curved distal region. Maxillule nodular,
bearing setules; maxilla 2-segmented; postoral process spinelike,
distal region more slender in mature specimens than in immature.
Maxilliped 2-segmented, with clawlike terminal process. Thoracic
legs I-IV biramous although endopodite of first and fourth legs re-
duced; ramal count 2-2, 3-3, 3-3, 3-2. Fifth legs present, setiform.

Remarxks.—The similarity of EHuryphorus and Elytrophora is re-
markable. This similarity is principally in the general shape of the
cephalothorax, the female alae, genital plates, abdominal processes
(although they are much larger in Huryphorus), and number of seg-
ments in the rami of the first 4 pairs of thoracic legs (identical in
members of both genera).

Elytrophora brachyptera Gerstiecker

FicureEs 16-21

Elytrophora brachyptera Gerstiecker, 1853, p. 60, pl. 3, fig. 12—Nordmann, 1864,
p. 468.—Heller, 1866, p. 753.—Van Beneden, 1870a, p. 57.—Richiardi,
1880, p. 3.—Stossich, 1880, p. 257.—Valle, 1880, p. 60.—Carus, 1885, p. 360.—

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 43

Bassett-Smith, 1896a, p. 158; 1896b, p. 12, pl. 4, fig. 3—Brian, 1899a, p. 4.—
Bassett-Smith, 1899, p. 462.—Brian, 1906, p. 51, pl. 1, figs. 2-3; 1908, p. 3;
1912, p. 10.—Scott and Scott, 1913, p. 83, pl. 19, fig. 10; pl. 23, figs. 1-2;
pl. 31, figs. 1-6—Yamaguti, 1936b, p. 3—Bonnet, 1948, p. 7.—Delamare-
Deboutteville and Nunes-Ruivo, 1953, p. 202, fig. 1—Shiino, 1954b, p. 279,
figs. 3-4.—Heegaard, 1955, p. 46, figs. 9-11.—Shiino, 1957a, p. 364.—
Yamaguti, 1963, p. 102, pl. 123, fig. 1.

Dinematura Thynni Krgyer, 1863, p. 157.—Wilson, 1907a, p. 376 (as thynni).
[Nomen nudum, label name attributed to Kollar.]

Arnaeus thynni Kréyer, 1863, p. 157, pl. 8, fig. 5a-g.

MatTeriAu.—15+ host records:

locality hosts reference
Mediterranean Unknown Gerstaecker, 1853
Thynnus vulgaris Heller, 1866
Thynnus thynnus Brian, 1899a
“Germon”’ Brian, 1908
“Thon rouge” Delamare-Deboutteville
and Nunes-Ruivo, 1953
Northeastern Atlantic Thynnus vulgaris Van Beneden, 1870a
Thynnus thynnus Bassett-Smith, 1896a
Orcynus thynnus Scott and Scott, 1913
Subtropical Atlantic Plankton? Heegaard, 1955
Pacific Thynnus thynnus
T. alalunga Yamaguti, 1936a
Western Pacific Thynnus orientalis
Parathynnus sibi Shiino, 1954a
Neothunnus albacora
Parathunnus obesus Shiino, 1957a
Hawaii Thynnus species Bonnet, 1948

Materiau.—Two adult females and 4 adult males (USNM 112876)
from the gill cavity of Parathunnus sibi Schlegel from the Honolulu
Fish Market (USFWS collections). Two adult females and 1 adult
male (USNM 112877) from the gill cavity of Neothunnus macropterus
(Schlegel) captured by longline near Christmas Island (Line Islands)
(JRM cruise 27, longline station 2). Four adult females (USNM
112878) from inside operculum of an unknown host (probably N.
macropterus (Schlegel)) captured by longline north of Christmas
Island (Line Islands) (LL cruise 6). Additional specimens, retained
by the author, include 2 adult females and 2 adult males from inside
operculum of unknown host captured by longline north of Christmas
Island (Line Islands) (USFWS); 1 adult female and 1 adult male
from the external surface of N. macropterus (Schlegel) captured by
longline at 0°29’N, 157°49’W (USFWS); 3 adult males from inside
operculum of unknown host captured by longline north of Christmas
Island (Line Islands) (USFWS); 2 adult females from inside oper-
culum of unknown host captured by longline north of Christmas
Island (Line Islands) (USFWS).

44 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

MEASUREMENTS.—(In mm) 10 females and 10 males:

female male
mean range mean range

Total length, excluding caudal setae 8.37 6.23- 9.68 6.33 5. 10-7. 80
Length of cephalothorax, including frontal

region 4.43 2.85- 4.88 3.45 3. 23-4. 05
Width of cephalothorax 4.51 3.48- 4.95 3.10 2. 48-3. 60
Length of genital segment, excluding gen-

ital flap 1.93 1.13— 2.37 1.45 1. 18-1. 70
Width of genital segment 1.70 0.72—- 1.95 1.27 0. 58-1. 43
Length of abdomen 1.27 0.99- 1.52 1.04 0. 70-1. 44
Length of alae (N=9 in female) 1.21 1.00- 1.33 0.73 0. 56-0. 85
Length of genital flap 0.87 0. 70—- 1.15
Length of caudal rami 0.57 0.49- 0.63 0.53 0. 44-0. 63
Length of egg strings (4 strings) 9.13 7. 50-10. 50

ReEMARKS.—Two sets of descriptions are given below, the first for
the mature adult, the second for the immature adult. The author
feels that the changes that occur in the adult life history stage, as
indicated in the measurements, may have caused some confusion in
the taxonomy of the genus. The bases for this belief are given in
the discussion section, following the description of the species.

DescripTion.—(Mature adult): Female cephalothorax (fig. 16a)
ovoid, consisting of cephalon and first 4 thoracic segments. Frontal
region distinct, length approximately one-twelfth that of entire cepha-
lothorax, with median cleft and narrow membrane along anterior mar-
gin. Lateral cephalothoracic margins with indentation medially,
distinct in some specimens, indistinct in others; margin with fine
membrane. Posterior lateral cephalothoracic region broadly rounded,
extending past median posterior region, with socket-like depression
(not figured); posterior sinus (fig. 16c) distinct, U-shaped, with fine
membrane along outer margin. Median-posterior region with pair
of small lobes laterally. Major dorsal cephalothoracic grooves form-
ing irregular H, anterior legs extending past indistinct ocular region
(ocular region not figured). Dorsal cephalothoracic surface without
plumosities but with minute, spikelike irregularities laterally and
small denticulations posteriorly.

Free fourth pedigerous segment, excluding alae, approximately one-
fifth width of cephalothorax, widest medially, at origin of fourth
thoracic legs. Alae sharply indented on median lateral margin,
extending posteriorly over anterior end of genital segment. Divi-
sion between fourth pedigerous and genital segments distinct, demar-
cated ventrally by ridgelike transverse projection.

Female genital segment (fig. 16d) of variable shape, narrow anteri-
orly in some specimens, broad in others, posterior dorsal surface with
pair of lateral lappets, lappets distinct from segment. Dorsal sur-
face of segment lightly rugose, extending laterally slightly past ven-

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 45

tral region. Fifth legs (fig. 16f) arising from posterior lateral surface
of dorsal overlap, consisting of 2 nodules, anteriormost with single
setule, posterior with 3 setules.

H+ |
2.0mm.?

Ficure 16.—Elytrophora brachyptera Gerstaecker, 1853, mature adult, dorsal view: a, fe-
male; b, male; ¢c, posterior cephalothoracicsinus. Ventralview: d, female genital segment
showing spermatophore (sp), abdomen, and caudal ramus; e, male genital segment, abdo-
men and caudal ramus; f, female left fifth leg; g, male left fifth leg.

Abdomen 2-segmented, distinct from genital segment ventrally,
indistinctly separable dorsally. First segment, excluding lateral
projections, slightly shorter than second, with pair of distinct, flap-
like projections extending laterally and posteriorly, to middle of second

46 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

segment. Second segment convex laterally, flat posteriorly except
for slight, bilobed anal projection. Caudal rami broad, with 4 plu-
mose setae and 2 lightly plumose setules distally, with row of fine
denticulations around base of outermost seta.

Male cephalothorax (fig. 165) similar to that of female although
lateral indentations more distinct. Alae on free fourth pedigerous
segment tapered laterally, without sharp constriction present in
female. Genital segment (fig. 16¢) with flatly convex lateral margins
anteriorly, almost straight lateral margins posteriorly. Fifth legs
(fig. 16g) similar to those of female except posteriormost nodule with
2 lightly plumose setules instead of 3. Abdomen 2-segmented, first
segment narrow anteriorly, angled to widest point medially, lateral
margins straight posteriorly. Second segment and caudal rami
similar to that in female.

Female and male antennule (fig. 17a) 2-segmented, attached to
anterior-lateral ventral surface of cephalothorax and adjacent portion
of frontal region. First segment slightly less than twice the length
of second, tapered distally, bearing 23 naked or lightly plumose setae
and setules on distal half of anterior ventral surface and distal ven-
tral surface. Second segment club shaped, with 14 naked setules
distally. Female antenna (fig. 17) 3-segmented, situated posterior
and medial to antennule base. First segment short, irregular, with
well-developed, posteriorly projecting spine from lateral posterior
surface. Second segment broader proximally than distally, with
minute setule on distal inner surface (not figured); third segment
fused with clawlike terminal process, bearing flabby, setalike accessory
process proximally, more elongate, setalike accessory process from
median third, at break in sclerotization suggesting point of fusion of
segment and terminal process. Male antenna (fig. 17c) similar to
that of female although projection on first segment shorter, with sec-
ondary claw on terminal process of third segment.

Female and male mandible (fig. 175) rodlike, jointed in proximal
medial region, proximal portion tapered to joint, distal part flattened
distally, with indistinct break at beginning of flattened part; flattened
region curved inward, with 12 denticulations along inner margin Fe-
male and male postantennal process (fig. 176) consisting of 2 nodules,
each bearing several hairlike projections. Female and male postoral
process (fig. 176) broadly based, distally slender, spinelike, curving
laterally distally. Maxillule (fig. 176) situated lateral to mouth cone
base, immediately anterior to postoral process, consisting of node bear-
ing naked seta and 2 naked setules. Maxilla (fig. 17d) 2-segmented,
situated lateral and slightly posterior to postoral process. First
segment slightly less than four-fifths the length of second, excluding
terminal processes. Second segment elongate, broadest medially,

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 47

Ficure 17.—Elytrophora brachyptera Gerstaecker, 1853, mature adult, ventral view: a, right
antennule; b, female oral region, right side showing antenna, postantennal process, mouth
cone, mandible, maxillule, and postoral process; c, male right antenna; d, left maxilla; e,
female maxilliped; f, male maxilliped; g, sternal furca.

48 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

with small, fan-shaped projection from outer margin at widest point
and 2 larger, frilled membranes more distally, 1 on inner surface,
second on outer. Second segment also with small roughened area on
distal outer surface, similar to that of adhesion surface; distal end
bearing 2 setalike terminal processes, innermost slightly less than
twice the length of outer, with fine membrane along outer and inner
margin, outermost with finely frilled membrane along outer margin.

Female maxilliped (fig. 17¢) 2-segmented, situated posterior and
slightly medial to maxilla base. First segment strongly developed,
narrow proximally and distally, broad medially, with pair of small,
shelflike projections on medial posterior surface. Second segment
short, distinct from clawlike terminal process, with setule-like acces-
sory process on inner surface, at junction of segment and terminal
process. Male maxilliped (fig. 17f) differing from that of female by
presence of large, lobate projection from proximal inner surface of
first segment, projection containing heavily sclerotized, clawlike proc-
ess; ridges on first segment not as distinct as in female.

Female and male sternal furca (fig. 17g) V-shaped, tines slender,
extending posteriorly over interpodal plate of first thoracic legs.

For nature of legs and armature, see figures 18a—f and table 8.
Small, platelike projection present immediately behind interpodal
plate of first thoracic legs, with minute nodules in immature adults,
similar to those of Huryphorus nordmanni Milne-Edwards.

DescripTion.—Immature adult. Female cephalothorax (fig. 19a)
ovoid, consisting of cephalon and first 4 thoracic segments. Frontal
region approximately one-twelfth the total length of cephalothorax,
with median cleft and narrow membrane on anterior margin. Lateral
cephalothoracic margins convex, with membrane; with distinct inden-
tation medially, indentation formed by folding of dorsal cuticle, ex-
tending inward to median cephalothoracic region. Posterior lateral
cephalothoracic region broadly rounded, terminating at junction with
median cephalothoracic region, forming outer surface of posterior sinus
(fig. 19c). Sinus irregularly U-shaped, outer margin with fine, irreg-
ular membrane. Posterior median cephalothoracic region not extend-
ing to end of posterior lateral cephalothoracic region, margin lobed
laterally, flat medially, distinct from anterior end of free fourth pedig-
erous segment. Major dorsal cephalothoracic grooves forming irregu-
lar H, anterior legs of H extending to just posterior to ocular region,
turning inward sharply. Ocular elements distinct. Dorsal cephalo-
thoracic surface with numerous, elongate plumosities arranged in 2
groups, one extending anteriorly, from just posterior to crossbar of
dorsal cephalothoracic grooves, to anterior region of longitudinal
grooves; second group, containing fewer plumosities, located on poste-
rior lateral cephalothoracic region.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 49

<n
Oimm.2 hw

n

iin yO

-

¢

cae ieesen eral
0.2mm. 3

Figure 18.—Elytrophora brachyptera Gerstaecker, 1853, mature adult, right thoracic legs,
anterior view: 4, first; b, second segment of exopodite of first; c, second; d, third; ¢, exopo-
dite of third (posterior view); f, fourth.

226-321— 67-4

50 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Female free fourth pedigerous segment, excluding alae, slightly
more than one-fifth the width of cephalothorax, widest medially, at
attachment of fourth thoracic legs. Alae widest anteriorly, con-
stricted medially, broadly rounded posteriorly, covering anterior third
of genital segment.

Female genital segment (fig. 19d) broadly rounded anteriorly,
lateral margins flatly convex, lateral-posterior dorsal surface extending
posteriorly as pair of lobate projections, projections not distinct
from genital segment. Dorsal surface of genital segment platelike,
extending laterally past somewhat irregular posterior ventral surface.
Fifth legs (fig. 19/) arising from ventral lateral surface of platelike pro-
jection of dorsal surface of genital segment, consisting of pair of nod-
ules, anteriormost with single setule, posterior with 3 setules. Ovoid,
adhesion-like surface present just posterior to fifth legs.

Female abdomen 2-segmented, distinct from genital segment.
First segment slightly shorter than second, broader anteriorly than
posteriorly, with slight posterior lateral projections; second segment
with convex lateral margins and slight, bilobed, anal projection.
Caudal rami broad, with 4 plumose setae and 2 naked or very lightly
plumose setules from distal surface, with row of fine denticulations
around base of outermost seta.

Male cephalothorax (fig. 19d) similar to that of female although
somewhat narrower, without dorsal plumosities or distinct projections
indicating previous presence. Free fourth pedigerous segment more
drawn out medially than in female, alae not as large, without medial
constriction present in female. Genital segment (fig. 19e) basically
ovoid, without platelike dorsal part and posterior extensions of female;
fifth leg (fig. 19g) similar to that of female, genital sezgment without
adhesion-like surface present in female. Abdomen 2-segmented,
first segment slightly shorter than second, overlapping second dorsally;
second segment tapered anteriorly, with flatly convex lateral margins
and bilobed anal projection. Caudal rami as in female although
more plumose along inner margin.

Female and male antennule (fig. 20a) 2-segmented, most of anterior
surface of first segment overlapped by ventrally curved lateral surface
of frontal region. First segment slightly more than twice the length
of second, broad proximally, tapered irregularly to narrow distal end,
with small, knoblike projection from posterior distal surface, with 15
naked or lightly plumose setules from anterior ventral and distal
ventral surfaces in female, male with 19 naked or lightly plumose
setules. Second segment subrectangular, with 1 naked setule from
median dorsal surface, second from distal posterior surface, and 8 or 9
naked setules from distal surface. Female antenna (fig. 206) 3-seg-
mented, situated medial and posterior to antennule base. First

No. 3574 COPEPOD CRUSTACEANS—LEWIS dl

bos mm

Ficure 19.—Elytrophora brachyptera Gerstaecker, 1853, immature adult, dorsal view:
a, female; b, male; ¢, lateral posterior cephalothoracic region and free fourth pedigerous
segment. Ventral view: d, female free fourth pedigerous segment, genital segment
(showing attached spermatophore), abdomen, and caudal ramus; ¢, male free fourth
pedigerous segment, genital segment, abdomen, and caudal ramus; f, right fifth leg and
genital fold of female; g, male right fifth leg.

52 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

O1Amm8o

Ficure 20.—Elytrophora brachyptera Gerstaecker, 1853, immature adult, ventral view:
a, left antennule; b, oral region, left side showing antenna (female), postantennal process,
mouth cone, mandible, maxillule, and postoral process; c, male right antenna (lateral
view); d, right maxilla; e, membrane on right maxilla; f, female left maxilliped; g, male
left maxilliped; h, female sternal furca, interpodal plate of first thoracic leg (ip) and
projection between interpodal plates of first and second thoracic legs; 7, male sternal
furca.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 53

segment short, irregular, with triangular projection from lateral
posterior surface; second segment approximately 4 times the length
of first, broader proximally than distally. Third segment fused with
clawlike terminal process, with flabby, setalike accessory process
from proximal surface and elongate, setalike accessory process from
medial third, at break in sclerotization suggesting distal end of seg-
ment and proximal end of terminal process. First segment of male
antenna (fig. 20c) similar to that of female although triangular pro-
jection smaller; second segment with pair of minute, lappet-like
projections medially and small adhesion surface (not figured) on slight
swelling of distal posterior surface. Third segment separable from
clawlike terminal process, bearing same accessory processes as in
female; terminal process strongly curved distally, with small, pointed
secondary projection proximally.

Female and male mandible (fig. 200) 2-parted, first part short,
tapered distally, with indistinct break at beginning of taper. Second
part elongate, flattened distally, with indistinct break just proximal
to flattened part, distal inner surface with 12 denticulations. Female
and male postantennal process (fig. 205) consisting of pair of nodules,
each with several hairlike processes, just lateral to padlike swelling
situated lateral to antenna base. Female and male postoral process
(fig. 206) spinelike, with broad base and laterally curved tip, not as
elongate as in mature adult specimens. Female and male maxillule
(fig. 206) nodular, bearing 2 setules and 1 seta. Female and male
maxilla (fig. 20d) 2-segmented, situated lateral to postoral process.
First segment slightly less than four-fifths the length of second,
excluding terminal process; second elongate, with small, fan-shaped
projection (fig. 20e) from outer surface medially, 2 additional mem-
branes present distally; terminal processes setalike, innermost more
than twice the length of outer, with membranous band and fine plu-
mosities along margin.

Female maxilliped (fig. 20f) 2-segmented, situated posterior and
shghtly medial to maxilla base. First segment strongly developed,
with pair of small, shelflike projections from posterior medial surface;
second segment small, distinct from long, clawlike terminal process,
bearing setalike accessory process from inner distal surface. Male
maxilliped (fig. 20g) similar to that of female although shelflike pro-
jections of first segment smaller and proximal end of segment more
complex, with heavily sclerotized, clawlike projection from proximal
inner surface.

Female sternal furca (fig. 20h) V-shaped, overlapping interpodal
plate of first thoracic legs. Male sternal furca (fig. 207) similar to
that of female although tines not as regular and not appearing as
heavily sclerotized.

54 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

Thoracic legs I-IV differing from those of adult specimens in nature
of segment divisions and minor armature elements (e.g., division
between first and second segments of protopodite of second thoracic
leg indistinct in female; plumosities on ramal segments not as exten-
sive). For nature of legs and armature see figures 21a-f and table
8. Small, platelike projection present just posterior to interpodal
plate of first thoracic legs, projection bearing pair of minute nodules
from median distal surface.

TaBLE 8.—Armature of thoracic legs I-IV of the female and male of
Elytrophora brachyptera Gerstdecker, 1853

Inter- Protopodite Exopodite Endopodite
Leg |} Surface Dodal! |i s_ vel PTs DAA Let be LAR oC AAS 5 DAD AAIL DD Re Oe
fae 1 2 1 2 3 tildes 3
I | Outer s,p rh§ dm,3dmH Cc
Inner p e 4P 3p
IL | Outer m,p | m,d,dmH | dmH 2dmH ,Q Cre ae
m
Inner Sa m,s | ¢,P cP c§,5P Pater 4P
III*} Outer fm,dm,P dm,dmH | dm,fmH 3fmH,2P ec |e 2P
m
Inner dj,2d,5s, ee Ce c§,3P Pa cr2 ok
P,m,4s
IV*| Outer s§,p dm,fmH fm,dm,fmH eee c | ¢,3pt
jm,
Inner fm,s§ e§ ,4pt pt| pt

*Numerous, hairlike processes on protopodite not tabulated.

ee re ee

§Not present in immature adults.

Discuss1on.—The measurements, descriptions, and figures of
the 2 adult stages of Elytrophora brachyptera indicate that a mor-
phological change takes place in the transition from the immature to
the mature condition. This, however, is not the first time that
this has been indicated. Heegaard (1955) points out some of the
differences between his immature ‘‘Atlantide”’ material and figures
of the adult E. brachyptera. Among these he indicates (p. 46) that
the sternal furca in the adult “is delicate with two long pointed
branches with a V-shaped incision between them. In the young
‘Atlantide’ specimens the furea is not fully grown and _ therefore
more plump in the peduncle, shorter in the branches which are
more blunt at their points and together shaping a U instead of the
V in the adult.” This age variation, which is present to a lesser
degree in the immature adult specimens used for this description,

Ficure 21.—Elytrophora brachyptera Gerstaecker, 1853, immature adult, right thoracic
legs, anterior view: a, first; b, second segment of exopodite of first; c, second; d, third;
e, exopodite of third (posterior view); f, fourth.

50

0.05mm8a*

b
Orl
E
=
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, oO
YY
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& ip
: iv pre
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Oo (FZ

COPEPOD CRUSTACEANS—LEWIS

Ss
O3mm&ca

Wy
USS

NO. 3574

56 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

can also be found in many other characteristics. The presence of
plumosities on the dorsal surface of the cephalothorax in immature
adult (with spermatophore) female specimens is suggested only by
small, spikelike irregularities in ovigerous female specimens. The
dimensions of the female genital segment become larger as egg pro-
duction begins; the angle changes at the point where the genital flap
projects, and the flap becomes distinct from the segment in the ma-
ture adult female. The projections on the first abdominal segment
of the female greatly enlarge in the transition from the immature to
mature adult condition. The claw-containing process on the prox-
imal inner surface of the male maxilliped is poorly sclerotized: the
claw projects in the immature state while it is heavily sclerotized
and is covered by the cuticle in the mature adult. The described
differences in the armature of the thoracic legs are more difficult
to reconcile although specimens collected from the same species of
host in the same geographic area (Line Island specimens) exhibit this
variation. Most of the change, however, occurs in the genital seg-
ment, the abdomen, and the amount of sclerotization. These var-
iations are evident not only in the material used for this description
but also in specimens identified as EL. brachyptera by C. B. Wilson
(USNM 78931). It is suggested, therefore, that some of the char-
acteristics previously used in the taxonomy of the species in the genus
should be reexamined on the basis of the maturity of the specimens.

Gloiopotes Steenstrup and Liitken, 1861

DraGcnosis.—Cephalothorax ovoid, consisting of cephalon and first
4 thoracic segments, with spines or hairlike processes, or both, on
dorsal surface. Free fourth pedigerous segment with alae. Female
genital segment with pair of lobate projections posteriorly. Fifth
legs large, heavily sclerotized, projecting past genital segment proper,
either rodlike or lobate; sixth legs, if present, strongly reduced.
Abdomen elongate, 2-segmented, caudal rami rodlike. Antennule
2-segmented; antenna 3-segmented, with clawlike terminal process,
male terminal process modified. Mandible rodlike, distal surface
slightly flattened, with 12-13 denticulations along inner surface.
Maxillule nodular, with several setalike processes; maxilla 2-seg-
mented. Postantennal process consisting of plate bearing 1 or more
spinelike projections and 2 nodules, each with several hairlike proc-
esses; postoral process spinelike, bifurcate, or simple. Maxilliped
2-seemented, prehensile. Thoracic legs J-IJI biramous although
endopodite of first leg reduced; fourth thoracic leg uniramous. Ramal
count of first 4 thoracic legs 2-2, 3-3, 3-2, 3; second segment of exopo-
dite of first leg tipped by 1 simple and 2 bifid spines, bifid spines with
spinelike accessory process.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 57

Gloiopotes huttoni (Thomson)
Fiaures 22-24

Lepeophtheirus huttont Thomson, 1889, p. 354, pl. 28, fig. 10a—c; pl. 29, fig. la-m.

Gloiopotes huttoni (Thomson) Bassett-Smith, 1899, p. 458.—Hewitt, 1964a, p.
86, figs. 1-16— Yamaguti, 1963, p. 104.

Gloiopotes costatus Wilson, 1919, p. 313, pl. 21; 1937, p. 429.—Yamaguti, 1963
p. 103.

Caligus longicaudatus Marukawa, 1925, p. 1243, fig. 2396; 1947, p. 927, fig. 2654.

Gloiopotes watsoni Kirtisinghe, 1934, p. 167, figs. 1-17.—Yamaguti, 1963, p. 104,
pl. 126, fig. 1.

Gloiopotes species Yamaguti, 1936b, p. 4, pl. 2, fig. 20; pl. 3, figs. 21-35.

Gloiopotes zeugopteri Rao, 1951, p. 248, figs. 1-15.

Gloiopotes longicaudatus (Marukawa) Shiino, 1954b, p. 278, figs. 1-2; 1957a, p.
364; 1958, p. 105; 1959a, p. 348.—Heegaard, 1962, p. 174, figs. 151-153.—
Ho, 1963, p. 87, figs. 6-10.—Shiino, 1963a, p. 343.— Yamaguti, 1963, p. 104,
pl. 125, fig. 1—Kirtisinghe, 1964, p. 87, figs. 102-103.

?

Hosts AND DISTRIBUTION.—24 host records:

location

New Zealand

Australia

Indian Ocean

Eastern Pacific

South Pacific

Western Pacific

hosts
Histiophorus herschelli
Makaira mitsukurit
Marlina zelandica
Istiompax australis
Histiophorus brevirostris
H. gladius
Xiphias zeugoptert
Makaira indica
M. mazara
Tetrapturus mitsukurit
T. marlina
“Swordfish”
Istiophorus greyi ‘‘Marlin”’
“Black Marlin”
Makaira audax
Makaira species
NXiphias gladius
“Marlin”
Tetrapturus mitsukurit
Parathynnus sibt
M. mazara
Istiophorus orientalis
Xiphias gladius

reference
Thomson, 1889
Hewitt, 1964a

Heegaard, 1962
Bassett-Smith, 1899
Kirtisinghe, 1934
Rao, 1951
Kirtisinghe, 1964

Shiino, 1958, 1959a
Wilson, 1919
Wilson, 1937

Shiino, 1963a
Wilson, 1929?
Yamaguti, 1936b
Shiino, 1954b

Ho, 1963

MareriaL.—Two females and 2 males (USNM 112879) collected
by D. W. Strasburg (USFWS) from external surface of Makaira ampla
(Poey) captured by rod and reel off Kona, Hawaii. Thirteen females

| and 1 male (USNM 112880) collected by E. C. Jones (USFWS) from

external surface of Makaira audax (Philippi) captured off Waianae,
Hawaii. Two females and 2 males (retained by author) from external
surface of Makaira audaz (Philippi) captured off Waianae, Hawaii.

58 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Two females and 2 males (retained by author) from either Makaira
ampla? (Poey) or Istiompax marlina (Jordan and Hill) examined at
Honolulu Fish Market (USFWS). ‘Two females and 1 male (retained
by author) from external surface of Makaira ampla? (Poey) captured
in Hawaiian region (USFWS).

MEASUREMENTS.—(In mm) 18 females and 6 males:

female male

mean range mean range

Total length, excluding caudal setae 12.02 10.20-12.90 10.25 9. 53-10. 95
Length of cephalothorax, including

frontal region 5.81 5.08- 6.08 463 4.35- 4. 88
Width of cephalothorax 5.06 4.35- 5.638 4.00 3.60 4. 28
Length of genital segment, excluding

fifth legs 3.21 2.40 3.60 2.16 1.95— 2. 25
Width of genital segment 3.038 2.70—- 3.30 1.94 1. 88— 2.03
Length of abdomen 2.29 1.67—- 2.55 1.83 1.52 2.04
Length of alae (male N=5) 157697 1, 838= 92) 11 SIN 08 10) 9S eas
Length of fifth leg, measured along

outer margin (male N=5) 1.61 4-178 1.56" 1. 30-1. G3
Length of caudal rami 106 0.89- 1.30 1.23 1.04 1.37
Length of egg strings (12 strings) 3.34 2.40- 4. 65

DescripTIon.—Female cephalothorax (fig. 22a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region distinct,
with small, median, knoblike projection and frilled membrane on
anterior margin; heavily sclerotized articulation surfaces (articula-
tion with cephalothorax proper) present on posterior surface. Lat-
eral cephalothoracic margins flatly convex although slightly irregular,
with narrow frilled membrane and row of plumosities, membrane
extending around posterior end of broadly curved posterior lateral
cephalothoracic region, terminating adjacent to origin of posterior
sinus. Posterior sinus (fig. 22c) irregularly U-shaped, open end
constricted, lateral margin formed by heavily sclerotized extension
of median cephalothoracic region, small, lappet-shaped dorsal projec-
tion present at anterior end of sinus in addition to small membrane
along lateral margin of sinus and second, posteriorly projec ting mem-
brane, from dorsal lateral surface. Median cephalothoracic region
extending to or slightly past posterior end of lateral cephaloth oracic
regions, margin irregular. Dorsal surface of cephalothorax with
numerous hairlike projections laterally, giving fuzzy appearance; 3
rows of long, slender, bifid plumosities present on lateral regions,
lateral to ocular region. Dorsal surface also with numerous spinules,
primarily on median cephalothoracic region, extending around ocular
region and on posterior end of median cephalothoracic region. Dorsal
cephalothoracic grooves distinct, major grooves forming irregular H.
Eyes distinct, in anterior third of cephalothorax.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 59

Ficure 22.—Gloiopotes huttoni (Thomson, 1889), dorsal view: a, female; b, male; c, poste-
rior portion of female cephalothorax and free fourth pedigerous segment showing ala and
armature; d, male, same. Ventral view: , female genital segment, fifth leg, abdomen,
and caudal ramus; f, male genital segment, fifth leg abdomen, and caudal ramus.

60 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Female free fourth pedigerous segment less than half the width of
cephalothorax, tapered sharply anteriorly and posteriorly from widest
point, at fourth leg attachment. Alae broad anteriorly, tapered (in
figured specimen) to narrow, sharply rounded posterior end tipped
with single spinule; alae rounded posteriorly in some specimens, sub-
rectangular in others. Alae extending posteriorly, over anterior
region of genital segment, bearing several spinules dorsally; medial
lateral surface semimembranous.

Female genital segment (fig. 22e) indistinctly separable from fourth
pedigerous segment, broad medially, with large, lobate projection
from each lateral posterior surface, lateral surface of segment rough-
ened adjacent to lobes. Genital segment bearing strong, heavily
sclerotized fifth legs projecting from ventral lateral posterior surface,
at base of lobate projections; legs extending posteriorly and laterally
to end of abdomen, with row of spinules on distal ventral surface and
on indentation in distal medial region.

Female abdomen 2-segmented, distinctly separable from genital
segment. First segment subrectangular, approximately half the
length of elongate second segment; second segment with club-shaped
swelling on dorsal surface. Both segments with numerous spinules.
Caudal rami elongate, appearing 2-parted although “division” an
indentation in lateral surface (rami not 2-segmented as indicated by
Heegaard, 1962, p. 174). Distal lateral surface of rami with single
spine, distal ventral surface with 1 and distal surface with at least 3
spines; several spinules present on remaining ramal surface (not
figured).

Cephalothorax of male (fig. 226) similar to that of female although
number and arrangement of spinules on dorsal surface different. Alae
of free fourth pedigerous segment (fig. 22d) primarily lateral expan-
sions, without narrow, posterior extension present in female; with
semimembranous trailing edge. Genital segment (fig. 22f/) distinct
from fourth pedigerous segment, ovoid in outline, without lobate
posterior projections. Heavily sclerotized fifth legs projecting from
posterior ventral surface, without distal medial indentation present
in female, with small spinules and bearing 3 spinules from distal sur-
face. Abdomen of both sexes similar in general outline although male
without club-shaped dorsal swelling of second segment. Caudal
rami similar to those of female although inner terminal process longer.

Female and male antennule (fig. 23a) 2-segmented, first segment

Ficure 23.—Gloiopotes huttoni (Thomson, 1889), ventral view: a, left antennule; b, female
left antenna; c, male left antenna and postantennal process (pap); d, female left post-
antennal process (*=see text); ¢, female oral region, left side showing mouth cone, man-
dible, maxillule, and postoral process; f, male, same, right side; g, right maxilla (pop=
postoral process; a=adhesion pad, not present in male); h, left maxilliped.

NO. 3574

COPEPOD CRUSTACEANS—LEWIS

61

62 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

approximately 1% times the length of second, tapered from broad
proximal to narrow distal end, bearing approximately 22 lightly
plumose setae from anterior ventral and distal ventral surfaces.
Second segment elongate, rod shaped, with naked setule from distal
posterior surface, approximately 7 from distal surface. Female an-
tenna (fig. 23) 3-segmented, situated posterior and medial to anten-
nule base. First segment subtriangular, small, with roughened,
triangular projection from proximal posterior surface. Second seg-
ment almost twice the size of first, with small, lappet-like anterior
distal projection. Third segment indistinctly separable from clawlike
terminal process, with small, knoblike projection proximally, projec-
tion bearing setule-like accessory process; second setule-like accessory
process present at junction of segment and terminal process. Male
antenna (fig. 23c) 3-segmented, first 2 segments similar to those of
female, third with larger accessory processes and bifurcate terminal
process.

Female and male mandible (fig. 23¢) indistinctly 3-parted, rodlike; third
part flattened, distal inner surface with 12 denticulations. Small,
bilobed cuticular flap present in female adjacent to mandible base.
Postantennal process of figured female (fig. 23d) situated lateral and
slightly posterior to base of antenna, consisting of platelike region of
heavy sclerotization bearing large spine on lateral posterior surface,
smaller spine on inner posterior surface, and bifurcate spinous pro-
jection on posterior inner surface. Platelike region also bearing 2
nodules on medial surface, each with several hairlike projections;
additional nodule present just posterior to process, with several hair-
like processes. Other female specimens with condition similar to that
of male or with various combinations of spines (see discussion).
Small, spinelike projection present adjacent to anterior inner surface
of postantennal process, arising from edge of depression associated
with base of antenna. Male postantennal process (fig. 23¢) similar to
that of figured female although lacking smaller of 2 single spinelike
projections. Female postoral process (fig. 23¢) bifurcate, each ramus
spinelike; additional, minute, sharply pointed projection present
just lateral to bifurcation. Male postoral process (fig. 23f) spinelike,
not bifureate as in female. Possible remnant of postoral adhesion
pad present in female as minute, oblong protrusion just posterior to
postoral process and medial to maxilla base. Female maxillule
(fig. 23e) nodular, bearing single setule and cluster of hairlike processes.
Male maxillule (fig. 23f) nodular, with 2 or 3 setules. Female and
male maxilla (fig. 239) 2-segmented, first segment approximately
three-fourths the length of second, slender, width varying little
throughout length. Second segment narrow proximally and distally,
swollen medially, with indistinct membrane on medial inner surface;

No. 3574 COPEPOD CRUSTACEANS—LEWIS 63

distal surface bearing 2 saber-shaped processes, inner longer than
outer, both with fine membranes along margins.

Female maxilliped (fig. 23h) 2-segmented, situated posterior and
slightly medial to maxilla base. First segment strongly developed,
lobate proximally, widest medially, tapered to slightly narrower
distal end, without distinct irregularities. Second segment short,
distinct from clawlike terminal process, with single, setule-like ac-
cessory process on inner surface, at junction of segment and terminal
process. Male maxilliped similar in outline to that of female; first
segment with 2 minute, lappet-like projections on inner surface,
second segment with second setule-like accessory process proximally.

Sternal furca of female and male (fig. 24a) situated between and
slightly posterior to maxilliped bases. Process basically bifurcate,
arising from oval platelike area of heavy sclerotization, with spine-
like projection on either side, just proximal to apex of bifurcation.
Tines of bifurcation angled outward, bifurcate distally, each ramus
of distal bifurcation bluntly rounded. Cuticular flap present on
either side of sternal furca, with pointed projection laterally in both
female and male, with rounded projection medially in female.

For nature of armature and legs, see figures 24b-g and table 9.

TABLE 9.—Armature of thoracic legs I-IV of the female and male of
Gloiopotes huttoni (Thomson, 1889)

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal |e ee eee
Plate 1 2 1 2 3 1 2 3
I | Outer Sss,D h d* 2
Inner Dp c 3P,p,3dH p’
II | Outer m,p’ m,fmHf |} dH dH,H,Q,P| C c c,4P
m
Inner S32 m,s ese. ee 4p P c,2P Zi
Iit | Outer $,m,p H H 3H,2P Cc c,2P
m
Inner 2ss,P, m, iP. Cie c,3P e2P 2P
2s,
IV | Outert h,p || rh,3h, D,H D,3H,D
D,H

,

*Denticulations in clump in female, in row in male.

+ Spine denticulated in male, not in female.

$} Numerous spinules on protopodite and first segment of exopodite not tabulated. Spinule arrangement
slightly different in two sexes.

§ Elements between rami.

|| Elements in middle of segment, not on edge.

Discussion.—Gloiopotes huttoni (Thomson, 1889) exhibits a wide
variation in many characteristics. This variation may be a natural
variation or the result of breakage and regrowth of some of the heav-
ily sclerotized parts. The most striking example of this, in the
Hawaiian specimens, is the variation in the spines associated with
the postantennal process. In the figured female specimen (fig. 23d),

64 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

———
O.5mmPc

Ficure 24.—Gloiopotes huttoni (Thomson, 1889): a, sternal furca, ventral view (male with-
out pads lateral to furca). Thoracic legs, anterior view: J, right first; c, second segment
of exopodite of right first; d, endopodite of right first; e, right second; f, right third; g, left
fourth.

NO. 3674 COPEPOD CRUSTACEANS—LEWIS 65

the process has a single, posteriorly directed spine, an inwardly di-
rected bifid spine, and a single spine arising between the preceding
two. In a second female, from the same host specimen, the left
postantennal process is similar to that figured by Shiino (1954b, fig.
le) and by Hewitt (1964a, fig. 13), having only the posteriorly di-
rected spine and the bifid inwardly directed spine. Inthe same female
specimen, the postantennal process on the right side hasa posteriorly
directed spine and a bifid inwardly directed spine, but the tip of the
anteriormost tine of the latter is bifid, giving an appearance similar
to that shown in figure 23d although the bifurcation is not as well
developed. Many of the characteristics used to separate the species
within the genus Gloiopotes have been shown to be variable (Hewitt,
1964a). Hewitt, however (p. 93), states that “the shape of the
plates [alae] on the fourth thoracic [fourth pedigerous] segment does
not appear to vary but they may be inclined at various angles . . .”’.
The figured female specimen (figs. 22a, c) exhibits alae with a posterior
projection while the alae of some of the other Hawaiian specimens do
not have this projection and are rounded posteriorly or are almost
flat. It is felt that the use of alae shape as the primary means of
distinguishing between species is open to question, as is the use of
_ the inclination of these plates and the amount and position of plu-
-mosities on the dorsal surface of the cephalothorax (Hewitt, 1964a).
- Hewitt also discusses the variation in the number of spines present
in the material that he had available for study. This variation is
exhibited by the Hawaiian specimens and, from an examination of
material in the U.S. National Museum, appears to be characteristic
of members of the genus. This author feels that spines may be of
value, but only if a complete series is either absent or present (see
discussion of G. ornatus below). One of the problems in using this
characteristic, however, is that many of the spinules or even small
spines are not figured or described in the literature (see discussion of
— G. auriculatus below.)

An examination of the type material of G. costatus Wilson, 1919
(USNM 49772, 49773, not 51040 and 51041 as Wilson indicates
_[p. 313]), and other material identified as this species, indicated that
_ the variation that prompted Wilson to erect this species is also found
in the Hawaiian specimens of G. huttoni (the variation in the number
and position of spinules and the shape and inclination of the alae).
One of the differences used by Shiino (1954b, p. 278) to separate
G. longicaudatus (Marukawa) from G. costatus Wilson is the tripartite
spine figured by Wilson on the first segment of the exopodite of the
third leg of G. costatus. The female and male cotype specimens
used by Wilson for his original description comprised a copulating
pair, and the original specimens were in copulo until the present

2262301" 65

66 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

author separated them. An examination of this material suggests
that the original figures were made while the pair was in copulo as,
among other characteristics, the spine on the first segment of the third
leg exopodite is bipartite. An examination of the specimens identified
as G. longicaudatus by Ho (1963) and deposited in the U.S. National
Museum (USNM 111247, 111248) shows the same variation present
in the Hawaiian specimens of G. huttona and the variation described
by Hewitt (1964a) for the New Zealand specimens of this species.

Based upon these examinations and upon Hewitt’s discussion, it
is suggested that G. huttona is the species of Gloiopotes that is found
on billfishes in the Pacific. G. ornatus Wilson, 1905b, found on
billfishes in the Atlantic, appears to be the counterpart of G. huttoni
(e.g., Shiino, 1959a). The primary difference between these two
species is the presence of a row of spines on the lateral surface of
the genital segment of G. ornatus (absent in G. huttont).

Barnard (1957) describes a species of Gloiopotes, G. auriculatus,
from a Striped Marlin at Mossel Bay, South Africa. He indicates
that it is close to G. ornatus Wilson and figures the postcephalothoracic
region of both the female and the male. The figure of the female,
however, does not show the series of spines on the lateral surface of
the genital segment, the diagnostic characteristic for G. ornatus.
Mossel Bay is at the southern tip of Africa and G. auriculatus is
bordered on the Indian Ocean side by G. huttoni and on the Atlantic
side by G. ornatus. The association of G. auriculatus with the other
two species cannot be determined from the literature but offers an
intriguing zoogeographical problem.

Gloiopotes hygomianus Steenstrup and Liitken

Figures 25-27

Gloiopotes hygomianus Steenstrup and Liitken, 1861, p. 363, pl. 5, fig. 9—Bassett-
Smith, 1899, p. 458.—Stebbing, 1900, p. 670, pl. 74, fig. A.—Shiino, 1960b,
p. 538, figs. 4-6.— Yamaguti, 1963, p. 103, pl. 124, fig. 3.

Gloiopotes species Bonnet, 1948, p. 7.

DISTRIBUTION AND HOSsTS.—4 hosts:

locality hosts reference
Atlantic Unknown Steenstrup and Liitken, 1861
Pacific “Albacore” Stebbing, 1900
Acanthocybium solandri Shiino, 1960b
Hawaii Acanthocybium solandri Bonnet, 1948

MarTertau.—One female and 1 male (USNM 112881) collected by
E£. C. Jones (USFWS) from Acanthocybium solandri (Cuvier) captured
in the Hawaiian area. One female and 1 male (USNM 112882) from
the caudal fin of A. solandri (Cuvier) captured near the Line Islands

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 67

(USFWS). Two females and 1 male (USNM 112883) from A. solandri
(Cuvier) captured near Washington Island (Line Islands) (USFWS.
Two females and 1 male (retained by author) from external surface
of A. solandri (Cuvier) captured near Fanning Island (Line Islands)
(USFWS).

MEASUREMENTS.—(In mm) 6 females and 4 males:

female male

mean range mean range
Total length, excluding caudal setae 14.33 13.65-15.15 9.92 9. 38-10. 35
Length of cephalothorax, including
frontal region 6.85 6.68 7.18 4.82 4.50 5.25
Width of cephalothorax 5: 44. 5. 25= 5. 78 ~ 3532" 3. 08="3: 538
Length of genital segment, including
posterior projections but not fifth

legs 5.37 4.73- 6.15 1.86 1.73— 2.03
Width of genital segment 2.99 2.63—- 3.30 1.63 1.58- 1.65
Length of abdomen 3.15 2.89- 3.37 1.69 1.52- 1.78
Length of alae 3.68 3.44 4.00 1.56 1.52- 1.59
Length of fifth leg, measured along

outer margin ISS 1a U4 Giver AS S0= 52
Length of caudal rami 1a le 26= 0559) (vl oe. he 4i—. 1. 78
Length of egg strings (4 strings) 12.32 10. 95-13. 88

Description.—Female cephalothorax (fig. 25a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region distinct,
with pair of narrow membranes along median anterior edge. Lat-
eral cephalothoracic margins flatly convex, with narrow membrane
and row of fine, hairlike plumosities, membrane extending around
posterior lateral cephalothoracic regions to beginning of posterior
sinus. Posterior sinus ovoid, lateral surface heavily sclerotized, bear-
ing small, posteriorly projecting membrane from posterior half and
fine membrane along all of margin, membrane projecting across
most of sinus. Anterior end of sinus sharply rounded, with small,
lappet-like, posterodorsally projecting membrane; inner surface irreg-
ular. Median cephalothoracic region slightly irregular, extending
almost to end of lateral cephalothoracic regions. Dorsal surface of
posterior lateral cephalothoracic region with row of fine, hairlike
plumosities along posterior lateral surface. Dorsal cephalothoracic
grooves distinct, forming irregular H although cross groove V-shaped,
with apex directed anteriorly; anterior longitudinal grooves of H
turning inward sharply just anterior to ocular region. Ocular region
small, consisting of 2 darkly pigmented cup-shaped regions contiguous
on median longitudinal axis of body, each with spherical lens.

68

VOL. 121

PROCEEDINGS OF THE NATIONAL MUSEUM

|
|

SS aa,

a, female;

Ficure 25.—Gloiopotes hygomianus Steenstrup and Litken, 1861, dorsal view:
b, male. Ventral view: c, female genital segment, abdomen, and caudal ramus; d, male,

same; ¢, tip of female left fifth leg; f, male, same; g, male left sixth leg.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 69

Female free fourth pedigerous segment, excluding ala, slightly
more than one-third the width of cephalothorax, indistinctly separable
from genital segment. Ala broad, extending laterally past lateral
edge of genital segment, with 2 large, subrectangular posterior pro-
jections overlapping most of genital segment proper. Apex of sinus
between lobes of ala flatly concave, posterior edge of lobes sharply
angled laterally, with small, lobate projections medially.

Female genital segment (fig. 25c) convex anteriorly, at junction
with fourth pedigerous segment, lateral surface smoothly irregular,
with row of spinules along posterior two-thirds of segment. Posterior
end of segment drawn out into pair of elongate lobes extending pos-
teriorly to end of abdomen, lobes with several rows of spinules and mi-
nute spicules (latter not figured). Median lateral surface of lobes
also bearing laminate fifth leg projecting laterally and posteriorly.
Distal surface of fifth leg (fig. 25e) bearing row of spinules and 2
plumose setules.

Female abdomen 2-segmented, incompletely separated from gen-
ital segment dorsally and ventrally. First segment slightly more
than half the length of second, with distinct dorsal swelling; second
segment elongate, tapered slightly toward distal end, with pair of
minute spinules on each side in anterior half of segment. Fine frilled
membrane present on ventral-lateral posterior surface, at junction
with caudal rami. Caudal rami elongate, rodlike, with small lateral
indentation in proximal half, indentation bearing 2 plumose setules;
distal end with 3 or 4 setules, longer 2 plumose.

Male cephalothorax (fig. 255) generally similar to that of female
although more elongate. Apex of posterior sinus without lappet-
like membrane although remaining membranes present; cross groove
of major dorsal grooves with indistinct apex. Ala of free fourth
pedigerous segment with angular posterior edge but without lobes
present in female. Male genital segment (fig. 25d) more orbicular
than that of female, without lappet-like projection but with spikelike
fifth leg extending to middle of second abdominal segment, inner
dorsal surface of leg projecting medially as narrow, heavily sclero-
tized flap. Distal end of fifth leg (fig. 25f) with 3 spinules, proximal
inner surface with minute, plumose setule. Sixth leg (fig. 25g) evi-
dent as pair of spinules and single minute, plumose setule adjacent
to junction of abdomen and genital segment. Abdomen 2-segmented,
distinct from genital segment dorsally, indistinctly separable ven-
trally. Second segment approximately 2% times the length of first,
first without swelling present in female. Caudal rami similar to
those of female.

Female and male antennule (fig. 26a) 2-segmented, attached to
lateral-anterior ventral surface of cephalothorax and adjacent sur-

70

S|
0.3mm.? ) (A

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|}_+—_+——_| @]

O3mmg¢ad

é

O5mm¢ ¢

VOL. 121

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 71

face of frontal region. First segment slightly shorter than second,
broad proximally, narrow distally, distal half of anterior ventral and
anterior surfaces with approximately 24 lightly plumose setae and
setules. Second segment elongate, rodlike, with single, naked seta
from posterior surface in distal third of segment and 8 naked setae
from distal surface. Female antenna (fig. 266) 3-segmented, situ-
ated medial and posterior to antennule base. Antennal base in
close proximity to several flaplike projections of ventral cephalo-
thoracic cuticle. First segment small, subtriangular in ventral view,
with spike-shaped projection from posteriormost surface; second seg-
ment irregular, outer surface shorter than inner, proximal and distal
surfaces irregular. Third segment separable from clawlike terminal
process by break in heavy sclerotization, with small, setule-like acces-
sory process proximally, on knoblike projection of segment, and second
small, setule-like accessory process at region of break in sclerotiza-
tion. Male antenna (fig. 26c) with irregular first segment, inner
proximal surface with heavily sclerotized bifurcation; second segment
similar to that of female. Third segment fused with double-clawed
terminal process, bearing setule-like accessory process proximally
and second, long, setule-like process just proximal to proximalmost
claw and arising from slight indentation behind minute, flaplike
cuticular projection.

Female and male mandible (fig. 26e) 4-parted; first part broad
proximally, tapered distally, with flexible joint between first and sec-
ond parts. Second part short, tapered slightly; third part elongate,
approximately equal to combined lengths of first 2 parts. Fourth part
short, flattened distally, inner edge of flattened portion with 13 den-
ticulations. Female and male postantennal processes (figs. 26c, d)
consisting of heavily sclerotized plate lateral and slightly posterior
to antennal base, with large, spike-shaped projection from lateral
surface and small, spike-shaped projection from inner posterior sur-
face; 3 minute nodules present, 2 on plate, third posterior to plate,
each with at least 2 hairlike processes. Male with small, flabby,
conical projection just medial to inner surface of postantennal process,
on ridge of heavy sclerotization extending from lateral anterior margin
of cephalothorax to antennal base (fig. 26c)._ Female and male postoral
process (fig. 26e) large, spinelike, female with small spine arising from
inner proximal surface, process absent on male. Female and male
maxillule (fig. 26¢) nodular, with setule distally. Female with pair
Ficure 26.—Glotopotes hygomianus Steenstrup and Litken, 1861, ventral view: a, left

antennule; b, female left antenna; c, male left antenna and postantennal process; d, fe-

male right postantennal process; ¢, oral region, left side showing mouth cone, mandible

(mdbl), maxillule (ma-1), postoral process, maxilla base (ma-2), and postoral adhesion

process (pa); f, left maxilla; g, female right maxilliped; 4, male left maxilliped; i, sternal
furca.

i2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

of small, knob-shaped projections posterior to mouth cone base,
similar in position to postoral adhesion pads. Female and male
maxilla (fig. 26f) 2-segmented, situated lateral and posterior to post-
oral process. First segment approximately two-thirds the length
of second, proximal end with pair of subconical articulation projections,
segment tapered slightly to distal region. Second segment elongate,
broadest medially, with fine membranous flange on inner medial
surface, with 2 saber-shaped processes from distal surface, inner
more than twice the length of outer, with fine membrane along inner
margin, outer with membranes along both margins.

Female maxilliped (fig. 26g) 2-segmented, situated posterior and
medial to maxilla base. First segment strongly developed; second
segment short, distinct from clawlike terminal process, bearing single,
setule-like accessory process from inner surface, at junction of segment
and terminal process. Male maxilliped (fig. 26h) 2-segmented, prox-
imal end of first segment recurved, terminating in socket-shaped
articulation surface; segment with slightly roughened adhesion surface
on medial inner surface. - Second segment appearing bifurcate distally,
one ramus of bifurcation formed by scoop-shaped terminal process;
segment bearing long, flexible process just proximal to apex of bifur-
cation. Flexible process longer than first segment, possibly repre-
senting elongate accessory process, based on position of origin as well
as absence of small, setule-like accessory process present on female.

Female and male sternal furca (fig. 267) situated between and
posterior to maxilliped bases, consisting of plate bearing 2 widely
separated, bluntly pointed tines and 2 accessory spines, at base of
tines. Pair of lappet-like projections present just posterior and
lateral to furca.

For nature of armature and legs, see figure 27 and table 10.

TABLE 10.—Armature of thoracic legs I-IV of the female and male of
Gloiopotes hygomianus Steenstrup and Liitken, 1861

Protopodite Exopodite Endopodite
Inter-
Leg | Surface podal
Plate
2 1 2 3 1 2 3
I | Outer Ss,p h c c,2P
Inner p c 3P,p,3dH p
II | Outer m,p’ 2m,dH dH dH,mH, c c c,4P
m Q,P
Inner SE m,s Cae Cie 4P P ¢,2P. 2:
III | Outer m,p 3s,H h 3h,2P c 3P
m
Inner 2s, P.m,2s, iP iP 3P ¢,2P e
c
IV | Outert pt d,D,dH | D,dH D,3fim,3dH

*Elements between rami.
t+Element in middle of segment.
{Spinules along inner surface not tabulated.

O.1mm.¢¢o b

Ficure 27.—Gloiopotes hygomianus Steenstrup and Liitken, 1861, right thoracic legs, ante-
rior view: 4, first; b, endopodite of first; c, distal end of second segment of exopodite of first;
d, second; e, third; f, exopodite of third (posterior view); g, fourth; h, distal end of third
segment of exopodite of fourth.

74 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121
Family Caligidae
Dentigryps Wilson, 1913
DiaGnosis.—See Lewis, 1964b.

Dentigryps ulua Lewis

Dentigryps ulua Lewis, 1964b, p. 351, figures 2-4; 12a, e, i; 18a.

Hosts AND DISTRIBUTION.—1 host record:

location hosts reference

Hawaiian Islands Caranz melampygus? Lewis, 1964b

Matrriau.—Two females and 1 male (retained by author) from
external surface of Caranz melampygus? Cuvier and Valenciennes
captured in a fishtrap between Diamond Head and Koko Head, Oahu,
Hawaii, by Samuel Kaolulo.

MEASUREMENTS.—(In mm) 2 females and 1 male:

female male
Total length, excluding fifth legs and caudal setae 6. 60, 6. 83 5. 18
Length of cephalothorax, inchiding frontal region 4. 43, 4. 43 3. 45
Width of cephalothorax 3. 40, 3. 66 2. 92
Length of genital segment, including lobes but excluding
fifth legs 1. 67, 1. 67 0. 93
Width of genital segment 1.70) 70 1. 04
Length of abdomen 0. 61, 0. 76 0. 49
Length of caudal rami 0. 31, 0. 29 0. 29
Length of fifth legs, measured along outer margin 1. 19, 1. 28 0. 99
Length of fifth leg terminal process 0. 05 0. 06
Length of sixth leg, measured along outer margin 0. 15
Length of sixth leg terminal process 0. 14

D£EscrIPTION.—See Lewis, 1964b.

Dentigryps bifurcatus Lewis

Dentigryps bifurcatus Lewis, 1964a, p. 203, figs. 17a-c, e-k, m-q; 18a-c, e-f; 1964b,
p. 356, figs: 5;°12b, f, j; 13b.

DiIstTRIBUTION AND HOoSTS.—5 host records:

locality hosts reference
Hawaiian Islands Acanthurus olivaceus
A, triostegus
sandvicensis Lewis, 1964a
Naso hexacanthus
Fistularia petimba Lewis, 1964b

MareriaALu.—Three males (retained by author) from external
surface of Bodianus bilunulatus (Lacépéde) from the Honolulu
Aquarium. One male (retained by author) from external surface
of Chaetodon fremblii Bennett captured in a fishtrap by Samuel
Kaolulo between Diamond Head and Koko Head, Oahu, Hawaii.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 75

Two immature males (retained by author) from external surface of
Aulostomus chinensis (Linnaeus) captured in a fishtrap by Samuel
Kaolulo between Diamond Head and Koko Head, Oahu, Hawaii.

MEASUREMENTS.—(In mm) 4 males:

Total length, excluding fifth legs and caudal setae 2.15, 2.26, 2.22, 2.29
Length of cephalothorax, including frontal region (N=2) 1.64, 1.62

Width of cephalothorax 1.585 1055, 124891253
Length of genital segment, excluding fifth legs 0.36, 0.36, 0.36, 0.35
Width of genital segment 0.46, 0.41, 0.41, 0.38
Length of abdomen O20. 01 O11. O02
Length of fifth leg, measured along outer margin (N=3) 0.17, 0.19, 0.18
Length of fifth leg terminal process (N= 2) 0.04, 0.04

Length of sixth leg, measured along outer margin (N=3) 0.07, 0.09, 0.07
Length of sixth leg terminal process (N=3) 0.09, 0.08, 0.07
Length of caudal rami 0.13, 0.14, 0.18, 0.14

DescripTion.—See Lewis, 1964a.
Anuretes Heller, 1865

Diaanosis.—See Lewis, 1964a.

Anuretes serratus Shiino
Anuretes serratus Shiino, 1954a, p. 260, figs. 1-2.—Lewis, 1964a, p. 188, figs. 13-14.
DISTRIBUTION AND HOSTS.—2 host records:

locality hosts reference
Japan Xesurus scalprum Shiino, 1954a
Hawaiian Islands Naso hexacanthus Lewis, 1964a

DeEscrIpTion.—See Lewis, 1964a.
Anuretes menehune Lewis

Anuretes menehune Lewis, 1964a, p. 195, figs. 15, 16.
DisTRIBUTION AND Hosts.—2 host records:

locality hosts reference
Hawaiian Islands Naso hexacanthus
N. unicornis Lewis, 1964a

DescripTion.—See Lewis, 1964a.
Lepeophtheirus Nordmann, 1832

DraGnosis.—Cephalothorax consisting of cephalon and first 4
thoracic segments; frontal region distinct, without lunules. Free
fourth pedigerous segment without alae. Genital segment without
projecting plates; fifth legs present on female and male, setiform or
projecting slightly, without ornamentation; sixth legs present on male,
setiform. Abdomen 1-2 segmented. Antennule 2-segmented, an-

76 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

tenna 3-segmented, with clawlike terminal process and frequently,
in male, with adhesion or additional spinelike process. Mandible
rodlike, distal region flattened, with 12 denticulations on distal inner
surface. Postantennal process consisting of 3 nodules, each with
several hairlike processes, and clawlike projection; postoral process
spinelike, often with smaller, spinelike secondary projection on inner
surface in male. Maxillule nodular, with setule-like processes;
maxilla 2-segmented, with 2 saber-like terminal processes. Sternal
furca present. Thoracic legs J-IIT biramous although endopodite of
first legs rudimentary; fourth legs uniramous, typically with 4
segments.

Lepeophtheirus crassus (Wilson and Bere)
Figures 28-30

Gloiopotes crassus Wilson and Bere, in Bere, 1936, p. 590, pl. 5, figs. 109-111; pl. 6,
figs. 125-155.

Lepeophtheirus crassus (Wilson and Bere) Shiino, 1960c, p. 546, figs. 3-4.—Yama-
guti, 1963, p. 72.—Shiino, 1963a, p. 337, fig. 2.

DIsTRIBUTION AND Hosts.—4 host records:

locality hosts reference
Western Atlantic Rhombochirus osteochir Wilson and Bere, 1936
Eastern Pacific ““Remora’”’
Remilegia australis Shiino, 1968a
Indian Ocean Echeneis albescens Shiino, 1960c

Mareriau.—Five females (USNM 112884) from external surface
of Rhombochirus osteochir (Cuvier) taken from Tetrapturus angusti-
rostris Tanaka captured by longline at 21°04.5’N, 173°47.5’E (USF
WS). Two females and 1 male (USNM 112885) from external sur-
face of Rhombochirus osteochir (Cuvier) taken at 20°50.5’N, 167°34.5’E.

Remarks.—Although these specimens were taken well west of the
Hawaiian Islands, the hosts are found in the Hawaiian area. Based
upon this and the wide reported distribution of the copepod, the author
feels that the copepod is a member of the Hawaiian parasitic copepod
fauna.

MEASUREMENTS.—(In mm) 7 females and 1 male:

female male
mean range

Total length, excluding caudal setae 6.36 5.63-7.05 4.80
Length of cephalothorax, including frontal region 3.73 3.53-4.20 3.00
Width of cephalothorax 3.07 2.70-3.38 2.48
Length of genital segment, excluding fifth legs 1.69 0.98-2.06 0.82
Width of genital segment 1.91 1.50-2.18 0.75
Length of abdomen 0.81 0.53-0.98 0.53
Length of fifth leg, excluding setae 0.79 0. 72-0.86 0.40
Length of fifth leg subterminal seta (female N=6) 0.08 0.07-0.11 0. 04
Length of egg strings (4 strings) 2.83 2. 63-3. 15

NO. 3574 COPEPOD CRUSTACEANS—LEWIS W

Duscription.—Female cephalothorax (fig. 28a) ovoid, consisting of
cephalon and first 4 thoracic segments. Frontal region narrow,

eI
0,02mm.o

Ficure 28.—Lepeophiheirus crassus (Wilson and Bere, 1936),dorsal view: a, female; b, male.
Ventral view: c, female genital segment, fifth leg, abdomen, and caudal ramus; d, male,
same; é, male tip of fifth leg; f, male sixth leg.

anterior edge with median indentation and narrow, membranous
flange. Lateral margins of cephalothorax flatly convex except for

78 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

slight medial swelling, posterior lateral corners smoothly rounded,
lateral surface terminating just lateral to beginning of posterior cepha-
lothoracic sinuses, at origin of longitudinal legs of cephalothoracic
grooves. Posterior sinuses distinct, U-shaped, lateral margin heavily
sclerotized, bearing filmy membrane that projects into sinus. Poste-
rior margin of median cephalothoracic region with slight concavity
medially, lateral margins convex. Major cephalothoracic grooves
forming irregular H, posterior longitudinal legs forming line of divi-
sion between median and lateral cephalothoracic regions. Cross
eroove of flattened V-shape, apex pointing anteriorly. Anterior longi-
tudinal grooves extending anteriorly and slightly laterally to indis-
tinct termination lateral and slightly posterior to ocular region.
Ocular region distinct, consisting of 2 heavily pigmented cup-shaped
regions, contiguous on median longitudinal axis of body, each with
spherical lens. Small, heavily pigmented area present between poste-
rior ends of cup-shaped regions and small, heavily sclerotized V-shaped
region present at anterior end of eyes, apex of V extending into depres-
sion formed by inner anterior margins of cup-shaped pigmented region.

Female free fourth pedigerous segment short, width more than
twice the length. Dorsal surface heavily sclerotized, medial posterior
margin of segment distinct dorsally, flatly convex.

Lateral margins of genital segment (fig. 28c) flatly convex, posterior
lateral surfaces forming lobate projections extending posteriorly
almost to end of second segment of abdomen. Fifth legs originating
on posterior ventral surface, just lateral to oviducal opening, at base
of lobate projections. Fifth legs dactyliform although slightly irreg-
ular, not heavily sclerotized, extending posteriorly past posterior end
of lobate projection, bearing 3 plumose setules on dorsal surface in
distal third of leg; additional plumose setule present on ventral sur-
face of genital segment, just lateral to leg base.

Abdomen 2-segmented, distinct from genital segment. First seg-
ment approximately four-fifths the length of second, lateral margins
flatly convex, posterior margin convex ventrally, V-shaped dorsally,
overlapping anterior end of second segment. Lateral margins of
second segment flatly convex, posterior margin slightly irregular,
tapered to bilobed anal region. Caudal rami laminate, length slightly
less than twice the width. Both lateral margins flatly convex, inner
lateral surface plumose distally. Outer distal margin indented,
bearing single plumose seta; inner two-thirds of distal surface rounded,
bearing 4 plumose setae; knoblike projection present on inner distal
surface, bearing minute, spikelike terminal process.

Cephalothorax of male (fig. 286) generally similar to that of female.
Anterior margin of frontal region flatter than in female; V-shaped
region of heavy sclerotization at anterior end of ocular region also

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 79

flatter. Free fourth pedigerous segment as in female. Genital
segment (fig. 28d) slightly longer than wide, lateral margins flatly
convex, posterior surface of segment overlapping dorsal surface of
first abdominal segment slightly. Fifth legs dactyliform, originating
on anterior-medial ventral surface, extending posteriorly and slightly
laterally to posterior medial region of genital segment. Fifth legs
with 2 plumose terminal setules, one plumose subterminal setule, and
one plumose setule from outer dorsal surface approximately two-thirds
the distance from proximal to distal end of leg (fig. 28e). Sixth legs
(fig. 28f) small, somewhat irregular, projecting slightly from posterior-
lateral ventral surface of segment and tipped by 2 plumose setules.
Abdomen 2-segmented, similar to that of female except dorsal poste-
rior projection of first segment not as distinct. Caudal rami some-
what more rectangular in shape than those of female, otherwise
similar.

Female and male antennule (fig. 29a) 2-segmented; first segment
more than twice the length of second, proximal margin broad, ir-
regular, with several small concavities and projections, one concave
depression in posterior margin forming articulation surface for small,
knoblike projection of platelike area on anterior ventral surface of
cephalothorax. Anterior lateral margin of first segment convex,
posterior lateral margin almost straight except distally where poste-
rior lateral surface forms base of small, shelflike extension of distal
surface; distal surface narrow, irregular. Anterior and distal ventral
surfaces with at least 14 plumose setae. Second segment club shaped,
ovoid in cross section, narrow proximally and distally, widest me-
dially. Distal medial region of posterior surface with one naked
seta, distal surface with 11 naked setae and one spikelike process, on
anterior distal corner. Female antenna (fig. 296) 3-segmented, lo-
cated posterior and slightly medial to antennule base. First segment
irregular, proximal end narrow, distal medial region broad, bearing
posteriorly projecting lobe; second segment strongly developed,
slightly shorter than first segment, greatest width slightly more than
four-fifths the length, distal corner projecting slightly as heavily
sclerotized, rounded process (not figured). Third segment and claw-
like terminal process continuous, proximal posterior surface with
small, poorly developed, laminate lobe. Setalike accessory process
present on middle of combined segment and terminal process, pre-
sumably at junction of segment and terminal process. Male antenna
(fig. 29c) 3-segmented; first segment flattened, median third forming
adhesion surface. Second segment well developed, longer than first
segment, most of proximal inner surface forming adhesion surface,
inner distal surface with 2 heavily sclerotized swellings serving as
articulation surfaces for 2 swellings on inner proximal surface of

80 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

iil ter badd 2

b 0.2mm. ? C 02mm 36

& 02mm ¢ Db 'oimm.'s

Z 0.2mm. 2 a 0.2mm. ¢

Ficure 29,—Lepeophtheirus crassus (Wilson and Bere, 1936), ventral view: 4, left antennule;
b, female right antenna;c, male, same; d, right mandible; e, female left postantennal process;
f, male, same; g, female right maxillule and postoral process; h, male, same; 1, female post-
oral adhesion pads, postoral processes, posterior end of mouth cone and maxilliped bases
(mb); 7, male, same.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 81

third segment; posteriormost swelling on second segment with small,
slightly projecting adhesion surface. Third segment and terminal
process continuous, claw shaped, inner surface irregular, appearing
roughened or minutely denticulated distally; single, long, needle-like
accessory process present on inner surface just distal to small swelling
on proximal medial region of segment.

Female and male mandible (fig. 29d) 4-parted, fourth part short,
curved inward smoothly, bluntly rounded distally, inner margin with
12 denticulations although proximal 2 denticulations indistinct in
some specimens. Female postantennal process (figs. 29e, f) consist-
ing of platelike region of heavy sclerotization bearing spikelike
projection and 3 nodules, each nodule with several hairlike processes;
spikelike projection broad proximally, tapering sharply in proximal
half, gradually in distal half, curving medially sharply in male,
slightly in female, sharply rounded, or pointed, distally. Female
postoral process (fig. 29g) bifureate, both tines pointed; male postoral
process (fig. 29h) spinelike, with small spine projecting from middle of
outer surface and small, heavily sclerotized swelling on middle of
inner surface. Female and male maxillule (figs. 299, h) nodular, with
3 setules distally. Female and male maxilla (fig. 30a) 2-segmented,
situated just lateral and posterior to postoral process. First segment
approximately three-fourths the length of second, slender, lateral
margins almost parallel; second segment elongate, widest medially,
medial region with fine membrane and, in male, small, spinelike
projection just distal to membrane. Distal end of second segment
with 2 saber-shaped, flexible processes, innermost approximately 11%
times the length of outermost, inner bearing fine membrane on inner
surface, outer with denticulated, membrane-like process along outer
margin. Male with pair of ovoid adhesion surfaces (fig. 29f) extend-
ing posteriorly and medially from just posterior to postoral processes
to just anterior to maxilliped bases, surfaces distinctly rugose. Female
also with surfaces but not rugose (fig. 297).

Female maxilliped (fig. 306) 2-segmented, situated posterior and
slightly medial to maxilla base. First segment strongly developed,
proximal end lobate, extending into cephalothorax as articulation and
muscle attachment surface. Second segment and terminal process
continuous, length of segment and process slightly greater than half
that of first segment, terminal process clawlike, sharply pointed;
single small, setalike accessory process present on inner surface at
uregular division between segment and terminal process. Male max-
illiped (fig. 30c) similar to that of female although small, irregular
protrusion present on medial inner surface of first segment, protru-
sion receiving distal end of second segment terminal process when

226-321-676

82 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Wye
‘| 005mm

No. 3074 COPEPOD CRUSTACEANS—LEWIS 83

segment flexed. ‘Terminal process of second segment not as sharply
curved as in female, accessory process slightly longer.

Female and male sternal furca (fig. 30d) situated about halfway
between maxilliped bases and interpodal plate of first thoracic leg.
Process consisting of ovoid, heavily sclerotized plate with 2 winglike
anterolaterally projecting structures in addition to U-shaped furca.
Tines of furca either pointed or sharply rounded, outer margins
flatly convex.

For nature of legs and armature, see figure 30 and table 11.

TABLE 11.—Armature of thoracic legs I-IV of the female and male of
Lepeophtheirus crassus (Wilson and Bere, 1936)

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal
Plate 1 2 1 | 2 3 1 2 3
I | Outer ss,p rh 3H,P
Inner p c 3P
II | Outer m,p m,fmH | fmH H,qH,Q, | c¢ c c,3P
m 2P
Inner Sie m,s Ce Ce c,3P 12 ¢c,2P | ¢,3P
IiI | Outer m,P 3s ¢;p’ c,3p’,P c ¢,2p
m
Inner P,s,m,l1s* p,m Cee ¢,3P iP c,4P
IV | Outer s dm,h dm,fmH | 3dm,3fmH

*On both anterior and posterior surfaces of protopodite.
Lepeophtheirus dissimulatus Wilson

Lepeophtheirus dissimulatus Wilson, 1905b, p. 631, pl. 22.—Lewis, 1963, p. 195,
figs. 1-20; 1964a, p. 178, figs. 11-12. [For synonymy, see Lewis, 1964a.]

REPORTED DISTRIBUTION AND HOSTS.—24 host records:

locality hosts reference
Bermuda Epinephalus morio Wilson, 1905b
E. striatus
Mycteroperca apua Linton, 1907
Thynnus pelamys Heegaard, 1943b
Dry Tortugas Lactophrys triqueter Wilson, 1935b
Galapagos Islands Epinephelus labriformis Wilson, 1905b
Mycteroperca species Heegaard, 1943b
Eastern Pacific Bodianus diplotaenis
Epinephelus labrifrons Shiino, 1959d

Merluccius productus
Paralichthys californicus
Hypsopsetta guttulata
Sphyraena argentea

Ficure 30.—Lepeophtheirus crassus (Wilson and Bere, 1936), lateral view: a, maxilla; d,
female maxilliped; c, male maxilliped; d, sternal furca (ventral view). Right thoracic
legs, anterior view: ¢, first; f, second segment of exopodite of first; g, second; A, third; 2,
exopodite and endopodite of third; 7, fourth; , frilled membrane on second segment of
exopodite of fourth; /, frilled membrane on terminal spines of third segment of exopodite
of fourth.

84 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

locality hosts reference
Paralabrax nebulifer
Sphaeroides annulatus

Galeichthys guatemalensis Causey, 1960
Gadus macrocephalus Lewis, 1964a
Hawaii Acanthurus olivaceus

A. dussumiert
A. triostegus sandivcensis
Zebrasoma flavescens
Naso hexacanthus
Chaetodon quadrimaculatus Lewis, 1964a
Mauritania Labrus species sensu lato Brian, 1924
Marertau.—Four females and 4 males (retained by author) from
gill cavity of Parupeneus pleurostigma (Bennett) captured in fishtrap
by Samuel Kaolulo between Diamond Head and Koko Head, Oahu,
Hawaii. One female and 1 male (retained by author) from external
surface of P. pleurostigma (Bennett) captured by trap by Samuel
Kaolulo between Diamond Head and Koko Head, Oahu, Hawaii.
MEASUREMENTS.—(In mm) 4 females and 5 males:

female male

Total length, excluding caudal mean range mean range

setae 2.66 2.538-2.75 2.17 2. 03-2. 25
Length of cephalothorax, includ-

ing frontal region 1.78 1.75-1.83 1.60 1. 52-1. 75
Width of cephalothorax 1.68 1.65-1.70 1.43 1. 40-1. 45
Length of genital segment 0.60 0. 53-0.68 0.36 0. 34-0. 38
Width of genital segment 0.86 0. 78-0.95 0.35 0. 35 (all)
Length of abdomen 0.13 0.12-0.14 0.11 0. 10-0. 12
Length of caudal rami 0.08 0.07-0.09 0.10 0. 08-0. 11

Length of egg strings (2 strings) 1. 00, 1. 48
DEscRIPTION.—See Lewis, 1964a.

Lepeophtheirus? fallolunulus, new species
Figures 31-33

MareriaLt.—One female (holotype, USNM 112886) from the gill
cavity of Naso unicornis (Forskal) speared off Lehua Rock, Nihau,
Hawaii. One immature adult male (allotype, USNM 112887) from
buccal cavity of Naso unicornis (Forskal) speared near ‘‘the blowhole,”’
Oahu, Hawaii.

MEASUREMENTS.— (In mm) 1 female and 1 male:

female male
. 05 2. 25
. 43 1. 30

Total length, excluding caudal setae 3

Length of cephalothorax, including frontal region 1

Width of cephalothorax 1. 30 1. 23
Length of genital segment 1. 08 0. 54
Width of genital segment 0. 85 0. 45
Length of abdomen 0. 15 0. 14
Length of caudal rami 0. 07 0. 08
Length of egg strings 1

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 85

(8 MK/,
fs

fer ui TS dsmm ¢

=

ee
Z a

Os

|=) + ee ee

¢ 0.3mm. 005mm.

Ficure 31.—Lepeophtheirus? fallolunulus, new species, dorsal view: a, female; b, immature
adult male; ¢, posterior cephalothoracic sinus. Ventral view: d, female genital segment,
abdomen, and caudal ramus; ¢, male, same; f, female right fifth leg; g, male, same; h
male right sixth leg.

86 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 87

Descriprion.—Female cephalothorax (fig. 31a) oblong, consisting
of cephalon and first 4 thoracic segments. Frontal region with dor-
sally facing lunule-like modification laterally, with deep incision medi-
ally and fine membrane along anterior ventral surface median to modi-
fication. Lunular modifications (fig. 326) with denticulated, dorsally
facing anterior margin, bearing single setule posteriorly. Division
between frontal region and cephalothorax distinct; 2 large, subtri-
angular projections present, extending anteriorly to posterior region of
lunular modifications from origin on lateral anterior surface of cephalo-
thorax. Posterior two-thirds of median cephalothoracic region raised
dorsally, above lateral cephalothoracic regions. Lateral surface of
cephalothorax forming distinct flange ventrally, bearing fine membrane
along entire length. Posterior sinus (fig. 31c) distinct, slender, with
heavily sclerotized lateral surface, bearing fine membrane along outer
surface. Posterior end of median cephalothoracic region extending
well past posterior end of lateral regions, covering anterior end of free
fourth pedigerous segment. Major dorsal cephalothoracie grooves
running longitudinally, cross groove absent. Free fourth pedigerous
segment slender, without dorsal plates.

Female genital segment (fig. 31d) ovoid, indistinctly separable from
free fourth pedigerous segment. Posterior end of segment irregular,
irregularities primarily in region of oviducal openings. Fifth leg
(fig. 31f) consisting of 5 plumose setules on lateral ventral surface
just lateral to oviducal opening.

Female abdomen 1-segmented, distinct from genital segment.
Caudal rami subrectangular, irregular distally, inner surface plumose ;
distal region with 4 plumose setae and 2 plumose setules.

Male cephalothorax (fig. 31b) differing from female in shape (circu-
lar), in absence of lunular modification of frontal region, in absence of
deep median incision of frontal region, in membrane projecting from
anterior edge of frontal region instead of from anterior ventral surface.
Also differing in concave nature of posterior median cephalothoracic
region, with large, lappet-shaped projection extending over anterior
half of free fourth pedigerous segment and forming attachment surface
of segment. Median cephalothoracic region with fine cuticular exten-
sion dorsally, extension covering concave posterior margin of region
and most of lappet-shaped projection. Free fourth pedigerous

Ficure 32.—Lepeophitheirus? fallolunulus, new species, ventral view: a, left caudal ramus;
b, female frontal region, right side showing lunule-like structure (dorsal view); c, right
antennule; d, female oral region, right side showing antenna, postantennal process, mouth
cone, mandible, maxillule (ma-l), postoral process (pop), postoral adhesion process (pa),
and maxilla base (ma-2); ¢, male oral region, right side showing antenna (stippled),
postantennal process, mouth cone (hatched), mandible (hatched), maxillule (ma-l),
postoral process (pop), postoral adhesion process (pa) and maxilla base (ma-2); f,
right maxilla; g, female left maxilliped; h, male right maxilliped.

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VOL. 121

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 89

segment without plates. Segment-like projection present between
fourth pedigerous and genital segments, separable from fourth pedig-
erous segment (at least superficially) both dorsally and ventrally,
distinct from genital segment; projection without processes. Genital
segment (fig. 3le) barrel shaped, fifth and sixth legs (figs. 31g, h)
evident as series of plumose setules. Abdomen 1-segmented although
with annulations anteriorly. Caudal rami as in female.

Female and male antennule (fig. 32c) 2-segmented, attached to
ventral surface of subtriangular projection from lateral anterior sur-
face of cephalothorax. Anterior and distal ventral surface of first
seement with 20 plumose setae and setules, median dorsal surface with
2. Distal-medial posterior surface of elongate second segment with
single naked setule, distal surface with 10. Female antenna (fig. 32d)
3-segmented, situated posterior to subtriangular projection associ-
ated with antennule and frontal region. First segment short, irreg-
ular, attached to laterally extending, heavily sclerotized ridge. Third
segment fused with clawlike terminal process, bearing 2 setule-like
accessory processes. Male antenna (fig. 32e) 3-segmented, first seg-
ment originating on heavily sclerotized ridge, extending medially
and ventrally so that second segments of both antennae in close
proximity to each other. Second segment club shaped, with 1 large
and 1 small, spinelike projection on inner ventral surface. Third
segment short, distinct from long, saber-shaped terminal process,
shorter, spinelike subterminal process and setule-like accessory
process.

Female and male mandible (figs. 32d, e) 4-parted, rodlike, distal
part curving inward, inner surface with 12 denticulations. Female
and male postantennal process (figs. 32d, e) consisting of sharply
curved, spinelike process, with outer margin denticulated distally,
in addition to 3 nodules, 2 on base of spinelike process, third posterior
to process, each with several hairlike processes. Female postoral
process (fig. 32d) formed of heavily sclerotized plate bearing small
bifurcate projection, tines rounded distally. Male postoral process
(fig. 32¢) small, lappet-like projection of heavily sclerotized ridge.
Female maxillule (fig. 32d) nodular, with 3 setules distally; male
maxillule (fig. 32e) nodular, with 1 long and 1 short setule distally.
Male oral region well anterior and congested, mouth cone located
between projecting first segments of antennae; postoral process,
antenna base, mouth cone, maxillule, and maxilla in close proximity.

Ficure 33.—Lepeophtheirus? fallolunulus, new species, ventral view: a, female sternal furca
(sf), tinelike projections, and base of first thoracic legs (ip=interpodal plate); b, male
sternal furca (sf), tinelike projections, and first thoracic legs (ip=interpodal plate). Right
thoracic legs: first, posterior view; d, second segment of exopodite of first, posterior view;
e, second, anterior view; f, third, anterior view; g, fourth, posterior view.

90 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Small, heavily sclerotized, knoblike postoral adhesion process present
in both female and male (figs. 32d, e), arising from Y-shaped region
of heavy sclerotization, just posterior to postoral process. Female
and male maxilla (fig. 32f) 2-segmented, second segment with fine
membrane along medial posterior surface and 2 saber-shaped terminal
processes, innermost approximately twice the length of outermost.

Female and male maxilliped (figs. 32g, h) 2-segmented, situated
medial and posterior to maxilla base. First segment strongly devel-
oped, with indentation on proximal inner surface, identation with
pair of small, heavily sclerotized projections, adjacent surfaces of
projections forming groove receiving distal end of second segment
terminal process when second segment flexed. Second segment sepa-
rable from clawlike terminal process, bearing setalike and lappet-
shaped accessory processes.

Sternal furca (figs. 33a, 6) situated well posterior to maxilliped
bases. Bifurcate portion of female furca at end of necklike process,
bifureate portion of male furca also on process although process
much shorter. Pair of spinelike projections present, posterior to
furcal base, just anterior to interpodal plate of first thoracic legs.

For nature of legs and armature, see figures 33c-g and table 12.
First thoracic leg with 3 spinules on inner surface of second segment
of exopodite instead of 3 plumose setae normally found.

TABLE 12.—Armature of thoracic legs I-IV of the female and male of
Lepeophtheirus? fallolunulus, new species

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal
Plate
2 3
I | Outer
Inner
It | Outer c ¢,3P
m
Inner ¢,2P 3P
III | Outer Dj,s,a*,m,p H Pepe. c ¢,38P
m
Inner r§,2s,P,m 1 4P,p 3P
IV | Outer pil s,H 3mH,H

*Five-lobed adhesion pad.

{Single large denticulation.

tRamified spine; female with 6 rami (excluding tip), male with 7.
eee roughened areas.

||Several hairlike processes not tabulated.

Discussion.—The species is tentatively placed in the genus Lepeoph-
theirus because the general characteristics most closely approximate
those of Lepeophtheirus (e.g., fourth pedigerous segment free, not
completely covered dorsally by median projection of cephalothorax;
abdomen distinct, not fused with genital segment; first 3 thoracic legs

biramous although endopodite of first leg reduced, fourth thoracic leg

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 91

uniramous). ‘These characteristics plus the lunule-like structure are
also found in Caligus. ‘The lunule-like structure, however, is not iden-
tical with that of Caligus and other lunule-bearing genera (see below).

The species differs from the majority of the species in Lepeophtheirus
for the following reasons:

1. Part of the frontal region is modified to form a lunule-like
process.

2. The fourth thoracic legs are 3-segmented, not 4-segmented,
although Lepeophtheirus does include some species with 3-segmented
fourth thoracic legs (e.g., L. watanabei Shiino, 1954d).

3. There are 4 terminal spines on the fourth thoracic legs instead
of 3.

4. The endopodite of the first thoracic legs is larger than in most
members of the genus. Many authors state that the first and fourth
thoracic legs are uniramous in Lepeophtheirus (e.g., Wilson, 1905b,
Yamaguti, 1963). The endopodite of the first lez of members of
this genus is reduced or rudimentary but it is present (e.g., L. dissimu-
latus Wilson, 1905b; ZL. gonistii Yamaguti, 1936a) and the leg is
biramous.

5. The postoral process is reduced, especially in the male.

The lunule-like structure of L. fallolunulus is formed of the upturned
lateral anterior surface of the frontal region. Further, the structure
is partially separated from the rest of the frontal area and has a
strongly denticulated membrane instead of the frilled membrane found
on the rest of the region. In Caligus and other lunule-bearing caligoids,
the lunule is formed of part of the membrane found on the leading
edge of the frontal region. In these, however, the membrane is ei-
ther formed around a sinus in the frontal region or it itself forms a
sinus. This structure can be simple (e.g., C. enormis Wilson, 1913)
or the membrane can be well developed and the sinus remain open
(e.g., C. zeit Norman and Scott, 1906), or the membrane can be well
developed, the sinus closed, and the lunule of a distinct cup shape
(e.g., C. coryphaenae Steenstrup and Liitken, 1861). In all cases,
however, the lunule is associated with some type of sinus and is di-
rected ventrally. In ZL. fallolunulus there is no sinus, the structure is
directed dorsally and not ventrally, and it is found only on the female.

In several characteristics, the species is similar to some members
of the genus Anuretes and the genus Pseudoanuretes. -Anuretes par-
vulus Wilson, 1913, resembles this species in the general shape of
the fourth pedigerous segment and genital segment, antennule, and
maxilliped, but it differs in the less distinct abdomen, the absence of
the lunule-like structure, the reduction of the female postantennal
process, the reduction of the sternal furca, and in the nature of most
of the appendages. The 3 small, spinelike projections present on

92 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

the inner surface of the second segment of the exopodite of the first
thoracic leg in L. fallolunulus are also found in Pseudoanuretes chae-
todontis Yamaguti, 1936a, while the ramified spine on the first seg-
ment of the exopodite of the second thoracic leg is similar to the
condition of this spine in Anuretes serratus Shino, 1954a.

The name ‘fallolunulus” is derived from the Latin terms ‘‘fallo”’
(deceive) and ‘lunula”’ (small moon) and refers to the nature of the
lunule-like structure on the female.

Midias Wilson, 1911b

DraGcnosis.—Cephalothorax ovoid, consisting of cephalon and first
4 thoracic segments. Frontal region distinct, with lunules. Free
fourth pedigerous segment wider than long, tergal region heavily
sclerotized (as is most of the dorsal body surface), with flaplike
lateral extensions, segment without alae or separate plates in either
sex. Genital segment with short, heavily sclerotized, spinelike fifth
leg projections on posterior lateral surface. Abdomen of female
2-segmented, of male 1- or 2-segmented, basal segment of female
with lateral lobes. Antennule 2-segmented; antenna 3-segmented,
with clawlike terminal process, male with adhesion surfaces on first
and second segments. Mandible rodlike, distal region curved inward,
slightly flattened, with 12 denticulations along distal inner surface.
Postantennal process consisting of spinelike projection and 3 nod-
ules, each bearing several hairlike processes; postoral process spine-
like, either bifid or simple, male with adhesion surface on distal region
of process. Maxillule nodular, with setules; maxilla 2-segmented,
tipped by 2 saber-shaped processes. Maxilliped 2-segmented, with
clawlike terminal process; sternal furca well developed. Thoracic
legs I-III biramous although endopodite of first leg reduced, fourth
thoracic leg uniramous.

ReEMARKS.—Two species have been described in this genus and
are compared in the discussion section following the description of
M. lobodes.

Wilson (1911b, p. 628) placed M. lobodes in the subfamily Eury-
phorinae (now recognized as a distinct family) because, “among
general characters,” of the large size, “the possession of rudimentary
dorsal plates on the fourth” pedigerous segment, ‘‘a strongly inflated
genital segment . . . an abdomen with lateral lobes on the basal
joint and posterior lobes on the terminal joint,” the possession of a
2-seemented endopodite on the first thoracic leg, and the presence
of 3 segments in each of the rami of the third thoracic legs. Yamaguti
(1963) accepts this placement and includes the genus in the Eury-

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 93

phoridae and in his subfamily Tuxophorinae, which includes both
Midias and Tuxophorus, the latter of which has distinct alae on the
fourth pedigerous segment.

The division between the Caligidae and the Euryphoridae is tenuous,
both families sharing common characteristics. Dentigryps, a caligid
genus, for example, possesses euryphorid-like fifth legs; Pupulina, a
cealigid, has euryphorid-like extensions of the genital segment. Addi-
tionally, one of the characteristics of the euryphorids is the presence of
a biramous condition in the first 4 pairs of thoracic legs, yet Gloiopotes,
a distinct euryphorid, has a distinctly uniramous fourth leg. The
euryphorids are considered to be a transition group, between the cali-
gids and the pandarids, and as such there are characteristics of the
family that associate them with either or both the caligids and the
pandarids.

Midas, Dentigryps, and a few other genera exhibit some characteris-
tics which approach those of the Euryphoridae (see Lewis, 1964b, for a
discussion of the association of Dentigryps with the Euryphoridae).
Midas is characterized by having certain parts of the body heavily
sclerotized. This condition is particularly noticeable on the tergal
region and on some of the projections on the body and the appendages.
The ‘‘dorsal plates,’”’ used as a euryphorid characteristic by Wilson, are
not distinct plates but rather are thickenings of the tergum which are
heavily sclerotized and do not project as do the plates of the alae-
bearing groups. These thickenings are also found in other caligids
(e.g., Caligus productus Dana, 1853). Of the other characteristics used
by Wilson, the inflated genital segment is not unique (e.g., Synestius, a
caligid), the lateral lobes of the first segment of the abdomen are also
found in Dartevellia, and the posterior lobes on the abdomen are pres-
ent in Dentigryps curtus Wilson, 1913, both of the last two being
ealigids. Further, the posterior lobes are not described for Midias
carangis Rangnekar, 1956. The possession of a 2-segmented endopo-
dite on the first thoracic leg is a qualifying characteristic although the
endopodite is strongly reduced and no larger than that found in many
caligids.

Midias is here placed in the Caligidae because of the absence of dis-
tinctly euryphorid characteristics, the presence of a uniramous,
4-segmented fourth thoracic leg, the presence of well-developed lu-
nules, the presence of a spinelike projection as part of the postantennal
process, and the presence of a reduced endopodite on the first thoracic
leg. The author also accepts the fact that most of these characteris-
tics are as open to criticism as are those Wilson used to include the
genus in the Euryphoridae.

Q4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121
Midias lobodes Wilson

Figures 34-36

Midias lobodes Wilson, 1911b, p. 626, pl. 65; 1913, p. 225.—Causey, 19538b, p.
11.—Shiino, 1958, p. 98, figs. 1-3; 1963a, p. 343.—Yamaguti, 1963, p. 107,

pl. 127, fig. 1.
DistTRIBUTION AND Hosts.—4 host records:
locality hosts references
Gulf of Mexico Sphyraena barracuda Wilson, 1911b
Western Atlantic S. barracuda Wilson, 1913
Hawaii S. barracuda Shiino, 1963a
Indian Ocean Sphyraena species Shiino, 1958

Marrerrau.—One female and 1 male (USNM 112888) from the ex-
ternal surface of Sphyraena barracuda (Walbaum) captured 80 miles
south of Oahu, Hawaii (USFWS). Two females and 2 males (retained
by author) from the external surface of S. barracuda (Walbaum)
captured in the Hawaiian region (USFWS).

MBEASUREMENTS.— (In mm) 1 female and 3 males:

female male

Total length, excluding caudal setae 12.00 7.13, 6.90, 6.30
Length of cephalothorax, including frontal region 4.73 3.68, 3.68, 3.38
Width of cephalothorax 4.35 3.38, 3.38, 3.08
Length of genital segment, excluding fifth legs 3.29 1.55, 1.44, 1.33
Width of genital segment 3.30 1.81, 1.92), 1-85
Length of abdomen 3:29" A708 0.78, 1267
Width of first segment of abdomen 3.11

Length of fifth leg 0.27 0.48, 0.54, 0.56
Length of caudal rami 0.35 0.29, 0.31, 0.28

Descriertion.—Female cephalothorax (fig. 34a) subovoid, consist-
ing of cephalon and first 4 thoracic segments. Frontal region approx-
imately one-fifteenth the length of cephalothorax, with pair of ovoid
lunules and bearing fine membrane along anterior edge. Lateral
cephalothoracic margin smoothly irregular, with fine membrane
extending laterally and second membrane extending medially, from
ventral edge. Posterior lateral surface of cephalothorax with small,
pocket-like depression; posterior sinus (fig. 34c) narrow, outer and
inner margins converging at opening, outer surface with fine membrane
extending into sinus and second membrane extending laterally over
division between lateral and median cephalothoracic regions. Median
cephalothoracic region not extending past posterior end of lateral
cephalothoracic regions, posterior margin irregular. Major dorsal
cephalothoracic grooves forming irregular H; ocular region distinct in
anterior third of body. Dorsal surface with scattered nodules, each
bearing 1 or 2 hairlike processes.

|
|
|
|
|
|
|

No. 3574 COPEPOD CRUSTACEANS—LEWIS 95

MIND
. ae
~ a

PINNIN
<<

Ficure 34.—Midias lobodes Wilson, 1911, dorsal view: a, female; b, male; c, posterior cepha-
lothoracic sinus. Ventral view: d, female genital segment; ¢, male genital segment.
Dorsal view: f, female right fifth leg; zg, male, same. Ventral view: h, male right sixth
leg; 1, female caudal ramus and abdominal projection; 7, male, same.

96 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

Female free fourth pedigerous segment distinct from cephalothorax,
also from genital segment on ventral but not on dorsal surface. Seg-
ment short, expanded medially into small, flaplike projections over
place of attachment of fourth thoracic legs. Genital segment (fig. 34d)
broader posteriorly, tapered regularly from slightly narrower anterior
end, lateral posterior surface slightly lobate. Fifth legs (fig. 34f)
situated on posterolateral surface, consisting of minute, knoblike
swelling, bearing single plumose setule, additional spikelike projection
present immediately posterior to swelling, bearing 3 plumose setules,
2 on dorsal surface, third on distal inner surface.

Female abdomen indistinctly 2-segmented, incompletely separable
from genital segment. First segment butterfly shaped; second seg-
ment subrectangular except for lobate outer distal surfaces, lobes
denticulated. Caudal rami slightly narrower proximally than dis-
tally, distal half of inner margin plumose, distal surface with 3 plumose
setae medially and 1 laterally in addition to 2 plumose setules, 1 adja-
cent to both outer and inner setae.

Male cephalothorax (fig. 346) similar to that of female. Free fourth
pedigerous segment with smaller lateral projections situated more
posteriorly, but still over position of leg attachment. Genital seg-
ment (fig. 34e) distinct from both free fourth pedigerous seg-
ment and abdomen, lateral surfaces broadly convex, fifth legs (fig. 34f)
spikelike, projecting from posterior lateral surfaces, spike bearing
plumose setule from nodule on proximal dorsal surface, plumose
setule from distal dorsal surface, and plumose setule from distal
surface. Sixth legs (fig. 34h) present medial to fifth, consisting of
node bearing 2 plumose setules distally. (The condition of the fifth
and sixth legs of the male approaches that of the male of Dentigryps.)
Abdomen appearing as 1 segment although indistinct evidence of
second segment visible in posterior half. Abdomen broader anteriorly
than posteriorly, lateral margins flatly concave, dorsal surface strongly
concave. Outer distal surfaces lobate, lobes denticulated. Caudal
rami as in female.

Female and male antennule (fig. 355) 2-segmented, attached to
ventral-lateral anterior surface of cephalothorax, just posterior to
division between frontal region and remaining cephalothorax. First
segment approximately 1% times the length of second, distal half of
anterior and anterior ventral surface bearing approximately 26 lightly
plumose setae and setules. Second segment elongate, slightly longer
in male than in female, with single naked setule from posterior medial
surface, 11 from distal surface. Female antenna (fig. 35c) 3-segmented,
situated slightly medial and posterior to antennule base. First seg-
ment short, knoblike, with small, heavily sclerotized subconical pro-
jection from posterior surface. Second segment short, subrectangular ;

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 97

h

Ficure 35.—Midias lobodes Wilson, 1911, ventral view: a, right side of frontal region show-
ing lunule; 5, left antennule; c, female oral region, right side showing antenna, postantennal
Process, mouth cone, mandible, maxillule, postoral process, and maxilla base; d, male
right antenna; ¢, male right postantennal process; f, male right postoral process and max-
illule; g, left maxilla; h, right maxilliped; 1, sternal furca.

226-321—67—_—_7

98 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

third segment fused with clawlike terminal process, bearing 2 setule-
like accessory processes, 1 at indication of division between segment
and terminal process, second from proximal inner surface. Male
antenna (fig. 35d) basically similar to that of female although segments
and processes relatively longer and first 2 segments bearing adhesion
surfaces.

Female and male mandible (fig. 35¢) 4-parted, distalmost part
curved inward, with 12 denticulations on inner surface. Female post-
antennal process (fig. 35c) consisting of short, spikelike projection,
just lateral to antennal base, and 3 nodules, each with several hairlike
processes; 2 nodules situated on base of spikelike projection, third
just posterior to base. Male postantennal process (fig. 35¢) similar to
that of female although projection slightly thicker distally. Female
postoral process (fig. 35c) spike shaped, with small spine projecting
from distal half of inner surface. Male postoral process (fig. 35f) simi-
lar to that of female except with lappet-shaped accessory projection
and adhesion surface. Female and male maxillule (fig. 35c) nodular,
bearing 3 setules distally. Female and male maxilla (fig. 35g) 2-seg-
mented, situated lateral and slightly posterior to postoral process.
First segment approximately three-fourths the length of second, both
elongate. Second segment with large, oval, membranous projection
from distal half of inner surface, with 2 saber-shaped terminal proc-
esses. Innermost terminal process approximately 1 times the length
of outermost, both with very lightly frilled membranes along both
inner and outer margins.

Female maxilliped (fig. 35h) 2-segmented, situated posterior and
slightly medial to maxilla base. First segment strongly developed,
with elongate proximal projection serving as articulation and muscle
attachment surface. Second segment short, incompletely fused with
clawlike terminal process, bearing small, setule-like accessory process
from inner surface, at junction of segment and terminal process.
Male maxilliped similar to that of female except first segment with
transverse ridge on distal half of inner surface. Female and male
sternal furca (fig. 352) situated between and posterior to maxilliped
bases, associated with heavily sclerotized transverse band on cepha-
lothorax. Tines curving medially slightly toward distal ends, sinus
U-shaped, lateral surface of tines thinner than medial and inner surfaces.

For nature of legs and armature, see figures 36a—h and table 13.

Ficure 36.—Midias lobodes Wilson, 1911, right thoracic legs, anterior view: 4, first; b, distal
end of second segment of exopodite of first; c, second; d, third; e, exopodite of third (pos-
terior view); f, first segment of exopodite of female third; g, male fourth; h, protopodite of
female fourth.

99

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

do
FOr
E
E
Be
Oo

|

0.5 mm.@o"

0.5mm.2a"

100 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

TABLE 13.—Armature of thoracic legs I-IV of the female and male of
Midias lobodes Wilson, 1911b

Inter- Protopodite Exopodite Endopodite
bega|/ Surfaces ||ood al ieee see. SE eS es ke Oe ee
Plate
1 2 1 2 3 1 2 3
I | Outer ss,p fm,rh 3fm,3dH, ss
Inner Dp Cc 3P
II | Outer m,p | m,fm, dmH | 24mH,Q, ¢,C ic c,3P
m dmH ie
Inner Sue TS alco (op) 2 c,4P P ¢,2e c,3P
III | Outer a,d,m,p s,fmH 3s,¢,p’ ¢,3p’,P c ¢.2P
m
Inner d,3s, P,s, Ce c,3P iz c,4P
m,s
IV | Outer S,Ss,s*(pf) 2s,fm,fmH | fm,fmH 3fm,3fmH

*Female only.
{Male only.

Discusston.—Several parts of Wilson’s original description (1911b)
need to be mentioned and characteristics that he did not describe
should be discussed. ‘These are characteristics noted not only on
the Hawaiian material but also on the original type material. Wilson
does not figure or describe an accessory process on the third segment
of the antenna but a setule-like process is present in the female
and 2 are present in the male (figs. 36c, d). The postantennal process
is composed of 3 nodules, each bearing several hairlike processes, as
well as a spinule-like projection. There is a great deal of variation
in the spinelike projection forming the postoral process; basically,
it is bifid in the female and simple in the male, but some of the female
specimens in the collections of the U.S. National Museum have a simple
process, while in others the process is bifid on one side and simple on
the other (see discussion of Gloiopotes huttoni for similar case). Wilson
indicates that the endopodite of the third thoracic leg is 3-segmented
although in all of the specimens examined, including the type mate-
rial, the endopodite is 2-sezmented. There is a cuticular break in
the second segment in all of the specimens (fig. 36d) but this break
is Incomplete and there is no indication of a 3-segmented condition,
either in the cuticle or by the associated musculature.

Wilson deposited a series of cotypes (USNM 39613) in the type
collection. One of these (a female, USNM 39613) has been selected
as the lectotype, a second (male, USNM 112846) as the allolectotype,
while the rest (USNM 112847) have been designated paralectotypes.

Rangnekar (1956) described the only other species in the genus,
M. carangis. The feature which best distinguishes the two species
is the presence of posterior lobes on the second segment of the abdo-
men of M. lobodes and their stated absence in M. carangis. A second

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 101

characteristic is the described ‘“‘sickle-shaped claw” on the first seg-
ment of the exopodite of the third thoracic leg in M. carangis; this
claw is bifid in M. lobodes.

Caligus Miiller, 1785

Draqnosis.—Cephalothorax consisting of cephalon and first 4
thoracic segments; frontal region distinct, with lunules. Free fourth
pedigerous segment without alae, sometimes with small, lappet-like
lateral projections of tergum extending over proximal surface of fourth
thoracic leg. Genital segment of variable shape, infrequently with
extensions of posterior lateral surface, without elongate, heavily
sclerotized fifth leg projections. Abdomen 1-4 segmented, of variable
length but not longer than combined lengths of cephalothorax, free
fourth pedigerous segment and genital segment. Antennule 2-seg-
mented; antenna 3-segmented, with clawlike terminal process, vari-
ously modified in male. Mandible rodlike, distal end curved inward,
slightly flattened, distal inner surface with 12 denticulations. Post-
antennal process consisting of 3 nodules, each with several hairlike
processes, usually with spinelike projection. Postoral process spine-
like, simple or with small accessory spine. Maxillule nodular, with
hairlike or setular projections; maxilla 2-segmented, terminated by
2 saber-shaped projections. Maxilliped 2-segmented, with claw-
like terminal process and setule-like accessory process. Sternal furca
present. Thoracic legs I-III biramous although endopodite of first
reduced; fourth thoracic legs uniramous.

Caligus coryphaenae Steenstrup and Liitken
Fiaures 37-39

?Caligus scutatus Milne-Edwards, 1840, p. 453.

Caligus coryphaenae Steenstrup and Liitken, 1861, p. 360, pl. 4, fig. 7—Richiardi,
1880, p. 148.—Valle, 1880, p. 58.—Carus, 1885, p. 358.—Bassett-Smith, 1899,
p. 451.—Brian, 1899a, p. 4.—Wilson, 1905b, p. 556, 559 (key).—Brian,
1908, p. 2; 1912, p. 7.—Wilson, 1923, p. 5; 1937, p. 424.—Heegaard, 1949,
p. 241, figs. 6-10.—Pearse, 1952a, p. 15.—Capart 1959, p. 81, fig. 14a—d.—
Shiino, 1959b, p. 2, figs. 1-2; 1959d, p. 294, fig. 12.—Kurian, 1961, p. 68,
figs. 16-24.—Nunes-Ruivo, 1962a, p. 70.—Pillai, 1962b, p. 513, figs. 1, 2.—
Shiino, 1963a, p. 336, fig. 1—Yamaguti, 1963, p. 51, pl. 59, fig. 3.

Caligus bengoensis Scott, 1894, p. 130, pl. 14, fig. 19.

Caligus aliuncus Wilson, 1905b, p. 576, pl. 9; 1935b, p. 330.—Bonnet, 1948,
p. 7.—Causey, 1953b, p. 8.

Caligus elongatus Heegaard, 1943b, p. 11, figs. 21-31.

Caligus tesserifer Shiino, 1952, p. 89, fig. 5.

Not Caligus coryphaenae.—Brian, 1935, p. 51, figs. 19, 20.—Yamaguti, 1936a,
p. 5, pl. 4, figs. 40-54.— Barnard, 1955, p. 246, figs. 8a—d.

102 PROCEEDINGS

DISTRIBUTION AND nosts.— 19 + host records:

locality
Subtropical Atlantic

Gulf of Mexico

Eastern Pacific

Hawaii
Western Pacific

Indian Ocean

Mediterranean

hosts
Coryphaena
“Bonito”’
C. hippurus
Plankton
Parathunus obesus
Euthynnus alleteratus
Coryphaena hippurus
Sarda species
Coryphaena hippurus
Euthynnus lineatus
Katsuwonus pelamis
Euthynnus alleteratus
Coryphaena hippurus
Katsuwonus pelamis
Coryphaena hippurus
Unknown
Squalus acanthias
Neothunnus albacora
Coryphaena hippurus

OF THE NATIONAL MUSEUM VOL. 121

reference
Steenstrup and
Liitken, 1861
Brian, 1908
Scott, 1894
Nunes-Ruivo, 1962a
Wilson, 1935b
Pearse, 1952a
Wilson, 1937
Heegaard, 1949

Shiino, 1959
Bonnet, 1948
Shiino, 1959d

Shiino, 1959b
?Milne-Edwards, 1840
Wilson, 1923

Shiino, 1959b
Richiardi, 1880

Materitau.—One female and 1 male (USNM 112889) from the ex-
ternal surface of Katsuwonus pelamis (Linnaeus) collected near Fanning
Island (USFWS, LL cruise 9). One female (USNM 112890) from the
external surface of K. pelamis (Linnaeus) collected at 20°35’N, 157°-
45’W (USFWS, CHG cruise 30, station 30). One female and one
male (USNM 112891) from the external surface of K. pelamis (Lin-
naeus) collected in the Hawaiian Island region (USFWS, HMS cruise 34).
One female and 1 male (USNM 112892) from the external surface of K.
pelamis (Linnaeus) collected at 20°4.5’N, 160°37.5’W (USFWS, HMS
cruise 39, station 17). One immature male (USNM 112893) from the
plankton, 0°00’N, 157°42’W (USFWS, HMS cruise 2, station 52).
One female and one male (USNM 112894) from the external surface of
K. pelamis (Linnaeus) collected 120 miles north of Oahu, Hawai
(USFWS). One female and 2 males (USNM 112895) from the exter-
nal surface of Huthynnus yaito (Jordan and Evermann) collected at
French Frigate Shoals (USFWS, HMS cruise 39, station 32). One
male (USNM 112896) from the external surface of Coryphaena hippu-
rus Linnaeus from unknown locality (in collections of U.S. Fish and
Wildlife Service, Honolulu). Two females (USNM 112897) from the
external surface of Huthynnus yaito (J. and E.) taken near Moku Manu
Island, near Oahu, Hawaii (USFWS). One female and 1 male
(USNM 112898) from the external surface of Katsuwwonus pelamis
(Linnaeus) collected at 2°S, 132°W (USFWS, JRM cruise 34, station
5). One male (USNM 112899) from the external surface of Huthyn-
nus yaito (J. and E.) collected in unknown locality (USFWS, CHG
cruise 52). Five females and 2 males (retained by author) from the

No. 3574 COPEPOD CRUSTACEANS—LEWIS 103

external surface of K. pelamis (Linnaeus) captured by Robert
Stevenson near Oahu, Hawaii.

MEASUREMENTS.—(In mm) 13 females and 10 males:

female male
mean range mean range

Total length, excluding caudal setae 6.61 5.85-7.28 5.52 4 73-6. 00
Length of cephalothorax, including frontal

region 3.28 3.03-3.63 3.52 3. 11-3. 74
Width of cephalothorax 2.71 2.44-3.00 2.78 2. 52-3. 03
Length of genital segment 1.77 1.30-2.11 0.99 0. 81-1. 11
Width of genital segment 198 “1. 33=18h) 1.25 lit 33
Length of abdomen 1.53 1.06-1.80 0.82 0. 68-0. 94
Length of caudal rami 0.25 0. 23-0.30 0.27 0. 23-0. 30
Length of egg strings (4 strings) 6.15 2. 40-8. 78

Description.— Female cephalothorax (fig. 37a) consisting of ceph-
alon and first 4 thoracic segments, with several hairlike processes
scattered over dorsal surface. Frontal region approximately one-
fifteenth the length of cephalothorax, with narrow membrane along
anterior margin; lunules large, projecting posteriorly, on ventral
surface, past division between frontal region and remaining cepha-
lothorax. Lateral cephalothoracic margins slightly irregular, with
fine membrane, also with distinct socket-shaped depression poste-
riorly. Posterior sinus (fig. 37c) distinct, U-shaped, with finely serrat-
ed membrane along outer margin. Posterior median cephalothoracic
region projecting slightly past lateral cephalothoracic regions, poste-
rior margin flatly convex. Major dorsal cephalothoracic grooves
forming irregular H, with posterior and anterior longitudinal grooves
flaring outward, anterior extending to just posterior to ocular region.
Ocular region distinct, pigment cups, containing lens element, con-
tiguous on median longitudinal axis of body.

Female fourth pedigerous segment free, width approximately
three times the length, dorsal surface raised slightly, appearing
platelike. Genital segment (fig. 37d) distinct from fourth pedigerous
segment, anterior end narrower than posterior, lateral margins wavy.
Posterior lateral regions of genital segment lobate, projecting pos-
teriorly shghtly, bearing fifth legs on outer surface. Fifth leg (fig. 37/)
consisting of 2 adjacent nodes, first bearing single plumose seta,
second bearing 3 plumose setules. Median posterior surface of
genital segment indistinctly and incompletely separable from abdo-
men. Abdomen considered 4-segmented by most workers although
only 2 distinct divisions (3 segments) present, third division suggested
by constriction. Anterior part of abdomen with constriction and
may represent 2 segments although constriction superficial and
developing eggs extend from genital segment through full length of
first part of abdomen. Anterior part also with small, bean-shaped

104 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

= phy ll

~

Ficure 37.—Caligus coryphaenae Steenstrup and Liitken, 1861, dorsal view: a, female; b,
male; ¢, posterior cephalothoracic sinus. Ventral view: d, female free fourth pedigerous
segment, genital segment, and anterior end of abdomen; ¢, male, same; f, female fifth leg;
g, male fifth (V) and sixth (VI) legs. Dorsal view: A, left caudal ramus; 7, frontal region,
right side showing lunule and antennule base.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 105

process on anterior dorsal surface. Posteriormost segment with
deep indentation in posterior half of lateral surface, indentations
forming articulation and attachment surfaces for caudal rami.
Caudal rami (fig. 37h) short, not extending past posterior end of
last abdominal segment, bearing 3 large, plumose setae from medial
posterior surface, 2 additional plumose setae from outer posterior
surface; sixth plumose seta present on inner posterior surface.

Male cephalothorax and free fourth pedigerous segment (fig. 37b)
similar to female. Genital segment (fig. 37g) suborbicular, lateral
posterior surface indented, indentations bearing fifth legs. Fifth
leg (fig. 37g) similar to that of female; sixth leg (fig. 37g) arising medial
and slightly posterior to fifth, from broad, lappet-like pad, leg con-
sisting of single plumose seta and 2 plumose setules. Abdomen
distinct from genital segment, 2-segmented, first segment broader
anteriorly than posteriorly, second segment similar to posteriormost
segment of female. Caudal rami as in female.

Female and male antennule (fig. 38a) 2-segmented, attached to
lateral-anterior ventral surface of cephalothorax, immediately
posterior to division between frontal region and remaining cepha-
lothorax. First segment more than twice the length of second,
with approximately 17 plumose setae and setules in addition to 2
knobs along distal half of anterior ventral surface; second segment
club shaped, with 1 plumose and 9 naked setae and setules in distal
region. Female antenna (fig. 38b) 3-segmented, situated posterior
and slightly medial to antennule base. First segment short, irre-
gularly cup shaped, heavily sclerotized; second segment longer than
first, length approximately 1% times the width. Third segment
fused with clawlike terminal process, bearing knoblike accessory
process at indistinct break in sclerotization, in proximal third; second
knoblike accessory processs at proximal end of segment. Male
antenna (fig. 38c) 3-segmented, first 2 segments basically similar
to those of female, third segment terminal process distinct from
segment, with second claw at proximal end of process.

Female and male mandible (fig. 38b) 4-parted, fourth part flattened,
with 12 denticulations along inner margin. Female and male post-
antennal process (fig. 385) consisting of 3 nodules, each with several
fine, hairlike processes. Female and male postoral process (fig. 380)
consisting of subtriangular, spimelike projection, bluntly pointed or
rounded distally. Female and male maxillule (fig. 386) nodular, bear-
ing 3 setae, situated in depression of platelike area of heavy sclerotiza-
tion between posterior margin of first segment of antenna and
subtriangular postoral process. Ovoid area of heavy sclerotization
present, extending posteriorly and medially between bases of maxillae,
with ridgelike projection along axis. Female and male maxilla (fig. 38

106 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

~“GammXo') € yf

+4
0.2mm. 0.2mm(?ic*

Ficure 38.—Caligus coryphaenae Steenstrup and Litken, 1861, ventral view: 4, left anten-
nule; b, female oral region, left side showing antenna, postantennal process (pan), mouth
cone, mandible (mdbl), maxillule (ma-1), postoral process, postoral adhesion pad (pa),
and maxilla base (ma-2);c, male right antenna; d, left maxilla; e, right maxilliped; f, ster-
nal furca, maxilliped bases (mxpd), and interpodal plate of first thoracic leg (ip).

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 107

d) 2-segmented, situated just lateral and posterior to postoral process.
First segment slightly shorter than second, excluding terminal proc-
esses, although wider; second segment elongate, tapered proximally
and distally, bearing membrane from middle of inner margin and 2
saber-shaped terminal processes, each with membrane along margins.

Female maxilliped (fig. 38¢) 2-segmented, situated posterior and
medial to maxilla base. First segment approximately 2% times the
length of second, strongly developed, width slightly less than half the
length. Second segment distinct from clawlike terminal process,
bearing single, setule-like accessory process from posterior-inner distal
surface. Male maxilliped similar to that of female although first seg-
ment with small, ledge-shaped projection on proximal inner surface.
Female and male sternal furea (fig. 38f) situated between posterior
ends of maxilliped bases. Furcal tines diverging widely, heavily
sclerotized. Pair of lappet-shaped projections present, lateral to base
of furca.

For nature of legs and armament, see figure 39 and table 14.

TABLE 14.—Armature of thoracic legs I-IV of the female and male of
Caligus coryphaenae Steenstrup and Liitken, 1861

Inter- Protopodite Exopodite Endopodite
Leg | Surface || podal a
Plate
2 1 2 3 1 2 3
I} Outer Ss,S,p rh 3dmH,p’
Inner p c 3P
II | Outer m,p |m,m,mH | mH 2H,Q CHiee C,3P
m
Inner pe m,p | ¢,P Ge @5P Pa e2e ¢3P
III | Outer d,m,p mi Dp *:c;p! ¢,3p’,p Coalncsaik
m
Inner d,P,s,m,s CE 3P Py 4p
peel ae = 4 2
IV | Outer p fm,dHt fm,dHt fm,dH,fm,
dH,fm,dHf
|

*Element in middle of segment.
{Female has denticulated membrane, not separate denticulations as in male.
Discussion.—Steenstrup and Liitken (1861, p. 362) state that
“Our C. coryphaenae also seems closely related to C. scutatus M. Edw.
(hist. d. Crust. t. III P. 453 no. 7), but as this is from the Indian
Ocean it does not seem probable to us that this is the same species.”
The present distribution and hosts of Caligus coryphaenae suggest
that Steenstrup and Liitken erred in using the different location as a

108 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ny wn

ea AL gt
X WV \ NN |! ih
eae : Tr

Ficure 39.—Caligus coryphaenae Steenstrup and Liitken, 1861 , right thoracic legs, anterior
view: 4, first; b, distal region of second segment of exopodite of first; c, second; d, third;
é, exopodite of third (posterior view); f, fourth.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 109

primary reason for separating the two species. Milne-Edwards (1840),
however, does not provide a figure of C. scutatus and does not list
the host from which the female specimen(s?) (male unknown) was
collected. Further, C. scutatus is not given in a list of Milne-Edwards’
species contained in the Paris Museum where most of his specimens
were deposited (list sent to Dr. R. Parker, Fisheries Research Board of
Canada, Nanaimo, British Columbia). Because of the lack of a
definitive description and specimens of C. scutatus, it is felt best to use
the name only as a questionable entry in the synonymy of C. cory-
phaenae.

The deletion of Brian, 1935, and Barnard, 1955, from the synonymy
is based on Pillai’s 1962b discussion of the synonymy of C. coryphaenae.
Shiino, 1959b, also lists Caranz pelagica, Seriola dorsalis, Rachycentron
canadus, and Caranz hippos as reported hosts.

Caligus quadratus Shiino
Fiaurss 40-42

Caligus productus.—Rathbun, 1884, p. 487.—Wilson, 1905b, p. 597, pl. 14, figs.
162-170.

Caligus coryphaenae.—Yamaguti, 1936a, p. 5, pl. 4, figs. 162-170.

Caligus quadratus Shiino, 1954c, p. 26, fig. 1; 1959b, p. 8, figs. 3-5; 1960a, p.
472.—Yamaguti, 1963, p. 59, pl. 75, fig. 4.

DISTRIBUTION AND HOSTS.—8 host records:
locality host reference
Japan Neothynnus macropterus
Histiophorus orientalis
Coryphaena hippurus
Neothunnus albacora

Rhinobatus schlegelit Shiino, 1954ce, 1959b, 1960a
“‘Pacific”’ Coryphaena hippurus Yamaguti, 1936a
Atlantic Coryphaena hippurus Wilson, 1905b
Western North At- Coryphaena species Rathbun, 1884

lantiec

MatTeErIAL.—Five females and 2 males (USNM 112900) from the
gill cavity of Coryphaena hippurus Linnaeus captured at 15°N,
155°30’W (USFWS, HMS cruise 36, stations 26-27). Two females
(1 immature) (USNM 112901) from the gill cavity of C. hippurus
Linnaeus captured 120 miles south of Oahu, Hawaii (USFWS).
Two females (1 immature) (USNM 112902) from the gill cavity of
C. hippurus Linnaeus captured 130 miles south of Nihau, Hawaii
(USFWS, HMS cruise 39). Three females and 1 male (retained by
author) from the buccal cavity, external surface and gill cavity of C.
hippurus Linnaeus captured near Oahu, Hawaii.

110 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

MEASUREMENTS.—(In mm) 11 females and 3 males:

female male
mean range

Total length, excluding caudal setae 5.02 3.90-6.15 3. 50, 3. 25, 2. 85
Length of cephalothorax, including frontal

region 1.79 1. 28-2.08 1. 93, 1. 95, 1. 53
Width of cephalothorax 1865) AeS0-I 88) iGo eos. sa
Length of genital segment 1.55 1. 33-1. 80 0. 73, 0. 60, 0. 55
Width of genital segment 1.24 1.08-1.438 0. 58, 0. 55, 0. 43
Length of abdomen 1.64 1.20-2.18 0. 73, 0. 60, 0. 53
Length of caudal rami 0:15 0. 10-0.18 0. 18,0. 19,0. 13
Length of egg strings (4 strings) 2.33 1. 25-3. 05

DescrieTion.—Female cephalothorax (fig. 40a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region slightly
more than one-tenth the length of cephalothorax, with membrane
along most of anterior margin. Lunules extending posteriorly, on
ventral surface, past junction of frontal region and rest of cepha-
lothorax. Lateral margin of cephalothorax slightly irregular, with
numerous small indentations, bearing narrow membrane extending
laterally and second, frilled membrane extending medially, on ventral
surface. Posterior lateral cephalothoracic surface with small, pocket-
like depression on dorsal surface. Posterior sinus (fig. 40c) irregu-
larly V-shaped, with fine membrane along outer surface. Posterior
margin of median cephalothoracic area irregularly convex, area
extending posteriorly slightly past lateral areas, with 3 pairs of minute,
hairlike processes on median-posterior dorsal surface. Major dorsal
cephalothoracic grooves forming irregular H; dorsal surface with
scattered minute, hairlike processes, in addition to those on posterior
median surface. Ocular region distinct, in anterior third of cepha-
lothorax, pigmented cups contiguous on median longitudinal axis
of body.

Female free fourth pedigerous segment broadest medially, angled
toward anterior and posterior ends. Segment distinct from cepha-
lothorax, distinctly separable from genital segment ventrally, in-
distinctly separable dorsally. Genital segment (fig. 40d) variable in
shape, subovoid to rectangular, anterior lateral and lateral posterior
surfaces frequently slightly lobate. Fifth leg (fig. 40f) situated on
lateral ventral surface in posterior fourth of segment, consisting of
2 nodules, outermost with single lightly plumose setule, innermost
with 2 lightly plumose setules.

Abdomen elongate, consisting of 1 or more segments, distinct from
genital segment. Anterior end of abdomen narrower ventrally
than dorsally. Lateral surfaces irregular, with several distinct
constrictions in some specimens, constrictions absent in others. Ab-
domen narrower posteriorly than anteriorly, posterior region (ap-

a ee ee a

NO. 3574 COPEPOD CRUSTACEANS—LEWIS Ast

Ficure 40.—Caligus quadratus Shiino, 1954, dorsal view: a, female; b, male; c, posterior
cephalothoracic sinus. Ventral view: d, female genital segment; ¢, male, same; f, female
fifth leg; g, male, same; , male sixth leg; 7, caudal ramus.

1g PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

proximately one-seventh the total length) with almost parallel sides,
region with indistinct evidence of division from rest of abdomen.
Posterior end of abdomen with concave lateral surfaces, irregularly
U-shaped median region, at anal opening. Caudal rami (fig. 407)
subrectangular, inner margin plumose distally, distal surface with
3 plumose setae medially, fourth plumose seta laterally; 2 plumose
setules also present, one on either side of setae.

Cephalothorax of male (fig. 405) similar to that of female. Free
fourth pedigerous segment of similar shape although division between
segment and genital segment distinct, segment bearing ridgelike
area of heavy sclerotization posteriorly, extending from posterior
end of fourth leg attachment surface. Genital segment (fig. 40e)
barrel shaped, constricted slightly anteriorly. Fifth legs (fig. 409)
situated in posterior half of ventral lateral surface, consisting of 3
lightly plumose setules. Sixth legs (fig. 40h) situated on posterior-
lateral ventral surface, consisting of pair of lightly plumose setules
arising from nodule. Abdomen 2-segmented, distinct from genital
segment. Second segment slightly less than 1% times the length of
first, first subrectangular; second segment narrower anteriorly than
posteriorly, lateral margins flatly convex, posterior surface as in
female. Caudal rami as in female.

Female antennules (fig. 41b) 2-segmented, attached to lobate pro-
jection of lateral anterior-ventral surface of cephalothorax. First
segment slightly less than 1% times the length of second, broad proxi-
mally, tapered to narrow distal surface; distal half of anterior and
anterior ventral surface bearing approximately 26 lightly plumose
setules and setae. Second segment rod shaped, distal region with
approximately 12 naked setules. Male antennule similar to that of
female except second segment (fig. 41c) longer. Female antenna
(fig. 41d) 3-segmented, situated posterior and slightly medial to anten-
nule base. First segment stubby, irregular, base subtriangular,
bearing small, heavily sclerotized, subconical projection from outer
posterior surface. Second segment well developed, broader proxi-
mally than distally. Third segment indistinctly separable from
clawlike terminal process, bearing setule-like accessory process from
distal lateral surface and small, lappet-like process from proximal
posterior surface. Male antenna (fig. 41¢) 3-segmented, first segment
short, subovate, concave distally. Second segment club shaped,
narrow end distalmost, with small adhesion surface on proximal
anterior surface and second, ridge-shaped adhesion surface on distal
half of inner surface. Third segment short, indistinctly separable
from trifid terminal process, bearing setule-like accessory process from
anterior distal surface. ines of trifid terminal process short, sit-
uated at distal end of process.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 113

Ficure 41.—Caligus quadratus Shiino, 1954, ventral view: , frontal region, right side show-
ing lunule and antennule base; b, female right antennule; c, second segment of male right
antennule; d, female oral region, right side showing antenna, postantennal process, mouth
cone, mandible, maxillule, postoral process, and maxilla base; e, male right antenna, post-
antennal process and base of antennule;f, claw and accessory process of male left antenna;
g, left maxilla; h, female left maxilliped; 7, male, same; 7, sternal furca.

226-321—67——_8

114 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Female and male mandible (fig. 41d) 4-parted, distalmost part
curved medially, with 12 denticulations along inner surface. Female
postantennal process (fig. 41d) situated lateral to antennal base, con-
sisting of short, spinelike projection bearing pair of nodules on prox-
imal surface, third nodule present slightly posterior to base of projection,
all 3 nodules with several hairlike processes. Male postantennal
process (fig. 41e) also consisting of spinelike projection and 3
nodules although projection much longer and strongly curved. Fe-
male and male postoral process (fig. 41d) spinelike, elongate. Female
and male maxillule (fig. 41d) nodular, with 3 setules distally. Female
and male maxilla (fig. 41g) 2-segmented, situated lateral and slightly
posterior to postoral process. First segment more than four-fifths
the length of second; second segment elogate, with small, horseshoe-
shaped membrane from distal medial surface and 2 saber-shaped
terminal processes. Innermost terminal process slightly less than 1%
times the length of outermost, bearing fine, filmy membrane along
inner margin; outermost with very finely frilled membrane along
both outer and inner margins.

Female maxilliped (fig. 41h) 2-segmented, situated posterior and
slightly medial to maxilla base. First segment well developed, with
long, strongly curved proximal articulation and muscle attachment
projection. Second segment distinct from clawlike terminal process,
bearing setule-like accessory process on distal inner surface. Male
maxilliped (fig. 417) more strongly developed, proximal projection
of first segment shorter, inner medial surface of segment with small
nodes proximally, larger node distally (node spinelike on 1 maxilliped
of 1 of the 3 specimens); second segment and processes as in female.
Female and male sternal furca (fig. 417) situated between and slightly
posterior to maxilliped bases. Furca with slightly curved or straight
tines, arising from butterfly-shaped area of heavy sclerotization.
Second butterfly-shaped area present between interpodal plates of
first and second thoracic legs.

For nature of legs and armature, see figure 42 and table 15.

Discussion.—The Hawaiian specimens here described as C.
quadratus Shino differ from Shiino’s 1954c¢ and 1959b description
as follows:

1. The terminal process of the male antenna differs from that
shown by Shiino although he does not include a lateral view of this
structure in his figures. The terminal process of the Hawaiian
specimens more closely resembles that of C. kuroshio Shiino, 1959c,
although the characteristic denticulation of the outer margin of
the second segment of the endopodite of the second thoracic leg of
C. quadratus and the Hawaiian specimens is not indicated. The

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 115

>

ated

WG Heda UG
a

a

AYE
cS
Ny

4

Wi
ae)

NAAN
wy

'

\
AN

({

4

*

Ficure 42.—Caligus quadratus Shiino, 1954, right thoracic legs, anterior view: 4, first;
b, distal end of second segment of exopodite of first; c, second; d, third; e, exopodite of
third; f, fourth.

116 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

TABLE 15.—Armature of thoracic legs I-IV of the female and male of
Caligus quadratus Shiino, 1954c

Inter- Protopodite | Exopodite Endopodite
Leg | Surface podal a
Plate
1 | 2 1 | 2 3 1 2 3
I | Outer ss,D rh H,2dH,P
Inner d,p (Oe 3P
II | Outer m,p | m,fm,fmH fmH h,mH,Q Dye D,3P
m
Inner s,P m;s' | ¢,P. er c,5P Pai Pane or
III | Outer d,m,p H ¢,p’ ¢,3p’,P Ganjnc.2e
m
Inner 2s,P,s,m,s ce ¢,3P Pane, 42
IV | Outer $,p s,fm,fmH 4fm,4mH

Hawaiian specimens also differ from C. kuroshio in the nature of
other appendages and the body.

2. Shiino does not figure or describe a break in the plumosities
on the inner margin of the first segment of the exopodite of the
first thoracic leg in C. quadratus. This break, although slight, is
distinct in the Hawaiian specimens.

Caligus quadratus and the following 2 species of Caligus are diffi-
cult to separate. A discussion of the characteristics most readily
usable is given following the description of the third species.

Caligus productus Dana
Fiacures 43-45

Caligus productus Dana, 1853, p. 1354, pl. 94, fig. 4.—Steenstrup and Liitken,
1861, p. 357, pl. 3, fig. 6—Krgyer, 1863, p. 138. pl. 3, fig. 4a-i.—Brian,
1898, p. 10.—Bassett-Smith, 1899, p. 452.—Brian, 1906, p. 42.—Fowler, 1912,
p. 481.—Wilson, 1913, p. 208.—Brian, 1935, P. 53.—Pearse, 1952a, p.
15.—Causey, 1953a, p. 6; 1953b, p. 10; 1955, p. 5.—Shiino, 1959b, p.
14, figs. 6-8.—Causey, 1960, p. 329.—Shiino, 1960a, p. 471.—Hewitt
1963, p. 65.—Shiino, 1963a, p. 335.—Yamuguti, 1963, p. 58, pl. 73, fig.
3.—Kirtisinghe, 1964, p. 53, figs. 25-26.

Caligus monacanthi Krgyer, 1863, p. 59, pl. 3, fig. 2a-e——Wilson 1905b, p.
607; 1937, p. 424, figs. 6-19.

Caligus lobatus Wilson, 1935a, p. 1, pl. 1, figs. 1-10.

Caligus katuwo Yamaguti, 1936a, p. 6, pl. 4, fig. 55; pl. 5, figs. 56-68.—Shiino,
1954f, 1 fig—Nunes-Ruivo, 1956, p. 11, pl. 1, fig. b; pl. 2, fig. b.

DIsTRIBUTION AND HOSTS.—37+host records:

locality hosts references
Subtropical Atlantic Thymnus pelamys [sic] Dana, 1853
Thynnus Steenstrup and Liitken,
1861
Coryphaena
Monacanthus species Kroyer, 1863
Bonito Fowler, 1912
Naucrates ductor Wilson, 1935a

Neothunnus albacora Nunes-Ruivo, 1956

NO. 3574 COPEPOD CRUSTACEANS—LEWIS C17

locality hosts references
Gulf of Mexico Scomberomorus cavalla
Sarda sarda
Elops saurus

Coryphaena hippurus Pearse, 1952a

Scomberomorus maculatus Causey, 1953b

Pogonias cromis Causey, 1955
Eastern Pacific Bonito Wilson, 1937

Aucxis thazard
Coryphaena hippurus
Neothunnus albacora Shiino, 1959b
Seriola dorsalis
Paralabraz clathratus
P. maculatofasciatus
Verrunculus polylepos
Mackerel

Katsuwonus vagans
Sphyraena argentea
Scomberomorrus sierra
Coryphaena species
Seabass

Calamus brachysomus
Lutianus species

Centropomus species Causey, 1960
Japan Katsuwonus pelamis

Coryphaena hippurus Shiino, 1959b
‘Pacific’ Euthynnus pelamys Yamaguti, 1936a
Indian Ocean Coryphaena hippurus

Katsuwonnus pelamis

Euthynnus affinis Kirtisinghe, 1964
Mediterranean Chrysophrys aurata Brian, 1898

MarTERIAL.— Twenty-one females and 15 males (USNM 112904)
from the extenal surface of Katswwonus pelamis (Linnaeus) captured
near Oahu, Hawaii by Robert Stevenson. One male (USNM 112905)
from the external surface of Neothunnus macropterus (Schlegel)
captured by D. W. Strasburg off Kona, Hawaii. One immature
female (USNM 112906) from the buccal cavity of Auzis thazard
(Lacépéde) collected south of Nawiliwili, Kauai, Hawaii (USFWS, HMS
cruise 39, station 37). One female and 1 male (USNM 112907) from
the gill cavity of Huthynnus yaito (Jordan and Evermann) from an
unknown locality (USFWS). One male (USNM 112908) from the
external surface of Katsuwonus pelamis (Linnaeus) collected in the
Hawaiian region (USFWS, HMS cruise 34). One immature female
and 1 male (USNM 112909) from the external surface of 2 specimens
of K. pelamis (Linnaeus) captured at 20°41.5’N, 160°37.5’W (USFWS,
HMS cruise 39, station 17). One female and 1 male (USNM 112910)
from the gill cavity of K. pelamis (Linnaeus) captured at an unknown
locality (USFWS). Three females (retained by author) from the
gill cavity of 3 specimens of K. pelamis (Linnaeus) captured southeast

118 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

of Kauai, Hawaii (USFWS, HMS cruise 39). Three females and 1
male (retained by author) from the gill cavity of K. pelamis (Linn-
aeus) captured at 20°41.5’N, 160°37.5’W (USFWS, HMS cruise 39,
station 17). Four females and 2 males (retained by author) from the
gill cavity of K. pelamis (Linnaeus) captured 120 miles north of Oahu,
Hawaii (USFWS). One female and 1 male (retained by auther) from
the external surface and gill cavity of K. pelamis (Linnaeus) captured
at 2°S, 132°W (USFWS, JRM cruise 34, station 5).

MEASUREMENTS.— (In mm) 32 females and 22 males:

female male
mean range “mean range

Total length, excluding caudal setae 4.80 4.10-5.45 4.58 4.05-5.15
Length of cephalothorax including frontal

region 2.16 1. 83-2.33 2.39 2.10-2. 78
Width of cephalothorax 1.92 1.50-2.10 2.04 1. 88-2. 28
Length of genital segment 1.58 1.05-1.93 1.07 0. 90-1. 20
Width of genital segment 1.09 0.80-1.28 0.75 0. 65-0. 83
Length of abdomen 1.14 0.96-1.39 0.87 0. 47-1. 00
Length of caudal rami 0.24 0.19-0.27 0.26 0. 22-0. 29
Length of egg strings (17 strings) 2.86 1. 50-415

Derscription.— Female cephalothorax (fig. 43a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region approxi-
mately one-twelfth the length of cephalothorax, with membranous
margin along anterior edge; lunules extending the length of frontal
region. Lateral cephalothoracic margin slightly irregular, with mem-
brane extending laterally and second projecting medially, on ventral
surface; small but distinct concavity present in posterior lateral surface.
Posterior sinus (fig. 43c) narrower at opening than medially, with
fine membrane along outer margin. Medial posterior margin of
median cephalothoracic region smoothly convex, projecting poster-
iorly slightly past rounded lateral margins of median region and
slightly past posterior end of lateral cephalothoracic regions. Dorsal
cephalothoracic surface with numerous small, hairlike processes;
major dorsal grooves forming irregular H. Ocular region distinct,
in anterior third of cephalothorax; pigmented cups contiguous on
median longitudinal axis of body.

Female free fourth pedigerous segment distinct from cephalothorax,
indistinctly separable from genital segment dorsally, distinctly sepa-
rable laterally and ventrally. Segment broad medially, in region of
fourth leg attachment, narrower anteriorly and posteriorly. Genital
segment (fig. 43d) with anterior end narrow, anterior lateral margins
convex; posterior lateral surfaces forming pair of posteriorly pro-
jecting lobes. Fifth legs (fig. 43f) not visible dorsally, situated on
lateral ventral surface, at base of lobes; consisting of pair of knobs,
outermost with single plumose setule, innermost with 2 plumose set-

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 119

005mm.c

Ficure 43.—Caligus productus Dana, 1853, dorsal view: a, female; b, male; c, posterior
cephalothoracic sinus. Ventral view: d, female genital segment; ¢, male, same. Legs:
Ff, female fifth; g, male fifth; A, male sixth.

120 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

ules. Abdomen 1-segmented, elongate, indistinctly separable from
genital segment; with small concavity in anterior-ventral lateral
surface, concavity giving 2-segmented appearance but no complete
division indicated, either by cuticle or musculature. Posterior half
of abdomen broader than anterior, posterior margin angled laterally,
with anal concavity medially. Caudal rami (fig. 45h) width approxi-
mately nine-tenths the length, narrower proximally than distally,
distal half of inner margin plumose. Distal surface bearing 3 large
plumose setae medially, smaller plumose seta laterally and 2 plumose
setules, 1 on either side of setae.

Male cephalothorax (fig. 436) similar to that of female although
slightly larger. Free fourth pedigerous segment with small, plate-
like lateral extensions of tergal region, attached anteriorly to place of
fourth leg attachment. Genital segment (fig. 43¢) distinct from fourth
pedigerous segment, barrel shaped, posterior surface slightly irregular,
ventral surface with padlike swelling posteriorly. Fifth leg (fig. 43g)
situated on lateral ventral surface in posterior half of segment, simi-
lar to that of female; sixth leg (fig. 43h) situated on posterior ventral
surface, consisting of nodule bearing 2 plumose setules. Abdomen
2-segmented, distinct from genital segment dorsally, indistinctly
separable ventrally. First segment approximately five-ninths the
length of second, subrectangular; second segment slightly narrower
anteriorly than posteriorly, posterior end similar to that of female.
Caudal rami as in female.

Female and male antennule (fig. 446) 2-segmented, attached to
lateral-anterior ventral cephalothoracic surface. First segment ap-
proximately 1% times the length of second, broad proximally, distal
half of anterior surface angled to narrow distal surface, bearing
approximately 25 plumose setae and setules. Second segment rod
shaped, rounded distally, distal surface bearing approximately 12
naked setae and setules. Female antenna (fig. 44c) 3-segmented,
situated medial and posterior to antennule base. First segment
short, squat, almost completely fused with cephalothorax, with
spikelike posterior projection. Second segment broad proximally,
narrower distally; third segment fused with clawlike terminal process,
bearing setule-like accessory process proximally. Male antenna (fig.
44d) 3-segmented; first segment of each antenna in close proximity,
structurally similar to female first segment although spinelike pro-
jection smaller. Second segment elongate, with adhesion surface
extending as band in median third of segment, segment also bearing
spike-shaped projection from distal third of inner surface, projection
with adhesion surface. Third segment indistinctly separable from
clawlike terminal process, with setule-like accessory process proximally.

Female and male mandible (fig. 44c) 4-parted, distal part curved

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 121

Ficure 44.—Caligus productus Dana, 1853, ventral view: a, frontal region, right side show-
ing lunule; b, left antennule; c, female oral region, right side showing antenna, postan-
tennal process, mouth cone, mandible, maxillule, postoral process, and maxilla base;
d, male, same as c except mouth cone, mandible, and maxilla base not shown; g, left
maxilla; f, right maxilliped; g, sternal furca.

122 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

inward, inner surface with 12 denticulations. Female postantennal
process (fig. 44c) short, clawlike, with 2 nodules on proximal surface
and additional nodule just posterior to clawlike projection, all 3
nodules with several hairlike processes. Postantennal process of
male (fig. 44d) much longer than that of female, recurved; nodules
and hairlike processes as in female. Female postoral process (fig.
44c) long, spinelike; male (fig. 44d) as in female except for small,
knoblike projection on outer medial surface. Female and male max-
illule (figs. 44c, d) nodular, with 3 setules. Female and male maxilla
(fig. 44e) 2-segmented, situated lateral and slightly posterior to post-
oral process. First segment slightly shorter than second, second
rodlike, with fine membrane on posterior surface of distal half of
segment and 2 saber-like terminal processes. Innermost terminal
process approximately 1% times the length of outermost, with fine
membrane along inner margin; outermost process with finely frilled
membrane along both margins.

Female and male maxilliped (fig. 44f) 2-segmented, situated pos-
terior and medial to maxilla base. First segment strongly developed,
narrow proximally and distally, broad medially, with small, spinelike
projection from proximal half of inner surface. Second segment
short, distinct from clawlike terminal process, with small, setule-like
accessory process from distal inner surface. Female and male sternal
furca (fig. 44g) situated on median longitudinal axis posterior to
maxilliped bases. ‘Tines diverging, rounded distally.

Second segment of exopodite of first leg without large plumose
setae along inner margin but with 3 minute projections in position
normally occupied by setae. For nature of legs and armature, see
figures 45a-g and table 16.

TaBLE 16.—Armature of thoracic legs I-IV of the female and male of
Caligus productus Dana, 1852

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal Se Ee
Plate
2 1 2 3 i 2 3
I | Outer 2s*,p rh H,2dH,dpP’
Inner d,p C,C 3s
II | Outer m,p | mymH mH mH,2Q |C}|D 3P
m
Inner 23,2 m;8 |) .¢, GE c,5P IPaiGi2k ¢,3P
III | Outer d,m,p s,H cp’ @Sp,b C1) G2e
m
Inner d,3s,P,s,m,s Cie, ¢3P. PeiTeaP
IV | Outer | 4s 4s,dmHf 2dmH,2H

* More than one hairlike process.
t Very fine denticulations on membrane.

Discussion.—The synonymy does not show several references
included by Shiino (1959b) (Rathbun, 1884; Wilson, 1935a), and

No. 3574 COPEPOD CRUSTACEANS—LEWIS 123

0.05mm 9d

++
02mm Sc

Figure 45.—Caligus productus Dana, 1853, right thoracic legs: a, first, anterior view; },
proximalmost setule on inner margin of second segment of exopodite of first; c, distal
region of second segment of exopodite of first, anterior view; d, second, anterior view;
e, third, anterior view; f, exopodite of third, posterior view; g, fourth, anterior view;
h, caudal ramus, ventral view.

124 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Wilson’s 1937 reference (“C. monacanthi”) is changed. In an ex-
amination of the material identified by Rathbun as Caligus productus
(USNM 6109) for his 1884 publication, all of the specimens were
found to belong to C. quadratus Shino. The female specimen that
Wilson (1935a) identified as C. productus (USNM 69776) is not
C. productus and does not belong to either of the 2 closely related
species here described. ‘The material that Wilson (1937) described as
C. monacanthi from Panama Bay (USNM 69867, 69868) is all C. pro-
ductus. Shiino (1959b) indicates that the males identified as C. mona-
canthi by Wilson are C. quadratus and the females C. productus. This
indication is presumably based on Wilson’s statement (1937, p. 426)
that ‘the terminal segment of the first legs [of the male] carries the
usual 3 plumose setae on its posterior margin.”” This indicates that
the males are not C. productus inasmuch as the absence of the 3 setae
is characteristic of this species. From an examination of Wilson’s
material, however, it was found that the males as well as the females
do not possess any plumose setae on the posterior margin of the
second (terminal) segment of the exopodite of the first thoracic leg.
Additionally, all of these specimens have other characteristics that
associate them with C. productus. An examination of the type
material for C. lobatus Wilson, 1935a (USNM 64059, 64060) supports
Shiino’s conclusion that this species is also a synonym of C. productus.

The species described as Caligus dentatus Heegaard, 1962, shows
some of the characteristics found in C. productus, particularly the
absence of plumose setae on the second segment of the exopodite of
the first thoracic leg. Whether this species is synonymous with C.
productus or is distinct remains questionable, and further, undamaged
material should be adequately described, figured, and compared with
associated species before the relationship can be realized.

For a comparison of C. productus with closely related species found
in Hawaiian waters, see the discussion section following the descrip-
tion of the following species.

Caligus bonito Wilson

Figures 46—48

Caligus bonito Wilson, 1905b, p. 589, pl. 13, figs. 150-158, text figs. 5, 12-15, 29,
30, 35, 37, 38, 40-45.—Rathbun, 1905, p. 89.—Brian, 1924, p. 13.—Wilson,
1932, p. 407, fig. 255.—Brian, 1935, p. 188, fig. 14—Yamaguti, 1936a, p. 8,
pl. 5, figs. 69-71; pl. 6, figs. 72-85.—Bere, 1936, p. 582.—Causey, 1953a,
p. 4; 1953b, p. 8.—Nunes-Ruivo, 1956, p. 6, pl. 2, fig. a—Causey, 1960,
p. 328.—Shiino, 1960b, p. 527, fig. 1; 1963a, p. 336.—Yamaguti, 1963, p. 50,
pl. 56, fig. 3.

?Caligus kuroshio Shiino, 1959c, p. 51, figs. 1-2.—Pillai, 1963, p. 80, fig. 9.—
Yamaguti, 1963, p. 55, pl. 71, fig. 3.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 125

DISTRIBUTION AND Hosts.—20 host records:

locality hosts references
Western North Atlantic Bonito Wilson, 1905b
Gymnosarda pelamis Rathbun, 1905
Subtropical Atlantic | Cybium Brian, 1924
~ Katsuwonus pelamis Nunes-Ruivo, 1956
Gulf of Mexico Pomatomas saltatrix

Scomberomorus maculatus
Lutianus griseus

Mugil cephalus
Oligoplites saurus

Scomberomorus cavalla Bere, 1936

Gymnosarda alleterato Causey, 1953a

Sarda sarda Causey, 1953b
Eastern Pacific Sarda chilensis

?Lutjanus species Causey, 1960

Katsuwonus pelamis Shiino, 1963a
Japan Euthynnus pelamis Yamaguti, 1936a

Katsuwonus pelamis

Thunnus thynnus Shiino, 1959¢
Indian Ocean ~ Euthynnus affinis Pillai, 1963
Mediterranean Pelamys sarda Brian, 1935

MateriAL.—Three females (USNM 112911) from the gill cavity
of several specimens of Katsuwonus pelamis (Linnaeus.) captured
100 miles north of Oahu, Hawaii (USFWS).

MEASUREMENTS.—(In mm) 3 females:

Total length, excluding caudal setae 6.20, 6.50, 6.45
Length of cephalothorax, including

frontal region 2.70, 3.05, 3.10
Width of cephalothorax 2.50, 2.85, 2.85
Length of genital segment 2.00, 2.05, 2.00
Width of genital segment 1.50, 1.35, 1.60
Length of abdomen 1.25, 1.35, 1.25
Length of caudal rami 0.19, 0.20, 0.22

Length of egg strings (all 3 nonovigerous)

DESCRIPTION OF FEMALE.—Cephalothorax (fig. 46a) ovoid, con-
sisting of cephalon and first 4 thoracic segments. Frontal region
approximately one-tenth the total length of cephalothorax, with
fine membrane along anterior margin. Lunules distinct, slightly
longer than greatest length of frontal region. Lateral cephalotho-
racic margin slightly irregular, with narrow membrane projecting
laterally and second projecting medially. Posterior lateral surface
with small indentation. Posterior sinus (fig. 465) U-shaped, with
fine membrane along outer margin. Posterior medial surface extend-
ing posteriorly slightly past lateral cephalothoracic regions, with
narrow, median projection covering junction of cephalothorax and
free fourth pedigerous segment. Major dorsal cephalothoracic

126 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

grooves forming irregular H, transverse groove of H distinct, ir-
regular. Ocular elements distinct, contiguous on median longi-
tudinal axis of body, in anterior third of cephalothorax.

Free fourth pedigerous segment wedge shaped, distinctly separable
from genital segment ventrally, appearing fused dorsally. Genital
segment (fig. 46c) constricted anteriorly, with pair of lobate projec-
tions posteriorly, distinct from abdomen. Fifth legs (fig. 46d)

Ficure 46.—Caligus bonito Wilson, 1905, female: a, dorsal view; b, posterior cephalothoracic
sinus, dorsal view. Ventral view: c, free fourth pedigerous segment, genital segment,
and anterior region of abdomen; d, right fifth leg; ¢, caudal ramus.

minute, situated on ventral posterior lateral surface, consisting of
single, lightly plumose setule laterally, pair of lightly plumose setules
just medial to lateral setule.

Abdomen 1-segmented, anterior half appearing swollen, degree of
swelling varying in 3 Hawaiian specimens. Posterior half of abdomen
rounded on dorsal posterior surface, with pair of concavities on ven-

No. 3574 COPEPOD CRUSTACEANS—LEWIS 127

tral surface forming place of attachment of caudal rami, anal inden-
tation slight. Caudal rami (fig. 46e) narrower proximally than distal-
ly, distal half of inner surface slightly swollen, swelling terminating
posteriorly in slight knob bearing plumose setule. Distal inner
margin plumose, distal surface with 3 plumose setae originating ventral

Ficure 47.—Caligus bonito Wilson, 1905, female, ventral view: a, right side of frontal
region showing lunule and antennule base; ), left antennule; c, oral region, left side show-
ing antenna, postantennal process, mouth cone, mandible, maxillule, postoral process,
and maxilla base (ma-2); d, left maxilla; ¢, left maxilliped; f, sternal furca.

to small median lobe, plumose setule present lateral to setae and 2
minute setules present, 1 on each side of ramus.

Antennule (fig. 476) 2-segmented, attached to lateral-anterior
ventral surface of cephalothorax and lateral-posterior ventral surface
of frontal region. First segment slightly longer than second, proxi-
mal half broad, distal tapered to narrow distal end, anterior ventral

128 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

surface with approximately 23 plumose setae and setules, anterior
dorsa] surface with pair of plumose setules. Second segment rodlike,
with approximately 12 naked setules distally. Antenna (fig. 47c)
3-segmented, situated posterior and medial to antennule base. First
segment short, squat; second segment broad proximally, tapered to
lightly narrower distal end, with small, horseshoe-like projection on
lateral anterior surface in distal half of segment. Third segment and
clawlike terminal process fused, single setule-like accessory process
present.

Mandible (fig. 47c) 4-parted, rodlike, distalmost part curved me-
dially, with 12 denticulations on inner surface. Labrum with minute
denticulations on distal surface. Postantennal process (fig. 47c)
situated lateral to antenna base, consisting of clawlike process and
3 nodules, each with several hairlike processes. Postoral process
(fig. 47c) long, spinelike, distal end flattened slightly, appearing spade
shaped. Maxillule (fig. 47c) nodular, with 3 setules. Maxilla (fig.
47d) 2-segmented, situated lateral and slightly posterior to postoral
process. First segment approximately four-fifths the length of
second; second elongate, slightly swollen medially, with fine membrane
on medial swelling and 2 saber-shaped terminal processes. Inner-
most terminal process approximately twice length of outer, with fine
membrane along both margins; outer terminal process with frilled
membrane along outer margin.

Maxilliped (fig. 47e) 2-segmented, situated posterior and medial to
maxilla base. First segment strongly developed, proximal half ta-
pered to narrow proximal end, curved strongly; distal half tapered to
irregular distal end. Second segment short, separable from claw-
like terminal process, bearing single setule-like accessory process
from distal inner surface. Sternal furca (fig. 47f) situated on median
longitudinal axis of body approximately halfway between maxil-
liped bases and first thoracic legs. ines chisel shaped, diverging
slightly; furca attached to small, platelike projection.

For nature of legs and armature, see figure 48 and table 17.

Discussion.—The Hawaiian material differs from the original
description of Caligus bonito Wilson, 1905b, in one major respect, the
longer abdomen of C. bonito. In an examination of the type material
of C. bonito (USNM 6035, 41975) considerable variation was noted
in both the length and width of the abdomen. This variation appears
to be due to the maturity of the specimens. The length of the region
posterior to the swollen anterior portion of the abdomen of the
Hawaiian specimens approximates the length of the region behind
the slight constriction (joint between segments) of the abdomen of
C. bonito. Based upon this, upon the absence of egg strings in the
Hawaiian material (although formative strings are present in the

129

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

—

-. 2S i
y ; ttt
5 ; wee

HHH
prt

ow SSN
305 — Ss
Ss

Ye <

005 mm.

Ficure 48.—Caligus bonito Wilson, 1905, right thoracic legs, anterior view: a, first; b. second

segment of exopodite of first; ¢, second; d, third; e, exopodite of third (posterior view);

f, fourth.

9

6—321— 67

22

130 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

TABLE 17.—Armature of thoracic legs I-I ee the female of Caligus bonito Wilson,
1905

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal a
Plate
1 2 1 2. 3 1 | 2 3
1 | Outer $,p rh 3dH,P’ ye
Inner p c 3P
II | Outer m,p | m,fm,mH | mH fn oH; C|D D,4P
m Q,
Inner se DS Wiel: c,P 4P Pl ¢.2P CP
Iil | Outer d,m,p s,H c,S,p’ c,3p’,P e jc
Inner m d,2s,P,s, Ge ¢,3P PAC Gr
m,s
IV | Outer 5s,p s,fm,mH fae ein,
| 3

genital segment), upon the change that occurs in the length of the
abdomen in other species (e.g., Caligus coryphaenae), and upon
Wilson’s statement (1905b, p. 589) that ‘“‘the joints [are] about equal
in immature females’ (the 2-segmented condition is not evident in
the Hawaiian specimens), the present author feels that the difference
in the length of the abdomen between the Hawaiian material and
Wilson’s original description of C. bonito is due to the immature condi-
tion of the Hawaiian specimens.

The differences between the Hawaiian specimens and Caligus
kuroshio Shiino, 1959c, are primarily differences in the small armature
elements of the appendages. The caudal rami of the Hawaiian
specimens possess a small, nodular projection on the inner distal
surface that bears a plumose setule while Shiino’s figure does not
show the nodule. Shiino indicates (table I) that there is a large,
plumose seta on the second segment of the protopodite of the second
thoracic leg which is not present in the Hawaiian specimens; how-
ever, he does not figure this armature element and its presence would
make OC. kuroshio unique. Shiino describes a 2-segmented condition
for the exopodite of the third thoracic leg while the present author
describes a 3-segmented condition for the Hawaiian specimens of
C. bonito. Shiino’s “first segment” is what the present author calls
segments 1 and 2. The large, inwardly curved spine is at the distal
end of the first segment (fig. 48¢) while the plumose seta and
naked setule (1h in Shiino’s table J) are on the second segment.
Shiino also uses the smaller size of his specimens to differentiate C.
kuroshio from (©. bonito. The size of Wilson’s holotype female is
8.3 mm, the largest specimen in Shiino’s 1959c description is 6.92 mm
(range 5.39-6.92), Yamaguti (1936a, C. bonito) lists 4.5-5.6 mm,
Brian (1935, C. bonito) lists 8mm, Brian (1924, C. bonito) lists 4,5,7
and 8 mm females, and a male at 5 mm, Shiino (1960b, C. bonito) gives
the largest female at 6.7 mm, while the Hawaiian female specimens
range from 6.2-6.5 mm.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 131

Remarks.—Caligus quadratus, C. productus, and ©. bonito are
difficult to separate not only because of the similarity in their mor-
phology but also because they are found on the same species of hosts
and not too infrequently they may be on the same host specimen.
Although this leads to some speculation on the relationships of the
three, there is some evidence to support the belief that they are
distinct species. ‘This evidence, or part of it, can be used to separate
the species without going through a complete morphological analysis.
Assuming that the general body shape and number of segments in
the thoracic legs agrees with the figures (i.e., that the specimens
belong to this species complex), Caligus productus can be separated
by the absence of the 3 plumose setae normally present on the inner
margin of the second segment of the exopodite of the first thoracic
leg. Both Caligus quadratus and C. bonito possess these 3 setae,
but C. bonito possesses a series of distinct denticulations on the outer
margin of the first and second segments of the endopodite of the
second thoracic leg. Caligus quadratus, on the other hand,
possesses a patch of stiff plumosities in the same region. The dif-
ference between stiff plumosities and denticulations is not great
but it is a distinct difference and sufficiently diagnostic to be used
as a key characteristic.

Caligus asymmetricus, Kabata 1965b
Figures 49-51

Caligus thynni Pillai, 1963, p. 89, fig. 14.
Caligus asymmetricus Kabata, 1965b, p. 110, figs. 1B, D, E.
Caligus asymmetricus Pillaii—Kabata, 1965b, pp. 109, 110.

DisTRIBUTION AND Host.—Trivandrum, South India, Huthynnus
afinis (in Pillai, 1963); Queensland, Australia, Huthynnus alleteratus
(in Kabata, 1956b).

Remarks.—Dana (1853, p. 1353) described the caligid Caligus
thymni “from the external surface of a Bonito (Thymnus pelamys)

.’ Presumably the species was named after the host although
the eyeline of the generic part of the host name (Thymnus) has, to
my knowledge, never been used in the taxonomy of the tunas, the
name being annie The name Caligus thymni again appears
(p. 219) in a handwritten ‘Catalogue of Crustacea of the U.S. Expl.
Exped. during 1838-’42 . . . collected and described by James D.
Dana. Geologist and Zoologist Exp.,” dated 1856 by Dana and
located in the U.S. National Museum. Yamaguti (1963, p. 61)
uses “C. thymni [misprint for thynni] Dana, 1853 . . .”’ in his account.
It is highly probable that thymni was not a misprint but a lapsus
by Dana, based on his erroneous spelling, Thymnus, of the host

132 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Thynnus. In either case, Caligus thymnus is an incorrect original
spelling and must be emended to Caligus thynni Dana. Caligus
thynni Pillai (1963), a distinct species from Huthynnus affinis, thereby
becomes a junior homonym.

After the publication of Caligus thynni Pillai, the author of the
name became aware of the problem of homonymy and submitted
a manuscript (‘““Copepods parasitic on Indian marine fishes—a
review”) containing the replacement name Caligus asymmetricus.
During this period, however, specimens of the species had also been
found on a fish captured near Green Island, Queensland, Australia,
and had been submitted to Z. Kabata for identification. After
corresponding with Pillai and learning of the replacement name
that was then in press, Kabata submitted a manuscript containing
the replacement name as well as a description of the species. It
is unfortunate that the manuscript containing Pillai’s replacement
name was not published prior to the publication of Kabata’s manu-
script. Since, however, Kabata (1965b) was the first to publish
the replacement name, he, therefore, becomes the author of that name.

The present author deeply appreciates the helpful correspondence
with Dr. Pillai and the discussion with Dr. Kabata concerning some
of the above details.

Matrrrau.—One adult female and 3 males (USNM 112912) from
the external surface of Huthynnus yaito (Jordan and Evermann)
captured off Moku Manu Island, near Oahu, Hawaii. One adult
female (USNM 112913) from the gill cavity of Huthynnus yaito (Jor-
dan and Evermann) captured at French Frigate Shoals (USFWS,
HMS cruise 39, station 32).

MEASUREMENTS.—(In mm) 2 females and 3 males:

female male

Total length, excluding caudal setae 3.00,3.35 3.15, 2. 80, 2. 55
Length of cephalothorax, including frontalregion 1.95,2.05 2.038, 1. 88, 1. 68
Width of cephalothorax 1. 58,1.65 1. 55, 1. 53, 1. 30
Length of genital segment 0. 80,0.90 0. 60, 0. 55, 0. 50
Width of genital segment 0. 70,0. 83 0. 53, 0. 55, 0. 48
Length of abdomen 0. 31,0.34 0. 41, 0. 37, 0. 33

OL OL1L 0: 13; O81, 0610

~

Length of caudal rami
Length of egg strings (neither female ovigerous)

DescrieTion.—Female cephalothorax (fig. 49a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region approxi-
mately one-thirteenth the greatest length of cephalothorax, with
fine membrane along anterior margin and pair of lunules overlapping
division between frontal region and remaining cephalothorax ven-
trally. Lateral margin of cephalothorax slightly irregular, with fine
membrane projecting laterally and second membrane projecting med-
ially on ventral surface. Posterior lateral surface with small but

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 133

a
, Legg ND \

/ $4 4+ 1 4 1 4
[Sf Fee er Se i 1.0 mm.2 ar

eH HH
Q03mm.c"

+, _———_+—_—_ b+ 4
0.3mm.2 0.3 mm.c"

Ficure 49.—Caligus asymmetricus Kabata, 1965b, dorsal view: a, female; b, male; c, posterior
cephalothoracic sinus. Ventral view: d, female genital segment; e, male, same; f, female
right fifth leg; g, male left fifth leg; h, male left sixth leg; 2, caudal ramus.

134 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

distinct concavity. Posterior sinus (fig. 49c) narrow, bearing fine
membrane along outer surface. Posterior median cephalothoracic
region broadly convex, projecting posteriorly well past posterior lat-
eral cephalothoracic region, overlapping anterior end of free fourth
pedigerous segment. Major dorsal cephalothoracic grooves forming
irregular H; dorsal surface with several spinules. Ocular region
distinct, pigmented cups contiguous on median longitudinal axis in
anterior third of cephalothorax.

Female free fourth pedigerous segment wedge shaped, irregular
posteriorly, indistinctly separable from genital segment. Genital
segment (fig. 49d) narrower anteriorly than posteriorly, lateral margins
latly convex, posterior flatly concave. Fifth legs (fig. 49f) situated
on ventral-posterior lateral surface, consisting of pair of knobs, ante-
riormost with single plumose setule, posterior with 2 plumose setules.

Female abdomen indistinctly separable from genital segment, con-
sisting of 1 or 2 segments. Indication of 2-segmented condition given
by constriction anteriorly although indication superficial. Posterior
two-thirds of abdomen subrectangular, posterior end tapered at junc-
tion with caudal rami, indented at anal opening. Caudal rami (fig. 497)
short, division between abdomen and rami incomplete. Lateral mar-
gins of rami essentially parallel, distal surface irregular, with 3 large
plumose setae medially, 2 plumose setules on outer surface and 1
on inner surface; inner lateral margin of rami lightly plumose distally.

Male cephalothorax (fig. 495) and free fourth pedigerous segment
as in female although fourth pedigerous segment of figured male
retracted under median posterior region of cephalothorax more than
in female. Genital segment (fig. 49e) barrel shaped, distinct from
both fourth pedigerous segment and abdomen. Fifth legs (fig. 49g)
situated on posterior half of lateral surface, consisting of 3 plumose
setules grouped as in female but without knobs. Sixth legs (fig. 49h)
on slight ventral swelling posterior and medial to fifth legs, consisting
of 2 plumose setules. Abdomen distinctly 2-segmented, first segment
short, slightly more than half the length of second, second as in pos-
terior two-thirds of female abdomen. Caudal rami as in female.

Female and male antennule (fig. 506) 2-segmented, attached to
lateral-anterior ventral surface of cephalothorax and lateral-posterior
ventral surface of frontal region. First segment approximately 1%
times the length of second, broad medially, narrower proximally and
distally, distal two-thirds of anterior and anterior ventral surface
bearing approximately 26 lightly plumose setae and setules, including
1 Jong, anteriorly projecting seta. Second segment club shaped,
bearing approximately 12 naked setae and setules distally, including
5 long setae (length slightly shorter than length of segment). Female
antenna (fig. 50c) 3-segmented, situated posterior and medial to anten-

No. 3574 COPEPOD CRUSTACEANS—LEWIS 135

He}
02mm.2cs O.1mm28o" b

bee + +f
OAmm&o

Ficure 50.—Caligus asymmetricus Kabata 1965b, ventral view: a, frontal region, left side,
showing lunule and antennule base; b, right antennule; c, female oral region, right side
showing antenna, postantennal process, mouth cone, mandible, maxillule, postoral process,
and maxilla base; d, male left antenna and postantennal process; e, left maxilla; f, right
maxilliped, g, sternal furca.

136 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

nule base. First segment short, squat, heavily sclerotized, with
very slight indication of posterior projection at base; second segment
slightly more than two-thirds as wide as long, narrower distally
than proximally. Third segment fused with clawlike terminal process,
bearing small, lappet-like projection on proximal posterior surface
and small spinule-like accessory process at distinct break in scleroti-
zation in proximal half of fused segment and terminal process. Male
antenna (fig. 50d) 3-segmented, first segment elongate, heavily sclero-
tized, irregular; second segment shorter than first, broad proximally,
tapered to narrower distal end, with ridged, lappet-like adhesion pad
on distal inner surface, just proximal to third segment articulation
surface and apparently receiving terminal process of third segment
when segment flexed. Third segment fused with short, clawlike
terminal process, bearing setule-like accessory process on anterior
surface.

Female and male mandible (fig. 50c) 4-parted, distalmost part
flattened, curved inward, with 12 denticulations along inner surface.
Female postantennal process (fig. 50c) situated lateral to antenna,
consisting of short, clawlike projection and 3 nodules, 2 on base of claw,
third posterior to claw, each with several hairlike processes. Male
postantennal process (fig. 50d) immediately lateral to antennal base,
clawlike process much longer than in female and strongly curved dis-
tally, nodes as in female. Female and male postoral process (fig. 50c)
long, spinelike, curved outward distally. Female and male maxillule
(fig. 50c) nodular, with 3 setules. Female and male maxilla (fig. 50e)
2-segmented, situated lateral and slightly posterior to postoral process.
First segment slightly shorter than second, more strongly developed;
second segment elongate, tapered proximally, with fine membrane
along inner medial surface and 2 saber-shaped terminal processes.
Innermost terminal process slighty less then 1% times the length of
outermost, with fine membrane along inner margin; outermost process
with finely frilled membrane along both margins.

Female and male maxilliped (fig. 50f) 2-seemented, situated poste-
rior and medial to maxilla base. First segment strongly developed,
broad medially, tapering proximally and distally, proximal end
strongly curved, heavily sclerotized; small, subconical projection pres-
ent on inner medial surface. Second segment distinct from clawlike
terminal process, with setule-like accessory process on distal inner
surface. Female and male sternal furea (fig. 50g) situated on median
longitudinal axis of body, posterior to maxilliped bases. Base of furca

Ficure 51.—Caligus asymmetricus Kabata, 1965), right thoracic legs, anterior view: a, first;
b, second segment of exopodite of first; c, second; d, third; e¢, exopodite of third (posterior
view); f, fourth.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 137

ae s ‘ a Oe
a» x Uf, ype = : a

}) M1)
SULA =
Z J NGS

S/
a (27 Ly ,
- AS re

AZ.

ORR SSS

(Oi =F
ACG ESS eS,
NSS Kees i:

) = S ak xX . ve
\\ y RN . NG WS 7 = A re ee
coy = \ \ \ < ‘ fe!

a oS YS js A
NX bt Y\\4 i ;

> it
>>
o2>>
Seely Sea

fp; 4 H H
0.05 mms"

138 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

between 2 concave, lappet-like projections; tines sharply pointed,
essentially parallel (right tine broken on figured female specimen, tines
more irregular on second female specimen).

For nature of legs and armature, see figures 51a—f and table 18.

TABLE 18.—Armature of thoracic legs I-IV of the female and male of Caligus asym-
metricus Kabata, 1965b

Inter- Protopodite Exopodite Endopodite
Leg | Surface || podal
Plate
2 1 2 3 1 2 3
I | Outer ss,p rh 3fmH,P’
Inner p c,c iP
II | Outer m,p m,mH mH 2mH,Q C | Dye ¢,3P
m
Inner s-2 m,s Cae CE 5P Pera 3P
Iit | Outer d,m,P mH ¢,p’ c,3p’,P Ceine2e
m
Inner d,2s,P,s, cae c,3P P | 4P
m,s
IV | Outer p sfm,mH | fm,mH,
fm,mH,
fm,H

Discusston.—The Hawaiian specimens differ from the original
description of Caligus asymmetricus Kabata in several respects. The
genital segment is more swollen, a characteristic that may be due
to maturity. The tines of the sternal furca of the figured specimen
are not quite as angular as those figured by Pillai (1963), although the
tines on the other female are similar to Pillai’s figure. The terminal
processes of the exopodite of the first thoracic leg are structurally
similar to those figured by Pillai but they are at the end of the seg-
ment, not subterminal as Pillai figures them. The denticulations
on the outer margin of the second segment of the endopodite of the
figured second thoracic legs differ in number (6 instead of 9) but
not in character. There is, however, some variation in the number of
denticulations in the Hawaiian material, even in the same specimen,
the number ranging from 5-14 denticulations. Pillai (1963) based
his description on 1 female specimen (nonovigerous?) from a speci-
men of Luthynnus affinis along with several specimens of C. kuroshio
(=?C. bonito Wilson, 19055), while the Hawaiian material came
from a specimen of Huthynnus yaito.

Caligus pelamydis Kr¢gyer
Fieures 52, 53

Caligus pelamydis Krgyer, 1863, p. 124, pl. 4, fig. 4a-—g.—Richiardi, 1880, p. 148.—
Valle, 1882, p. 245.—Carus, 1885, p. 357.—Bassett-Smith, 1899, p. 452.—
Brian, 1899b, p. 198.—Wilson, 1905b, p. 594, pl. 13, figs. 154-161; pl. 14, fig.
16la.—Brian, 1906, p. 43.—Brady, 1910, p. 589, fig. 69.—Stebbing, 1910,

No. 3574 COPEPOD CRUSTACEANS—LEWIS 139

p. 558.—Guiart, 1913, p. 7.—Scott and Scott, 1913, p. 57, pl. 7, figs. 2,3;
pl. 9, figs. 1-5; pl. 71, fig. 14—Scott, 1929, p. 89.—Wilson, 1932, p. 406,
fig. 254.—Heegaard, 1943b, p. 5.—Causey, 1953a, p. 5; 1953b, p. 10; 1955,
p. 4.—Nunes-Ruivo, 1956, p. 5, pl. 1, fig. a—Barnard, 1955, p. 245.—
Causey, 1960, p. 329.—Hewitt, 1963, p. 78, fig. 6.

Caligus scomberi Bassett-Smith, 1896b, p. 11, pl. 3, fig. 2; 1899, p. 450.

Caligus scombri Scott T., 1901, p. 148, pl. 5, fig. 9-10.—Scott A., 1906, p. 196,
pl. 6.—Guiart, 1913, p. 6.

DISTRIBUTION AND HostTs.—12 host records:

locality hosts references
North Atlantic Pelamys sarda Krgyer, 1863
Scomber scombrus Scott and Scott, 1913
East coast of North Unknown Wilson, 1905b
Atlantic
South Atlantic Unknown Brady, 1910
Stebbing, 1910
Sarda sarda Nunes-Ruivo, 1956
Gulf of Mexico Pogonias cromis
Scomberomorus cavalla Causey, 1953a
Sarda sarda Causey, 1953b
New Zealand Thyrsites atun Hewitt, 1963
Mediterranean Scomber scomber
Pelamys sarda Richiardi, 1880
Unknown Guiart, 1913

Marertau.—One female (USNM 112914) from the gill cavity of
Euthynnus yaito (Jordan and Evermann) captured at French Frigate
Shoals (USFWS, HMS cruise 39, station 32).

MEASUREMENTS.—(In mm) 1 female:

Total length, excluding caudal setae 4. 05
Length of cephalothorax, including frontal region 1. 38
Width of cephalothorax 1. 20
Length of genital segment 1.18
Width of genital segment 110
Length of abdomen 1. 45

On 07,

Length of caudal rami
Length of egg strings (female nonovigerous)

DeEscRIPTION OF FEMALE.—Cephalothorax (fig. 52a) ovoid, con-
sisting of cephalon and first 4 thoracic segments. Frontal region
narrow, middle of anterior surface distinctly indented, margin with
fine membrane. Lunules (fig. 52f) large but filmy, extending
posteriorly, on ventral surface, past junction of frontal region and
remaining cephalothorax. Lateral margin of cephalothorax slightly
irregular, bearing fine membrane extending laterally and second
extending medially on ventral surface. Posterior lateral cephalo-
thoracic surface with small concavity. Posterior sinus (fig. 526)
narrow, constricted at opening, with fine membrane along outer
surface. Posterior median cephalothoracic surface extending past

140 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

lateral surface, posterior end overlapping junction of cephalothorax
and free fourth pedigerous segment, margin flatly convex. Major
dorsal cephalothoracic grooves forming irregular H, posterior longi-
tudinal legs and crosssbar continuous, heavily sclerotized. Ocular
region distinct.

Free fourth pedigerous segment short, distinct from cephalothorax,
incompletely separable from genital segment; segment sharply angled
posteriorly and anteriorly, from region of fourth leg attachment.
Genital segment (fig. 52c) broad for most of length, anterior end con-
cavely angled inward, posterior end with pair of broad lappets dor-
solaterally, lappets extending posteriorly past junction of genital
segment and abdomen. Fifth legs (fig. 52d) consisting of single,
plumose setule arising from ventral lateral surface at anterior end of
lappets. Spermatophores visible on single Hawaiian specimen,
attached immediately ventral to junction of genital segment and
lateral surface of abdomen.

Abdomen indistinctly 2-segmented, junction with genital segment
distinct. First segment approximately five times the length of
second, broadest in anterior medial portion, irregularly curved to
anterior and posterior ends; second segment with flatly convex lateral
margins, anal indentation slight. Caudal rami (fig. 52¢) approxi-
mately 1% times as long as wide, distal half of inner surface plumose;
three plumose setae present on distal surface, single plumose seta on
outer distal surface and 1 plumose setule on either side of setae.

Antennule (fig. 52g) 2-segmented, attached to lateral-anterior
ventral surface of cephalothorax. First segment slightly less than
twice the leneth of second, broader proximally than distally; ventrally
curved distal half of anterior surface with approximately 18 plumose
setules. Second segment rodlike, distal end with approximately 12
naked setules. Antenna (fig. 52h) 3-segmented, attached medial and
posterior to antennule base. First segment squat, irregular, heavily
sclerotized; second segment tapered slightly from broad proximal to
slightly narrower distal end, proximal outer surface with small, sub-
triangular projection. Third segment and clawlike terminal process
indistinctly separable, segment elongate, with small, lappet-like pro-
jection from posterior medial surface.

Mandible (fig. 52h) 4-parted, rodlike, distalmost part curved
inward, with 12 denticulations on inner surface. Postantennal process

Figure 52.—Caligus pelamydis Krgyer, 1863, female: a, dorsal view (broken line=nick in
specimen). Ventral view: b, posterior cephalothoracic sinus; c, genital segment (anterior
end directed downward); d, fifth leg; e, caudal ramus; f, right side of frontal region showing
lunule and antennule base; g, right antennule; h, oral region, left side showing antenna,
postantennal process, mouth cone, mandible, maxillule, postoral process, and maxilla
base; 7, left maxilla; 7, left maxilliped; &, sternal furca.

141

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

142 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

(fig. 52h) situated lateral and slightly posterior to antenna base,
consisting of short, heavily sclerotized, dagger-shaped projection and
3 nodules, 2 at base of projection, third slightly posterior, each with
several hairlike processes. Postoral process (fig. 52h) large, heavily
sclerotized, spinelike. Maxillule (fig. 52h) nodular, situated adjacent
to inner proximal surface of postoral process, with 3 setules distally.
Maxilla (fig. 327) 2-segmented, situated slightly lateral and posterior
to postoral process. First segment approximately four-fifths the
length of second, second elongate, with narrow, frilled membrane
on medial inner surface, with 2 saber-shaped terminal processes.
Innermost terminal process approximately 1% times the length of
outermost, both with fine membranes along inner and outer margins.

Maxilliped (fig. 527) 2-segmented, situated medial and posterior
to maxilla base. First segment strongly developed, with narrow,
heavily sclerotized, strongly curved proximal articulation and muscle
attachment surface. Second segment indistinctly separable from
clawlike terminal process, with single, small, setule-like accessory
process from distal inner surface. Sternal furca (fig. 52k) situated
between and slightly posterior to maxilliped bases, tines broad,
parallel, terminating in rounded surface. Furea associated with
platelike area of heavy sclerotization extending posteriorly almost
to interpodal plate of first thoracic legs and extending laterally past
line running along longitudinal axis of body, through region of
attachment of maxilliped bases.

Fourth thoracic leg uniramous, poorly developed, similar, in seg-
ment constitution, to that of caligids in last chalimus stages. For
nature of legs and armature, see figure 53 and table 19.

TABLE 19.—Armature of thoracic legs I-IV of the female of Caligus pelamydis
Krgyer, 1863

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal a
late
1 2 1 2 3 1 2 3
I | Outer Sss,D h 4H Ss
Inner p c 3P s
II | Outer m m,fm,H H H,mH,Q Cc C,3P
m
Inner s,P m,s | ¢,P e,P 5P a iCre 3P
III | Outer m,d,p H C.ps c,3p’,P Ce e2P
m
Inner 2s,P,s,m,s P 3P Para
Iv* | Outer 2s,p s,fm,pH fm,2pH,p’,
3pH,fm

i eee mentation of this leg in doubt: protopodite and first segment of exopodite recognizable, others ques-
ionable.

Ficure 53.—Caligus pelamydis Kréyer, 1863, female, right thoracic legs, anterior view:
a, first; b, distal end of second segment of exopodite of first; c, second; d, third; ¢, exopodite
of third; f, fourth; g, exopodite of fourth.

RS

LAL

7
&

XY
XX]
|

Zz

F
y

p

a

COPEPOD CRUSTACEANS—LEWIS

144 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Discusston.—Wilson (1905b) suggests that Caligus pelamydis
Kroyer, and C. scombert Bassett-Smith are synonymous but they
were not synonymyzed until 1913 (Scott and Scott). Guiart (1913),
however, feels that there is some evidence for maintaining both
species. In comparing the two, he states: ‘Cette espéce [C. pelamy-
dis] est beaucoup plus volumineuse que la précédente [C. scombri,
T. Scott’s (1901) modification of Bassett-Smith’s C. scomberi], le
segment génital beancoup (sic) plus long et l’abdomen nettement
forme de deux segments. Beaucoup d’autres caractéres s’opposent
du reste 4 la fusion de cette espéce avec la précédente, comme le
voulait Wilson (8, p. 596).’’ Scott and Scott (1913, p. 59), however,
state: “We think there can be no reasonable doubt that the form
obtained by Krgyer on Pelamys sarda, and described by him in the
work referred to under the name of Caligus pelamydis, is identical
with that from the mackerel [C. scombri]; the general structure of the
animal, the shape of the sternal fork and of the genital segment, and
the structure and armature of the fourth pair of thoracic legs, are
similar in both forms.’”’ A comparison of specimens of these species
in the U.S. National Museum suggests, to the present author, that
the two are synonymous.

Caligus pelamydis is most readily distinguished by the nature of
the fourth thoracic leg. The shape of the female genital segment
is variable. In some of the specimens in the U.S. National Museum
(USNM 74284 and 74358) the lobate condition of the posterior end
of the genital segment is present, as shown in figures 52a,c, while
in others the lobes are reduced or absent.

The fourth thoracic leg is complex, the armature elements are
poorly defined, and, in most cases, lightly sclerotized. The figure
given by Wilson (1905b, pl. 13: fig. 161) does not show the frilled
membranes adjacent to the bases of at least the first 2 spines and does
not show the fine accessory projection at base of the second spine

that was evident in a stained mount of the leg made from a specimen
in the USNM collection.

Caligus kanagurta Pillai, 1961, exhibits a very close similarity to
C. pelamydis. The fourth thoracic leg, however, is figured as having
three small, plumose spines on the distal segment which are not
present in C. pelamydis.

Caligus longipedis Bassett-Simth
Fiacures 54-56
Caligus longipedis Bassett-Smith, 1898b, p. 359, pl. 10, figs. 2-3.—Yamaguti,

1963, p. 56.
Caligus longipes Bassett-Smith, 1899, p. 452.—Wilson, 1905b, p. 556 (in key).

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 145

Caligus amplifurcus Pearse, 1952b, p. 199, figs. 29-35.—Shiino, 1959d, p. 280,
figs. 7-8.—Yamaguti, 1963, p. 49.

Caligus amplifurcatus Pearse, 1952b, p. 200 (figure title) (lapsus).

Caligus lucidus Heegaard, 1962, p. 158, figs. 54-61.

DISTRIBUTION AND HostTs.—4 host records:

locality hosts references
Gulf of Aden Caranz melamphigqus Bassett-Smith, 1898b
Australia Cantherhines ayraud Heegaard, 1962
Eastern Pacific Caranz lugubrius Shiino, 1959
Gulf of Mexico Caranx crysos Pearse, 1952b

MareriAu.—Twenty-six females and 3 males (USNM 112915)
from the external surface of Caranx melampygus Cuvier and Valen-
ciennes, captured in fishtrap by Samuel Kaolulo between Diamond
Head and Koko Head, Oahu, Hawaii. One female (USNM 112916)
from the external surface of C. melampygus Cuvier and Valenciennes,
captured in fishtrap by Samuel Kaolulo between Diamond Head
and Koko Head, Oahu, Hawaii. Four females (one damaged) and
1 male (retained by author) from the external surface of C. melampygus
Cuvier and Valenciennes, captured in fishtrap by Samuel Kaolulo
between Diamond Head and Koko Head, Oahu, Hawaii.

MEASUREMENTS.—(In mm) 28 females and 4 males:

female male
mean range mean range

Total length, excluding caudal setae 5.06 4.58-5.48 5.24 5. 03-5. 48
Length of cephalothorax, including fron-

tal region 3.16 2.89-3.33 3.47 3. 07-3. 74
Width of cephalothorax 2.94 2.63-3.11 3.32 3. 15-3. 52
Length of genital segment 112 0.941.385 0.78 0. 76-0. 83
Width of genital segment 1.29 1.00-1.52 0.83 0. 81-0. 89
Length of abdomen 0.56 0.49-0.67 0.60 0. 56-0. 63
Length of caudal rami 0.29 0. 27-0.33 0.39 0. 38-0. 41
Length of egg strings (10 strings) 3.24 2, 22-3. 96

Descrirtion.—Female cephalothorax (fig. 54b) ovoid, consisting
of cephalon and first 4 thoracic segments. Greatest length of frontal
region approximately one-tenth that of entire cephalothorax, with
fine membrane along anterior margin, bearing pair of ovoid lunules
(fig. 55a) extending posteriorly, on ventral surface, to junction of
frontal region and rest of cephalothorax. Lateral margin of cephalo-
thorax slightly irregular, with narrow membrane extending laterally
and second membrane extending medially, on ventral surface. Pos-
terior-lateral dorsal surface of cephalothorax with small, posteriorly
facing depression. Posterior sinus (fig. 54c) irregularly U-shaped,
with thickened membrane-like projection along outer margin. Pos-
terior median cephalothoracic surface projecting well past lateral
surfaces, posterior end of surface with short, lappet-like projections
laterally. Major dorsal cephalothoracic grooves incomplete, partially

oer aet 6710

146 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

replaced by heavily sclerotized bands forming irregular H. Ocular
region distinct, in anterior third of body.

Female free fourth pedigerous segment distinct from cephalothorax,
incompletely separable from genital segment dorsally and ventrally,

Figure 54.—Caligus longipedis Bassett-Smith, 1898, dorsal view: a, male; b, female; c;
posterior cephalothoracic sinus. Ventral view: d, female genital segment; ¢, male, same,
f, female fifth leg; g, male fifth and sixth legs; h, female caudal ramus; 1, male, same.

overlapped anteriorly by lappet-like projections of posterior median
cephalothoracic surface. Segment widest medially, angled toward
anterior end, convexly curved posteriorly. Genital segment (fig. 54d)
obcordate. Fifth legs (fig. 54f) situated on posterior lateral surface,

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 147

consisting of 2 nodules, anteriormost with single plumose setule,
posterior with 2 plumose setules. Abdomen 1-segmented, broadest
anteriorly, constricted at anterior end, tapered to biconcave posterior
end. Caudal rami (fig. 54h) subrectangular, with slight concavity
on medial inner surface, inner surface plumose. Distal end of rami
with 3 long, plumose setae medially, additional seta on outer distal
surface, plumose setule on inner surface and second on ventral surface
just proximal and medial to outermost seta.

Male cephalothorax (fig. 54a) similar to that of female, free fourth
pedigerous segment slightly shorter, distinct from both cephalothorax
and genital segment, with flat or slightly concave dorsal posterior
margin instead of convex margin of female. Genital segment (fig.
54e) of general barrel shape, lateral surface indented posteriorly,
at origin of fifth legs. Fifth legs (fig. 54g) consisting of nodule
bearing 3 plumose setules; sixth legs (fig. 54g) situated on posterior-
lateral ventral surface, consisting of 2 closely associated plumose
setules. Abdomen 1-segmented although incomplete V-shaped band
of heavy sclerotization in anterior portion giving 2-segmented ap-
pearance. Posterior region of abdomen widest, posterior surface
biconcave, with distinct anal depression. Caudal rami (fig. 54%)
larger than those of female, inner surface convex, plumose in distal
half, remaining armature as in female.

Female and male antennule (fig. 555) 2-segmented, attached to
knoblike extension of ventral-lateral anterior surface of cephalothorax,
not appearing to be attached to frontal region. First segment
slightly more than twice the length of second (male second segment
slightly longer), anterior margin bent almost at right angle medially,
anterior and anterior ventral surface of distal half of segment bearing
approximately 21 lightly plumose setae and setules. Second segment
club shaped, distal end with approximately 12 naked setules. Female
antenna (fig. 55¢) 3-segmented, situated medial and posterior to
antennule base. First segment short, squat, irregular, with spike-
like projection from posterior distal surface. Second segment strongly
developed, broad proximally, tapered to narrower distal end. Third
segment fused with clawlike terminal process, bearing setule-
like accessory process from proximal inner surface, second from
anterior surface. Male antenna (fig. 55d) 3-segmented, first segment
irregular, second irregularly club shaped, with indistinct adhesion
surface on medial inner surface, with distinct swelling just distal
to adhesion surface, swelling bearing well-defined adhesion surface.
Third segment abrupt, terminal process fan shaped, with setule-
like accessory process on distal inner surface.

Female and male mandible (fig. 55c) 4-parted, distalmost part
curved inward, bearing 12 denticulations along inner surface. Female

148 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

postantennal process (fig. 55c) situated lateral to antenna base, con-
sisting of elongate, spine-shaped projection and 3 nodules, 2 on base
of projection, third slightly posterior to base, each with several hair-
like projections. Male postantennal process similar to that of female
except spine-shaped projection longer. Female and male postoral
process (fig. 55c) spine shaped, proximal end broad, medial region
narrow, distal region expanded, flattened. Female and male
maxillule (fig. 55c) consisting of pseudosegmented node bearing 3
setules distally. Female and male maxilla (figs. 55¢, f) 2-segmented,
situated slightly lateral and posterior to postoral process. Second
segment approximately 1 times the length of first, both segments
elongate, second thinner than first, distal end of outer surface knob-
like, distal half of outer surface finely denticulated. Second segment
bearing membranous conical process distally and 2 saber-shaped
terminal processes, innermost slightly less than twice the length of
outermost, with fine membrane along both margins; outermost ter-
minal process with finely frilled membrane along both margins.

Female maxilliped (fig. 55g) 2-segmented, small in comparison with
that of other members of the genus, situated medial and posterior to
maxilla base. First segment well developed although of general elon-
gate nature, tapered proximally to narrow proximal end. Second
segment slender, approximately four-tenths the length of first, dis-
tinct from short, clawlike terminal process, bearing setule-like acces-
sory process from distal inner surface (not figured) and small, hairlike
process from nodule on distal posterior surface. Male maxilliped
(fig. 55h) slightly larger than that of female, first segment more
strongly developed, with 2 ridge-shaped projections on inner surface,
distalmost with small secondary ridge bearing adhesion surface.
Second segment and processes as in female. Female and male ster-
nal furea (fig. 557) situated between and posterior to maxilliped bases.
Furcal base elongate, tines angled outward slightly, with broad sur-
face, bluntly rounded distally.

Inner 2 terminal spines of second segment of exopodite of first
thoracic leg bifid, both parts of equal length. For armature and
nature of thoracic legs, see figure 56 and table 20.

Discussion.—One of the distinguishing characteristics of C. longi-
pedis is the nature of the second segment of the maxilla, with its broad
distal end, denticulated margin, and membranous, spine-shaped
subterminal process. Additionally, the broad tines of the sternal
furca and the long processes on the exopodite of the fourth thoracic
leg are diagnostic characteristics. Although Bassett-Smith’s original
description (1898b) and figures leave much to be desired, these
characteristics, as well as the body shape, are evident. An exam-
ination of the type material of C. amplifurcus Pearse (USNM 93710)

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 149

Ficure 55.—Caligus longipedis Bassett-Smith, 1898, ventral view: a, frontal region, left
side showing lunule and antennule base; d, left antennule; c, female oral region, right side
showing antenna, postantennal process, mouth cone, mandible, maxillule, postoral process,
and maxilla base; d, male right antenna; ¢, left maxilla; f, distal half of second segment
of left maxilla; g, female right maxilliped; h, male, same; 7, sternal furca.

150 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

TaBLE 20.—Armature of thoracic legs I-IV of the female and male of
Caligus longipedis Bassett-Smith, 1898b

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal 3 oe
Plate
1 2 1 2 3 if 2 3
I | Outer $,p Th 3Hi,P’
Inner p ce 38P
II | Outer m,p* m,fm,mH | mH mh,mH, ec jie | ¢,2P
m
Inner s,P m,s Ce cE 5P Poise2P 4P
III | Outer 2s,m,p m,s,H ¢,2s,p’ ¢,3p’ elie
m
Inner 2s,P,s,m,s P 4P Pa WiGe
IV | Outer ss,s,P dm,H 3dm,2H,
mH

*Plumosities not visible but minute fragments of dirt clustered around setule suggest fine plumosities
are present.

showed the presence of these features as well as other characteristics
of C. longipedis. In Pearse’s original description (1952b), however,
the figures do not indicate the relationship. The shape of the genital
segment in the figure of the female, the presence of 2 segments in
the abdomen of the figure of the male, and the ‘‘appendage on basal
segment of second leg’ (Pearse, 1952b, fig. 31) all suggest a species
distinct from C. longipedis. Pearse, however, apparently made his
description and figures from permanent whole mounts. The absence
of spacers between the slide and the coverslip has distorted the shape
of the body and, in one specimen at least, changed the shape of the
appendages. The so-called appendage on the basal segment of the
second leg could not be definitely found although, presumably in the
original mounting of the specimens, the proximal end of one of the
sclerites on the surface of the second segment of the female second
thoracic leg has torn free. This sclerite projects anteriorly and
medially, giving an appearance similar to that shown in Pearse’s
figure 31 and the appearance that it arises from the first segment
of the protopodite.

Although the author has not examined specimens of Caligus lucidus
Heegaard, 1962, the suggested nature of the maxilla (Heegaard’s
“first maxilliped,” no figure number given but identified as MxP,
in figure series 54-61), the nature of the armature on the fourth tho-
racic leg (fig. 58; although the segmentation is not shown), and the
shape of the body of the female all indicate that the species is synony-
mous with C. longipedis.

The description by Shiino, 1959d, of C. amplifurcus Pearse differs
in few details from that given here. The one important difference
is that Shiino indicates the presence of 2 segments in the exopodite
of the third thoracic leg while the present author indicates that
this ramus is 3-segmented. As in C. quadratus, however, Shiino

‘

No. 3574 COPEPOD CRUSTACEANS—LEWIS Pt

H+ a
d o3mm2o 0.5mm.¢o" f

Ficure 56.—Caligus longipedis Bassett-Smith, 1898, right thoracic legs, anterior view: a,
first; b, distal region of second segment of exopodite of first; c, second; d, third; e, exopodite

of third (posterior view); f, fourth.

152 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

indicates that the first segment includes not only the segment bearing
the hook-shaped spine but also the succeeding segment and it is here
suggested (fig. 56¢) that these are 2 distinct segments.

The distribution of C. longipedis is intriguing. Originally described
from near Aden, it has since been described from Australia, the Revilla-
gigedos, the Gulf of Mexico, and now Hawaii. Peculiarly enough,
there has, to the author’s knowledge, never been a specimen described
from either India or Ceylon. Many of the collections of parasitic
copepods made from the latter, however, were from market fish, and
in a good many cases these collections were from the gill cavities and
buccal cavity of the hosts. The only reports giving parasite location
of C. longipedis indicate that it is found on the external surface of the
host. In collections from Hawaiian fishes it was noted that specimens
of this species move rapidly and have a tendency to crawl off the sur-
face of the host when it starts to dry.

C. rugosus Shino, 1959d, exhibits characteristics which closely
ally it with C. longipedis (maxilla, thoracic legs). The exact relation-
ship, however, can not be presently determined.

Caligus kapuhili, new species
Figures 57-59

MarrrrAu.—One female (holotype, USNM 112918) from the gill
cavity of Chaetodon miliaris Quoy and Gaimard, captured in a fish-
trap by Lester Zukeran in Kaneohe Bay, Oahu, Hawaii. One male
(allotype, USNM 112919) from the holotype host specimen. One
female and 1 male (paratypes, USNM 112920) from the gill cavity of
C. miliaris Q. and G., captured in a fishtrap by Lester Zukeran in
Kaneohe Bay, Oahu, Hawaii. Two females and 1 male (paratypes,
USNM 112921) from the gill cavity of an unknown chaetodontid
captured in the Oahu area, Hawaii. One female (paratype, USNM
112922) from the gill cavity of Chaetodon frembli Bennett, speared by
Carolyn Lewis off Rabbit Island, Oahu, Hawaii. One female (para-
type, USNM 112923) from the gill cavity of C. miliaris Q. and G.
from the Honolulu Aquarium. One female (paratype, USNM 112924)
from the gill cavity of C. miliaris Q. and G., captured in a fishtrap
by Lester Zukeran in Kaneohe Bay, Oahu, Hawaii. One female
(paratype, USNM 112925) from the gill cavity of C. miliaris Q. and
G., captured in a fishtrap by Lester Zukeran in Kanohe Bay, Oahu,
Hawaii. Five females and 2 males (one immature) (paratypes,

Figure 57.—Caligus kapuhili, new species, dorsal view: a, male; b, female; c, posterior
cephalothoracic sinus. Ventral view: d, female free fourth pedigerous segment, genital
segment, abdomen, and caudal ramus; ¢, male, same; f, male fifth and sixth legs; g, female
fifth leg; h, caudal ramus

153

005mm.ee
b

0.05 mm. 9
§

COPEPOD CRUSTACEANS—LEWIS

NO. 3574

154 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

USNM 112926) from the gill cavity of several specimens of C. fremblit
Bennett, captured in fishtraps by Samuel Kaolulo between Diamond
Head and Koko Head, Oahu, Hawaii. One female and 1 male (male
damaged) ( paratypes, retained by author) from the gill cavity of C.
fremblii Bennett, speared by Carolyn Lewis off Rabbit Island, Oahu,
Hawaii. Two females (1 damaged) (paratypes, retained by author)
from the gill cavity of C. miliaris Q. and G., captured in a fishtrap
by Lester Zukeran in Kaneohe Bay, Oahu, Hawaii. Four females and
2 males (1 male damaged) (paratypes, retained by author) from the gill
cavity of 4 specimens of C. miliaris Q. and G., captured in fishtraps
by Lester Zukeran in Kaneohe Bay, Oahu, Hawaii.

MEASUREMENTS.—(In mm) 17 females and 5 males:

female male

Total length, excluding caudal mean range mean range

setae 1.94 1.67-2.37 1.52 1. 44-1. 59
Length of cephalothorax, in-

cluding frontal region 1.03 0.92-1.10 0.90 0. 86-0. 92
Width of cephalothorax 0.91 0. 79-1.01 0.78 0. 76-0. 85
Length of genital segment 0.57 0.45-0.72 0.32 0. 32-0. 34
Width of genital segment 0.63 0.50-0.74 0.26 0. 25-0. 27
Length of abdomen 0.19 0. 13-0.23 0.14 0.140.15
Length of caudal rami 0.06 0.05-0.07 0.05 (all)

Length of egg strings (3 strings) 0. 74, 0. 56, 0. 72

Description.—Female cephalothorax (fig. 575) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region distinct,
approximately one-twelfth the total length of cephalothorax, with
fine membrane along anterior margin. Lunules (fig. 58a) large though
finely membranous, extending posteriorly, on ventral surface, almost
to junction of frontal region and remaining cephalothorax. Lateral
margins of cephalothorax smooth, slightly irregular, with slight
indentation in anterior third, with fine membrane extending later-
ally. Posterior sinus (fig. 57c) narrow, irregularly V-shaped, with
finely frilled membrane on outer margin. Posterior median cepha-
lothoracic region projecting well past posterior lateral regions, with
broad, lappet-like projection extending from posterior surface over
anterior end of free fourth pedigerous segment. Major dorsal cepha-
lothoracic grooves forming irregular H, posterior legs and crossbar of
H continuous. Ocular region distinct, in anterior third of cepha-
lothorax.

Female free fourth pedigerous segment distinctly separable from
cephalothorax, indistinctly separable from genital segment; segment
angled sharply toward anterior end from region of fourth leg attach-
ment, in posterior part of segment. Genital segment (fig. 57d)
subtriangular, distended by eggs in figured female, posterior surface
overlapping anterior end of abdomen dorsally. Fifth legs (fig. 579)

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 155

| 4-1 ++}
0.05 mm. Qo

Ficure 58.—Caligus kapuhili, new species, ventral view: a, left antennule and frontal
region showing lunule; b, female oral region, left side showing antenna, postantennal
process, mouth cone, mandible (mdbl), maxillule (ma-1), postoral process, and maxilla
base (ma-2); ¢, male left antenna; d, right maxilla; ¢, left maxilliped; f, sternal furca and
maxilliped bases (mxpd).

156 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

situated on posterior lateral surface of genital segment, consisting of
2 nodules, each in slight depression; anteriormost nodule with single
plumose setule, posterior with 3 plumose setules.

Abdomen 1-segmented, distinct from genital segment, anterior half
slightly swollen, posterior half straight, anal identation in medial
posterior surface formed by pair of small, lappet-like projections.
Caudal rami (fig. 57h) short, rounded distal half of imner margin
plumose, distal surface with 4 plumose setae and 2 plumose setules,
1 on either side of setae.

Male cephalothorax (fig. 57a) similar to that of female. Free
fourth pedigerous segment distinct from cephalothorax, fused with
genital segment, shape as in female. Genital segment (fig. 57e)
narrow anteriorly, broader posteriorly, with slight indentation at fifth
leg and with small, lobate projections from posterior ventral surface.
Fifth legs (fig. 57f) consisting of 2 plumose setules; sixth legs (fig.57/)
arising from 1 of lobate projections on posterior ventral surface of
genital segment, consisting of 3 plumose setules. Abdomen and
caudal rami similar to those of female although anterior end of abdo-
men not swollen as in female.

Female and male antennule (fig. 58a) 2-sezmented, attached to lat-
eral-anterior ventral surface of cephalothorax and adjacent frontal
region. First segment approximately 1% times the length of second,
narrow proximally and distally, broad medially, distal half of anterior
and anterior ventral surface with approximately 21 lightly plumose
setules. Second segment rodlike, with 12 naked setules distally. Fe-
male antenna (fig. 58b) 3-segmented, situated posterior and medial to
antennule base. First segment socket-like, irregular; second segment
broader proximally than distally. Third segment fused with clawlike
terminal process, bearing setule-like accessory process proximally.
Male antenna (fig. 58c) with similar first segment, second segment larg-
er; third segment indistinctly separable from double-clawed terminal
process, with setule-like accessory process at base of proximalmost
claw, at junction of segment and terminal process.

Female and male mandible (fig. 585) 4-parted, distalmost part flat-
tened, with 12 denticulations along inner surface. Female and male
postantennal process (fig. 58b) consisting of small, lappet-like projection
and 3 nodules, 2 at base of projection, third slightly posterior to base,
each with several hairlike processes. Female and male postoral proc-
ess (fig. 58b) elongate, spinelike. Female and male maxillule (fig. 580)
nodular, with 3 setules distally. Female and male maxilla (fig. 58d)
2-segmented, situated lateral to postoral process. Both segments elon-
gate, second more slender, slightly longer than first, with membranous
projection from middle of inner surface and 2 saber-shaped terminal

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 157

Ficure 59.—Caligus kapuhili, new species, right thoracic legs. Posterior view: a, first.
Anterior view: 5, second; c, third; d, exopodite and endopodite of third; ¢, fourth.

158 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

processes. Innermost terminal process approximately 1% times the
length of outermost, both with fine membranes along margins.

Female and male maxilliped (fig. 58¢) 2-segmented, situated pos-
terior and slightly medial to base of maxilla. First segment strongly
developed, with distally concave ledge on inner surface that receives
tip of terminal process of second segment when segment flexed. Sec-
ond segment distinct from clawlike terminal process, with single,
setulelike accessory process on distal inner surface. Female and male
sternal furca (fig. 58f) situated approximately halfway between max-
illiped bases and interpodal plate of first thoracic legs, on small,
heavily sclerotized platelike region projecting laterally as 2 small
lobate processes, adjacent to base of furca. Furcal tines diverging,
sinus irregularly V-shaped, tines broad, flat.

Spine at distal end of first segment of exopodite of third thoracic
leg straight, not curved. For nature and armature of legs, see figure
59 and table 21.

TaBLE 21.—Armature of thoracic legs I-IV of the female and male of
Caligus kapuhili, new species

Inter- Protopodite Exopodite Endopodite
Megs MS mEtace's|||( OC al |: 2 ca. erate tee S| a ee a ee ek Pee
Plate
2 1 2 3 1 2 3
I | Outer p h h,mH,2H rh?
Inner Dp c 3P rh?
II | Outer m,p | m,H H 2h,Q Cc C, 3P
m
Inner P,s MSs |MCye: ce 5P 12) | ia 2 3P
III | Outer a,m,P 2ss,H ¢,p’ ¢,3p’ c | ¢,3P
m
Inner 3s,P,m,s P 4P Paes
IV*| Outer 3s,p mH mh 3mH,m |

*Division between second and third segments of exopodite indistinct, incomplete.

Discusston.—Caligus kapuhili exhibits several characteristics closely
approximating those of C. laticaudus Shiino, 1960a. The similarity
of the lunule, the female and male antenna (male C. laticaudus de-
cribed by Pillai, 1961), the maxilliped with its cup-shaped projection
on the first segment, and the similarity of the first 3 thoracic legs
suggest that these 2 species are closely related. The differences
between the species are in the body shape, the size relationship of
the body parts, and the characteristics of the appendages other than
those mentioned above. Shiino’s female and only specimen of C.
laticaudus is reportedly 3.51 mm in length. Pillai reports a length
of 2.6 mm for his female specimens (no length given for the male),
the average length of the Hawaiian female specimens is 1.94mm,
and the range is from 1.67-2.37mm. The abdomen is distinctly
2-segmented in C. laticaudus and 1-segmented in C. kapuhili, while
its length (female only) is 0.71mm in C. laticaudus and an average

|
:

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 159

of 0.19 mm (range 0.13-0.23 mm) in C. kapuhili. The relationship
of the length of the abdomen to the length of the body (average
total length divided by average length of abdomen) is 4.94 in C.
laticaudus and 10.2 in C. kapuhili. The spinelike projection of
the postantennal process in C. laticaudus is elongate and distally
pointed, while in C. kapuhili it is short and distally rounded. The
fourth thoracic leg of C. laticaudus has a frilled membrane at the
base of each spine; with the exception of the innermost terminal
spine, these membranes are absent in C. kapuhilt.

The species name is derived from ‘‘Kapuhili,”’ the Hawaiian name

for several of the chaetodontids or butterfly fishes, the host of the

species.
Caligus kala Lewis

Fiaures 60, 61

Caligus kala Lewis, 1964a, p. 142, figs. 2-3

DIsTRIBUTION AND Hosts.— Hawaiian Islands, Naso hexacanthus.

Mareriau.—One female and 1 male (USNM 112927) from the
buccal cavity of Dascyllus albisella Gill, captured by spear by
Robert Stevenson, Oahu, Hawaii. One male (USNM 112928) from
the buccal cavity of D. albisella Gill from the Honolulu Aquarium.
One immature male (USNM 112929) from the gill cavity of Saurida
gracilis (Quoy and Gaimard), captured in a fishtrap by Samuel
Kaolulo between Diamond Head and Koko Head, Oahu, Hawaii.
Two males (USNM 112930) from the gill cavity of D. albisella Gill,
captured by rotenone off Waikiki, Oahu by William Gosline. One
female (USNM 112931) from the gill cavity of D. albisella Gill, from
the Honolulu Aquarium. One immature male (USNM 112932) from
the buccal cavity of D. albisella Gill, from the Honolulu Aquarium.
One female and 1 male (retained by author) from the buccal cavity
of Pomacentrus jenkinsi (Jordan and Evermann), speared by N.
Ferris in Hanauma Bay, Oahu, Hawaii. One damaged female (re-
tained by author) from the gill cavity of D. albisella Gill, from the
Honolulu Aquarium.

MEASUREMENTS.— (In mm) 5 females and 7 males:

female male
mean range mean range

Total length, excluding caudal setae 4.20 3.95-4.40 2.79 2. 20-3. 20
Length of cephalothorax, including

frontal region 2.27 2.10-2.43 1.87 1. 70-2.05
Width of cephalothorax 1.85 1.75-2.00 1.50 1.30-1. 63
Length of genital segment 1.27 1.18-1.38 0.53 0. 50-0. 60
Width of genital segment 1.18 1.00-1.45 0.63 0. 58-0. 78
Length of abdomen 0.68 0.61-0.77 0.31 0. 25-0. 37
Length of caudal rami 0.14 0.13-0.15 0.12 0. 10-0. 14

Length of egg strings (2 strings) 1. 30, 1. 48

160 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

DescripTion.—Kemale and male cephalothorax (figs. 60a,b) ellip-
tical, consisting of cephalon and first 4 thoracic segments. Frontal
region less than one-ninth the length of cephalothorax, with fine
membrane along anterior margin. Lunules large, extending poste-
riorly, on ventral surface, slightly past division between frontal region
and rest of cephalothorax. Posterior margin of median cephalotho-
racic region trilobed, not convex as originally described, median lobe
projecting over anterior end of free fourth pedigerous segment. For
description of rest of cephalothorax and fourth pedigerous segment,
see original description.

Female genital segment (fig. 60d) of general obcordate shape al-
though anterior end broadly rounded. Posterior lobes projecting to
middle of abdomen, tipped by fifth legs. Fifth legs (fig. 60f) consist-
ing of 2 knobs, outermost with single plumose setule, inner with 2
plumose setules. Female abdomen indistinctly 2-segmented, broad,
lateral margins irregularly convex. First “segment” shorter than
second, second with flatly angled distal end, with deep anal depression.
Caudal rami of female and male (fig. 602) as origionally described
except for additional small setule present on outer distal ventral
surface and hairlike process from middorsal surface.

Male genital segment (fig. 60e) of varible shape, variation appear-
ing due to longitudinal contraction of free body segments. Dis-
tinct separation of fourth pedigerous and genital segments in holotype
specimen, separation not visible in presently figured specimen but
present in specimens exhibiting contracted condition. ‘Posterior
margin” of original description (p. 144) thus formed by cuticular
fold. Sixth leg (fig. 60g) present as small, naked setule and small
plumose setule, both situated immediately ventral to junction of
abdomen and genital segment.

In the description of the appendages and processes given below,
those of the heretofore undescribed female are given only if they
differ from the male. Additional descriptions are given if there
appeared to be some error in the original description (Lewis, 1964a).

Female and male antennule (fig. 61a) as originally described except
attachment to both frontal region and anterior-lateral ventral surface
of cephalothorax, not just to frontal region. Female antenna (fig. 61d)
3-segmented, situated posterior and medial to antennule base. First
segment short, squat, with posteriorly directed lobate projection;
second segment broader proximally than distally, well developed.
Third segment indistinctly separable from clawlike terminal process,
with setule-like accessory process from proximal inner surface, second
from inner distal surface. Male antenna (fig. 61a) as originally de-
scribed except only one accessory process, on anterior inner surface.
Female postantennal process (fig. 615) situated lateral to antenna base,

NO: 3574 COPEPOD CRUSTACEANS—LEWIS 161

Ficure 60.—Caligus kala Lewis, 1964, dorsal view: a, female; 6, male; c, posterior cepha-
lothoracic sinus. Ventral view: d, female genital segment; ¢, male, same; f, female right
fifth leg; g, male right fifth (V) and sixth (VI) legs; h, caudal ramus.

226-321— 67——11

162 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

consisting of short, slightly curved, spinelike process, with 2 nodes
proximally, third node present slightly posterior to spinelike process,
all 3 nodes with several hairlike processes. Male postantennal

Sas eee
YS == 0.05mm8c"

Ficure 61.—Caligus kala Lewis, 1964, ventral view: a, male oral region, left side showing
antenna, postantennal process, mouth cone, maxillule, postoral process, and maxilla base;
b, female right antenna and postantennal process; ¢, sternal furca. Right thoracic legs:
d, distal end of second segment of exopodite of first, anterior view; e, exopodite of third,
posterior view.

process (fig. 61a) similar although spinelike process much longer than
in female. Female and male maxillule (fig. 61a;=part of postoral
process in original description) nodular, with 1 large and 2 small,

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 163

setules. Female and male maxilla as originally described except lo-
bate process on middle of second segment a folded membrane. Female
maxilliped similar to that of male except with single tuberculate
projection from middle of inner margin of first segment, not with 2
as in male.

Female and male sternal furca (fig. 61c) attached to ventral surface
of cephalothorax between pair of small, lobate, anteriorly directed
projections. Additional pair of somewhat larger lobate projections
present between interpodal plates of first and second thoracic legs.
First thoracic leg with distinct break in plumosities along inner sur-
face of first segment of exopodite; both inner spines on distal surface of
second segment of exopodite with flexible accessory processes, not
just middle spine as originally indicated. Exopodite of third thoracic
legs 3-segmented, not 2 as originally noted, protopodite with plumose
setule at base of exopodite. Endopodite of third legs includes lobate
projection previously indicated as arising between rami. For nature
of armature and thoracic legs, see figures 61d, e and table 22 in present
publication, figures 3a-f in Lewis, 1964a.

TABLE 22.—Armature of thoracic legs I-IV of the female and male of
Caligus kala Lewis, 1964a

Inter- Protopodite Exopodite Endopodite
Leg | Surface podal are
plate
1 2 1 2 3 1 2 3
I | Outer sss,P rh 3H,P
Inner d,p c,¢ 3P
II | Outer m,p|m,m,mH |} mH h,H,Q,P CCD 3P
m
Inner Se m,s | ¢,P c,P c,4P 2 iP c,3P
III | Outer d,m,p H c,p’ c,3p’,P e C,3P
m
Inner d,2s,P,s, e;P ¢,3P iE ¢,3P
m,s
IV | Outer 2s,p | s,m,mH 4m ,4mH
\

Caligus ligatus Lewis
FicureEs 62-64
Caligus ligatus Lewis, 1964a, p. 164, figs. 8-9.

DIsTRIBUTION AND HoSTS.—Hawaiian Islands, Acanthurus dussu-
meri, Naso hexacanthus.

MaATERIAL.—Five females and 2 males (USNM 112933) from the
gill cavity and buccal cavity of Dascyllus albisella Gill, from the
Honolulu Aquarium. One male (retained by author) from the buccal
cavity of Aulostomus chinensis (Linnaeus), captured in fishtrap by
Samuel Kaolulo between Diamond Head and Koko Head, Oahu,
Hawaii. One male (retained by author) from the gill cavity of

164 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Holocentrus zantherythrus Jordan and Evermann, from the Honolulu
Aquarium. One immature female and 5 males (retained by author)
found in bait well after loading Pranesus insularum (Jordan and
Evermann) at French Frigate Shoals (USFWS, JRM 31).

MEASUREMENTS.—(In mm) 5 females and 3 males:

female male
mean range
Total length, excluding caudal setae 3.29 3. 20-3.35 2. 25, 2. 55, 2. 65

Length of cephalothorax, including

frontal region 1.62 1.55-1.68 1. 40, 1. 58, 1. 60
Width of cephalothorax 1.49 1.43-1.53 1.18, 1.30, 1. 33
Length of genital segment 0.95 0.93-0.98 0. 43, 0. 50, 0. 53
Width of genital segment 0.93 0.88-1.03 0. 38, 0. 40, 0. 43
Length of abdomen 0.59 0. 58-0.61 0. 29, 0. 48, 0. 34
Length of caudal rami 0.13 (all) 0. 12, 0. 15, 0. 14
Length of egg strings (2 strings) 1.55, 0:73

Description.— Female cephalothorax (fig. 62a) ovoid, consisting
of cephalon and first 4 thoracic segments. Frontal region approxi-
mately 15 percent of total length of cephalothorax, with broad mem-
brane along anterior margin and pair of lunules (fig. 63a) extending
posteriorly, on ventral surface, slightly past junction of frontal region
and rest of cephalothorax. Lateral margin of cephalothorax slightly
irregular, with fine membrane extending laterally and second mem-
brane extending medially, on ventral surface; posterior lateral surface
with small but distinct concavity. Posterior sinus (fig. 62c) irregularly
U-shaped, with fine membrane along outer surface. Posterior me-
dian cephalothoracic region flatly convex, extending posteriorly well
past lateral cephalothoracic regions, connected to free fourth pedig-
erous segment by flaccid arthrodial membrane. Major dorsal ceph-
alothoracic grooves forming irregular H although anterior longitudinal
grooves of H extending posteriorly and medially, past cross groove
(more pronounced in female than in male). Ocular region distinct, in
anterior third of cephalothorax.

Female free fourth pedigerous segment narrow anteriorly, lateral
margins angled outward sharply to region of fourth leg attachment.
Segment indistinctly separable from genital segment. Genital seg-
ment (fig. 62d) apple shaped, broader posteriorly, with fifth legs from
ventral-posterior lateral surface. Fifth legs (fig. 62f) consisting of 2
knobs, anteriormost with 1 plumose setule, posteriormost with 2.

Female abdomen indistinctly separable from genital segment, con-
sisting of 1 or 2segments. Indication of segment division indistinctly
present in middle of abdomen. Abdomen subrectangular, width ap-
proximately half the length, tapered slightly toward posterior end.
Posterior end angled sharply at attachment of caudal rami, anal region
bilobed. Caudal rami (fig. 624) subrectangular, with small indenta-

{
NO. 3574 COPEPOD CRUSTACEANS—LEWIS 165

Three plumose setae present on distal surface; single plumose setule

tion on medial outer surface, indentation bearing single plumose seta.
on inner distal surface, second on outer surface, just medial to indenta-
|

0.05mm8c" /h

Figure 62.—Caligus ligatus Lewis, 1964, dorsal view: a, female; b, male; c, posterior ceph-
alothoracic simus. Ventral view: d, female genital segment; e¢, male, same; f, female left
fifth leg; g, male right fifth (V) and sixth (VJ) legs; h, caudal ramus.

tion; distal half of inner surface plumose.
Male cephalothoax (fig. 626) similar to that of female, free fourth
pedigerous segment also similar except for irregular posterior lateral

166 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

margins. Genital segment (fig. 62¢) subovoid, fifth legs (fig. 629)
arising from ventral lateral surface, consisting of 2 knobs, each with
single, plumose setule; sixth legs (fig. 62g) arising from slight swell-
ing on lateral-posterior ventral surface, consisting of minute nodule
bearing 3 plumose setules. Abdomen distinctly 2-segmented, first
segment approximately one-third the total length, constricted sharply
posteriorly; second segment slightly wider posteriorly than anteri-
orly. Caudal rami as in female.

Female and male antennule (fig. 635) 2-segmented, attached to
anterior ventral surface of cephalothorax, not attached to adjacent
frontal region. First segment slightly less than 1% times the length
of second, narrow proximally and distally, broad medially; distal
half of anterior and anterior ventral surface with approximately
23 plumose setae and setules. Second segment club shaped, with
single naked setule from distal half of posterior surface, approximately
11 naked setules from distal surface. Female antenna (fig. 63c)
3-segmented, situated posterior and medial to antennule base. First
segment dactyliform, ventrally facing surface irregular; second seg-
ment broader proximally than distally. Third segment short, incom-
pletely fused to clawlike terminal process, with single setule-like
accessory process from posterior medial surface. Male antenna (fig.
63d) 3-segmented, first segment elongate, irregular; second segment
broader proximally than distally, with adhesion surface on proximal
half of anterior surface and small, shelflike adhesion surface project-
ing from distal half. Third segment short, incompletely fused with
short, bifureate terminal process, tines of bifurcation sharply
pointed. Single, setule-like accessory process present on distal
anterior surface of segment.

Female and male mandible (fig. 63c) 4-parted, distalmost part
curved medially, with 12 denticulations. Postantennal process of
female (fig. 63c) situated lateral and slightly posterior to antenna
base, consisting of short, clawlike projection with 3 associated nodules,
2 on base of projection, third just posterior to base, each with several
hairlike processes. Male postantennal process (fig. 63d) consisting of
long, strongly curved, clawlike projection with 2 nodules proximally,
each with several hairlike processes, third nodule not visible. Female
and male postoral process (fig. 63c) with broad base, tapered abruptly
to slender, slightly curved spinelike projection. Female and male
maxillule (fig. 63¢) nodular, with 2 small and 1 large setule. Female
and male maxilla (fig. 63e) 2-segmented, situated lateral and slightly
posterior to postoral process. First segment slightly more than
four-fifths the length of second, narrower proximally and distally
than medially. Second segment elongate, with small, lobate mem-
brane on distal half of posterior surface and 2 saber-shaped terminal

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 167

Het 4 pe
O.1mm&o" Olmm2o

Ficure 63.—Caligus ligatus Lewis, 1964, ventral view: a, frontal region, left side showing
lunule and antennule base; J, left antennule; c, female oral region, right side showing
antenna, postantennal process, mouth cone, mandible, maxillule, postoral process, and
maxilla base; d, male left antenna and postantennal process; ¢, left maxilla; f, female right
maxilliped; g, male left maxilliped; h, maxilliped bases (mxpd), sternal furca, interpodal
plate of first thoracic leg (ip), and processes between interpodal plates of first and second
thoracic legs.

168 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

a

bret
Oimm.?c

Ficure 64.—Caligus ligatus Lewis, 1964, right thoracic legs, anterior view: a, first; }, distal
region of second segment of exopodite of first; c, second; d, third; ¢, exopodite of third;
f, fourth.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 169

processes. Outermost terminal process approximately 1) times the
length of innermost, with fine membrane along both margins; inner-
most with finely serrated membrane along both margins.

Male maxilliped (fig. 63g) 2-segmented, situated posterior and
medial to maxilla base. First segment large, strongly developed,
with knoblike projection on distal half of inner surface, distal surface
of knob grooved, appearing to receive distal part of terminal process of
second segment when segment flexed. Second segment short, tapered
from proximal end to indistinct junction with clawlike terminal
process, single, setule-like accessory process present on distal inner
surface of segment. First segment of female maxilliped (fig. 63f/)
not as well developed as that of male, without projection of distal
inner surface. Second segment and terminal process as in male.
Sternal furca situated on median longitudinal axis of body, posterior
to maxilliped bases, base of furca long, lobate, tines diverging in
curved manner from point of bifurcation, terminating in rounded tips.

For nature of legs and armature, see figure 64 and table 23.

Remarxs.—The presence of mature females and the variation
exhibited in the males in the present collection indicated that a re-
description of the species was in order.

TaBLE 23.—Armature of thoracic legs I-IV of the female and male of
Caligus ligatus Lewis, 1964a

Inter- Protopodite Exopodite Endopodite
Leg | Surface || podal Je ae
Plate
1 2 1 2 3 1 2 3
I | Outer ss,p rh 3HEP
Inner p c 3P
It | Outer m,p | m,fm,mH | dmH h,mH,Q,P} ¢ Ce ¢,3P
m
Inner s,P eS Cake Ge 4P 1 PAR ae
Til | Outer m,d,p s,H ¢,S,p’ ¢,3p’ ChalnCy ak
m
Inner s,P,s,m,s cae c,4P P | ¢3P
IV | Outer p sfm,mH | mH,fm,mH,
fm,H,fm,H

* Condition on male, female=D.
Caligus flexispina Lewis

Caligus flexispina Lewis, 1964a, p. 149, figs. 4-5.

DISTRIBUTION AND HOSTs.—Hawaiian Islands, Acanthurus trio-
stegus sandvicensis.

Mareriau.—One female (retained by author) from the external sur-
face of Aulostomus chinensis (Linnaeus), captured in a fishtrap by
Samuel Kaolulo between Diamond Head, and Koko Head, Oahu, Ha-
waii. One male (retained by author) from the external surface of an
unknown labrid captured in the Hawaiian region.

170 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

MEASUREMENTS.—(In mm) 1 female and 1 male:

JSemale male
Total length, excluding caudal setae 2. 37 2. 29
Length of cephalothorax, including frontal region 1. 53 1. 67
Width of cephalothorax eZ 1. 26
Length of genital segment 0. 59 0. 38
Width of genital segment 0. 68 0. 38
Length of abdomen 0. 20 0. 20
Length of caudal rami 0. 20

Length of egg strings (female nonovigerous)

DescripTion.—See Lewis, 1964a.

Caligus randalli Lewis

Caligus randalli Lewis, 1964a, p. 156, figs. 6a, b, d-f, h-l, n-q; fig. 7.
DIsTRIBUTION AND HostTs.—Hawaiian Islands, Acanthurus trio-
stegus sandvicensis.
DerscriIpTION.—See Lewis, 1964a.

Caligus kalumai Lewis

Caligus kalumai Lewis, 1964a, p. 171, fig. 10.

DisTRIBUTION AND HosTs.—Hawaiian Islands, Acanthurus guttatus.
DeEscriIPTION.—See Lewis, 1964a.

Order Lernaeopodoida

Family Lernaeopodidae

Charopinopsis Yamaguti, 1963

Diacnosis.—Female: Cephalothorax, including posteriorly _re-
moved maxilla-bearing segment, in line with rest of body, anterior
portion of tergum heavily sclerotized. Trunk broader than cephalo-
thorax, flattened dorsoventrally, without indication of segmentation,
with subconical or conical projection at each posterolateral corner;
with pair of long, dactyliform posterior processes medially, ventral to
oviducal openings. Abdomen and caudal rami not visible. Anten-
nule 3-segmented, subconical; antenna biramous, not chelate, ex-
opodite broadly rounded, without externally projecting armature,
endopodite 2-3 segmented (status of distalmost “segment’’ ques-
tionable). Maxillule with small palp, palp tipped by setule (spine?),
distal surface of maxillule with 3 setules (spines?). Maxillae posterior
to maxillipeds, joined at bulla, distal region convoluted, giving
knoblike appearance. Maxillipeds 2-segmented, prehensile, second
segment with short, clawlike terminal process.

Male: See Yamaguti, 1963, p. 251.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS U7
Charopinopsis quaternia (Wilson)

FIGURE 65

Charopinus quaternius Wilson, 1935b, p. 343, pl. 4, figs. 42-49.—Causey, 1953a,
Deletios lo:

Brachiella coryphaenae Pearse, 1952a, p. 35, figs. 129-135.—Pillai, 1962a, p. 85,
fig. 18 —Yamaguti, 1963, p. 248, pl. 270, fig. 7.

Charopinopsis quaternia (Wilson) Yamaguti, 1963, p. 251, pl. 271, fig. 3.

DISTRIBUTION AND HOSTS.—4 host records:

locality hosts references
Gulf of Mexico Peristedion gracilis
Coryphaena hippurus Wilson, 1935b
Scomberomorus cavalla Causey, 1953a
Indian Ocean Coryphaena hippurus Pillai, 1962

MateriAu.—Nine females (USNM 112934) from the gill filaments
of Coryphaena hippurus Linnaeus, captured 120 miles south of Oahu,
Hawaii (USFWS, HMS cruise 34).

MEASUREMENTS.—(In mm) 9 females:

mean range

Total length, excluding posterior processes 6. 75 6. 23-7. 35
Length of cephalothorax 1. 54 1. 37-1. 70
Width of anterior region of cephalothorax 0. 78 0. 74-0. 85
Width of posterior region of cephalothorax (at level of

maxillae) 0. 54 0. 47-0. 63
Length of “neck” between maxillae and trunk 0. 95 0. 81-1. 11
Length of trunk, excluding posterior processes 4, 38 4, 138-4. 65
Width of trunk 1. 75 1. 30-2. 00
Length of lateral-dorsal posterior processes 0. 66 0. 44-0. 85
Length of median-ventral posterior processes 2. 91 2. 48-3. 55
Length of egg strings (8 strings) 7. 54 6—53-8. 18

DESCRIPTION OF FEMALE.—Cephalothorax (fig. 65a) elongate, ante-
rior part bearing small, terminally concave projection from median
anterior surface, in addition to cephalic appendages (except maxillae)
and maxillipeds. Posterior part of cephalothorax necklike, slightly
narrower than anterior part. Trunk elongate, broader than cephalo-
thorax, constricted at junction with cephalothorax, without indication
of segmentation. Posterior end of trunk with 2 pairs of processes (fig.
650), 1 short, dactyliform pair that may be curved on outer posterior
surfaces; 1 long, filamentous pair on ventral posterior surface, adjacent
to oviducal openings. Median posterior ventral surface of trunk with
small, padlike projection appearing to contain droplets of viscous ma-
terial, projection not appearing to be associated with reproductive
elements. Median posterior surface essentially concave although
heavily sclerotized, forming place of attachment for ovoid spermato-
phores that project slightly and give biconcave outline to surface.

Li2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Antennule (fig. 65c) 3-segmented, attached to concave frontal
projection, immediately anterior and slightly lateral to labrum.
Appendage flaccid, first segment broad proximally, approximately
half as wide distally, with flexible, subconical projection on inner
surface. Second segment short, approximately two-thirds the
length of third segment; third segment rounded distally, distal sur-
face with 2 setules. Antennae (fig. 65d) large, biramous, attached
lateral and slightly posterior to antennule. Protopodite subrec-
tangular, with irregular areas of heavy sclerotization. Exopodite
separable from distal surface of protopodite by breaks in scle-
rotization, with 2 minute, knoblike projections on inner part of
rounded distal surface; entire ramus, including projections, covered
by filmy cuticle. Endopodite distinctly separable from distal inner
surface of protopodite, apparently 3-segmented although status of
terminal “‘segment”’ questionable. First and second segments sub-
cylindrical, first slightly less than twice the length of second. Third
“seoment” approximately two-thirds the length of second, concave
distally, with minute, setule-like projection arising from concavity.

Mandible (fig. 65d) rodlike, distal end flattened, inner edge rounded,
without apparent denticulations. Maxillule (fig. 65¢) situated
adjacent to posterior lateral edge of mouth cone base, consisting
of stalk with palp on distal half of inner surface, palp tipped by spine-
like projection. Distal end of maxillule rounded, with single short,
setalike process on outer edge, 2 long, setalike processes on medial
and inner edge. Maxillae (fig. 65a) subcylindrical, flabby, distal
region (fig. 65f) tapered to bulbous swelling formed by convolution
of distal end of each maxilla; swellings enclosing tip of capstan-
shaped bulla.

Maxilliped (fig. 65g) 2-segmented, situated posterior and medial
to maxillule base, arising from small, apron-shaped projection of
ventral surface of cephalothorax. First segment well developed,
approximately twice the length of second, medial inner surface with
knob-shaped swelling bearing small, conical projection. Second
segment with wavy outline, distal third of inner surface denticulated,
inner distal surface bearing setalike process. Terminal process of
second segment originating on outer distal surface of segment, claw-
like except for spike-shaped accessory projection on medial inner
surface.

Discussion.—The Hawaiian specimens differ from Wilson’s type-
material (USNM 64009-64011) and Pearse’s holotype and paratype
slides (USNM 92663, 92688) in size, the Gulf of Mexico specimens
being approximately four-fifths the length of the Hawaiian specimens
(excluding posterior processes). There is also some minor variation
in the size and shape of the various body regions and posterior proc-

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 173

Ficure 65.—Charopinopsis quaternia (Wilson, 1935), female: a, dorsal view (eggs drawn
in free hand). Ventral view: b, posterior end of trunk showing posterior processes,
region of genital openings, and part of one egg string; c, right antennule; d, anterior end
of cephalothorax, left side showing antennule (a-1), antenna (a-2), mouth cone, mandi-
ble, maxillule (ma-1), and maxilliped; ¢, left maxillule; f, distal end of maxilla showing
attachment with host tissue (h); g, right maxilliped.

174 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

esses, both in the 3 collections as a whole and individually, but
the author attributes this to natural variation. Pillai (1962a, p. 86)
points out that the posterior processes on the trunk of his specimens
‘“‘are more narrowing towards the tip”’ than those described by Pearse
(1952a). Pearse, however, made permanent mounts of his specimens
and flattening of the specimens and their component parts has taken
place, as evidenced by his figures.

Brachiella Cuvier, 1829

Diaagnosis.—Female: Cephalothorax elongate, frequently at angle
to trunk, anterior region of tergum heavily sclerotized. Trunk
swollen, flattened dorsoventrally, with 1 or 2 pairs of posterior proc-
esses and single genital process (1 or more of these may be reduced
or lacking); without recogmizable abdomen or caudal rami. Anten-
nule 2-4 segmented, subconical; antennae biramous, exopodite
with or without armature, endopodite reduced. Maxillule with
palp; maxillae removed well behind maxillipeds, distally united,
with bulla. Maxillipeds close to oral region, 2-segmented, second
segment with clawlike terminal process.

Male: Body divisible into prosome (~cephalothorax) and urosome,
separated by constriction or internal indication of constriction.
Urosome longer than prosome, usually narrower, caudal rami small.
Antennule 3—4 segmented, antennae biramous, exopodite 1-segmented,
lobate, endopodite 2-segmented. Maxillule similar to that of female;
maxilla 2-segmented, tipped with clawlike terminal process. Max-
illiped 2-segmented, second segment with clawlike terminal process.

Brachiella thynni Cuvier
Fiacures 66, 67

B. thynni Cuvier, 1829, p. 257, pl. 15, fig. 5—Guérin-Méneville, 1829-1844, pl.
9, fig. 6a-c.—Nordmann, 1832, p. 90.—Milne-Edwards, 1840, p. 512.—
Steenstrup and Liitken, 1861, p. 420, pl. 15, fig. 36—Van Beneden, 1851,
p. 128; 1861, p. 153.—Heller, 1866, p. 756.—Van Beneden, 1870a, p. 37;
1870b, p. 244.—Vogt, 1877, pl. 3, fig. 9—Richiardi, 1880, p. 7.—Stossich,
1880, p. 268.—Valle, 1880, p. 77.—Carus, 1885, p. 375. Bassett-Smith,
1896a, p. 162.—Brian, 1899a, p. 6.—Bassett-Smith, 1899, p. 502.—Brian,
1901, p. 1, fig. 1—Graeffe, 1902, p. 16—Thompson and Scott, 1903,
p. 294.—Stenta, 1904, p. 345.—Miculicich, 1905a, p. 600; 1905b, p. 733.—
Rathbun, 1905, p. 102.—Brian, 1905, p. 8; 1906, p. 105, pl. 9, fig. 1—Scott
and Scott, 1913, p. 204, pl. 64, figs. 4-6——Wilson, 1915, p. 703, pl. 25, fig. C;
pl. 53, figs. 209-215.—Leigh-Sharpe, 1926, p. 386.—Kirtisinghe, 1935,
p. 342, figs. 40-42.—Bere, 1936, p. 613.—Bonnet, 1948, p. 7—Causey,
19538b, p. 15.—Delamare-Deboutteville and Nunes-Ruivo, 1953, p. 217.—
Shiino, 1956b, p. 2838, figs. 8-9; 1958, p. 112; 1960b, p. 539.—Pillai, 1962a,
p. 81, figs. 15, 16.—Shiino, 1963a, p. 346—Yamaguti, 1963, p. 247, fig.1.—
Kirtisinghe, 1964, p. 119, figs. 171, 172.

Thynnicola ziegleri Miculicich, 1904, p. 48, figs. 1-3.

NO. 3574

DIsTRIBUTION AND HostTs.— 28 host records:

locality
Unknown

North Atlantic

Equatorial Atlantic

Baltic
Gulf of Mexico

Hawaii

Japan
Indian Ocean

Mediterranean

COPEPOD CRUSTACEANS—LEWIS

hosts
Scomber thynnus
“tuna”
Thynnus thynnus
T. vulgaris
“tuna’’
Orcynnus thynnus
“Albacore”
Sciaena aquila
“‘Baracottaer”’

Thynnus vulgaris
Pomatomus saltatrix
Scomberomorus cavalla
Thynnus species
Thunnus albacares
Parathunnus sibt
Thunnus obesus
Acanthocybium solandri
Chirocentrus dorab

Thunnus (Germo) macropterus
Parathunnus obesus
Neothunnus albacora
Acanthocybium solandri
Indocybium lineolatum
Neothynnus macropterus
Thynnus vulgaris

Thynnus thynnus

‘funa’”’

Thunnus thynnus

175

references
Nordmann, 1832
Milne-Edwards, 1840
Bassett-Smith, 1896a
Rathbun, 1905
Wilson, 1915
Scott and Scott, 1913

Steenstrup and Liitken,
1861

Van Beneden, 1870a

Bere, 1936

Causey, 1953b

Bonnet, 1948

Shiino, 1963a

Shiino, 1956b

Thompson and Scott,
1903

Kirtisinghe, 1935

Shiino, 1958

Shiino, 1960b

Pillai, 1962a

Kirtisinghe, 1964

Carus, 1885

Brian, 1899a

Brian, 1901

Delamare-Deboutteville
and Nunes-Ruivo,
1953

Mareriau.— Three females and 2 males (USNM 112935) from the
external surface of Acanthocybium solandri (Cuvier) caught in the
Hawaiian region (USFWS, HMS cruise 38).

MeAsuUREMENTS.— (In mm) 3 females and 1 male:

Total length, excluding posterior processes
Length of cephalothorax, including ‘‘neck”’
Width of anterior region of cephalothorax

Width of posterior region of cephalothorax (at maxillae)

Length of trunk
Width of trunk

Length of dorsal posterior processes
Length of ventral posterior processes

Length of egg strings

female

9.30, 14.55, 16.13
5.85, 11.25, 10.50

0.93, 1.07, 0.96
1.30, 1.26, 1.30
3.45, 3.30, 5.63
2.29, 3.48, 3.77
5.93, 9.15, 9.15
3.90, 8.18, 9.15

6.23, 13.05, 15.90

176 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

male
Total length 1. 85
Length of prosome 0. 76
Width of prosome 0. 53
Length of urosome 0. 91
Width of urosome 0. 53
Width at constriction between prosome and ursome 0. 24
Length of caudal rami 0. 12

DESCRIPTION OF FEMALE.—Body (fig. 66a) consisting of 2 parts,
anteriormost (cephalothorax) elongate, vermiform, posteriormost
(trunk) flattened dorsoventrally, swollen laterally. Cephalothorax,
except for maxilla-bearing segment, with heavily sclerotized tergum;
maxillae arising from posterior end of cephalothorax, well posterior
to maxillipeds. Swollen trunk slightly narrower anteriorly than
posteriorly, lateral margins irregular. Dorsal surface of trunk flatly
convex, with slight irregularities, distinctly separable from vermi-
form cephalothorax, without distinct evidence of segmentation.
Ventral surface of trunk (fig. 66b) with 3 pairs of bosses, with dis-
tinct though incomplete line of division posterior to each pair, giving
4-segmented appearance. Lateral posterior surfaces projecting past
slightly biconvex median portion as lobate extensions; median pos-
terior surface bearing 4 elongate, lanceolate projections, 2 dorsally, 2
ventrally, each member of dorsal pair with knob-shaped proximal swell-
ing. Trunk with small, knoblike projection between and ventral to
base of processes. Egg strings projecting from posterior surface,
between dorsal and ventral posterior processes.

Antennule (fig. 66¢c) 3-segmented, club shaped, arising from anterior
ventral surface of cephalothorax, just lateral and anterior to mouth
cone. First segment swollen, flabby, slightly less than twice the
combined lengths of remaining 2 segments. Second segment cylin-
drical; third segment approximately 1% times the length of second,
rounded distally, with single, minute spinule from medial anterior
surface and 2 hairlike processes distally. Antennae (fig. 66¢e) bira-
mous, situated posterior and lateral to antennule base, extending
anteriorly around anterior end of cephalothorax. Division between
antenna base and cephalothorax indistinct, incomplete. Protopodite
and exopodite separable only by breaks in sclerotization, forming
dactyliform projection. Endopodite rudimentary, knoblike, with 3
spinules distally.

Mandible (fig. 66f) rodlike, wavy, flattened distally, distal inner
surface with 6 denticulations (including distal end). Mazxillule (fig.
66g) situated adjacent to lateral posterior surface of mouth cone,
consisting of stalk (with indistinct evidence of segmentation) bearing
node from medial posterior surface, node tipped by spinule; stalk
flattened distally, distal surface with 3 nodes, each tipped by spinule.

NO. 3574

COPEPOD CRUSTACEANS—LEWIS

a

Be ea PTY

Ficure 66.—Brachiella thynni Cuvier, 1829, female: a, dorsal view. Ventral view: b, pos-

terior end of neck and trunk showing maxillae (m-2), bosses on trunk (crosshatched areas),
posterior processes (pp), and attached male; c, left antennule; d, anterior end of cephalo-
thorax, left side showing antennule, antenna (a-2), mouth cone, mandible (mdbl), maxillule
(ma-l), and maxilliped; ¢, left antenna; f, left mandible; g, right maxillule (posterior view);
h, left maxilliped, showing musculature.
226-321—67——_12

178 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Shiino (1956b) describes an accessory lamella associated with the
maxillule (Shiino’s ‘‘maxilla’’) which was not found on the dissected
appendage although a cuticular flap is present on the mouth cone
anterior to the maxillule. Maxilla (fig. 660) subconical, with numer-
ous superficial annulations, distal end concave on inner half and
enveloping cylindrical bulla (not figured).

Maxilliped (fig. 66h) situated posterior and slightly lateral to
mouth cone, segmentation indistinct and incomplete although mus-
culature suggests 3-segmented condition. Appendage divisible
into heavy basal part and clawlike distal part; basal part with knob-
shaped projection from medial inner surface, with minute, doughnut-
shaped projection just distal to knob with series of minute denticula-
tions on distal inner surface. Clawlike distal part bluntly pointed
distally, with setule-like accessory process on distal inner surface,
with row of minute denticulations extending from medial inner surface
to base of accessory process.

DerscriPTION OF MALE.—Cephalothorax or prosome (figs. 67a,})
ovoid in both lateral and dorsal view, consisting of cephalon and
maxilliped-bearing segment. Division between prosome and urosome
distinct, complete. Urosome broadest anteriorly, tapered to narrow
posterior end; posterior end with large, lappet-like anal laminae,
also bearing pair of subconical caudal rami, rami without armature
elements.

Antennule (fig. 67c) 3-segmented, situated on lateral-anterior dorsal
surface. First segment broad proximally, proximal width more than
twice medial width, approximately 4 times distal width; length approx-
imately twice combined lengths of second and third segments. First
segment with spinule on distal half of anterior dorsal surface. Second
segment approximately two-thirds the length of third, with spinule on
distal half of anterior dorsal surface. Third segment with indentation
on medial anterior surface, indentation bearing single spinule; rounded
distal end with 4 setules. Antennae (fig. 67d) biramous, situated on
anterior ventral surface of cephalothorax, below antennule base and
lateral to mouth cone. Protopodite 1-segmented, originating from
angular projection of ventral surface of cephalothorax. Exopodite
lamellate, with spine on medial posterior surface, second spine on
distal-outer posterior surface (spines not noted by Shino, 1956b).
Endopodite 2-segmented, first segment approximately twice the length
of second, second with sharp indentation on distal mner surface,
bearing clawlike terminal spine and small spinule from indentation.

Mouth cone projecting anteriorly from anterior ventral surface of
cephalothorax, labrum and labium indistinctly separable. Labrum

NO. 3674 COPEPOD CRUSTACEANS—LEWIS 179

flat, with membranous lateral surface; labium rounded, with heavily
sclerotized band distally, band bearing fine membrane projecting
medially and membrane-like projection directed distally, latter topped

OJmm.

Ficure 67.—Brachiella thynni Cuvier, 1829, male: a, lateral view; b, dorsal view. Ventral
view: c, left antennule; d, left antenna; e, oral region, left side showing mouth cone,
mandible (mdbl), and maxillule (ma-1); f, right maxilla; g, right maxilliped; h, projections
on right maxilliped.

by row of small, bifid pads, pads topped by finely frilled membrane.
Mandible (fig. 67e) rodlike, originating from padlike swelling adjacent
to mouth cone. Maxillule basically similar to that of female except

180 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

node on medial surface bearing 2 spinules instead of 1, distal 3 nodes
not as prominent as in female. Maxillae and maxillipeds situated on
irregular, apron-like projection, posterior half of projection separated
from anterior half by distinct ridge. Maxillae (fig. 67/) well developed,
2-segmented, situated on anterior half of projection. Both maxillae
connected at inner proximal end by posteriorly bifurcate band of heavy
sclerotization. First segment well developed, with heavily sclerotized
ring at proximal end, duct from maxillary gland visible inside proximal
inner surface of appendage. Second segment fused to clawlike
terminal process.

Maxilliped (fig. 67g) 2-segmented, situated on posterior half of
apron-like projection. First segment of maxilliped elongate, with
3 adhesion surfaces on inner surface. Adhesion surfaces formed by
numerous minute, padlike or hook-shaped projections (fig. 67h).
Proximal adhesion surface associated with heavily sclerotized ridge
and pad, medial surface associated with swelling, swelling with
spinule; distal adhesion surface associated with swelling on distal
inner surface. Second segment elongate, with adhesion process
on distal half of posterior surface; terminal process short, clawlike,
distinct from segment.

Discussion.—The Hawaiian specimens differ from the description
given by Shiino (1956b) primarily in the characteristics of the male:
the shorter cephalothorax and longer trunk, the presence of 3 instead
of 2 tuberculated bulges (adhesion pads) on the first segment of the
maxiliped, and the presence of 2 spinules on the exopodite of the
antenna. Whether or not these differences warrant specific con-
sideration remains questionable. The similarity of the female and
the males, with the exceptions here noted, and the broad distribution
of the parasite and the pelagic hosts that it characteristically
parasitizes suggest, to the present author, that these differences are
due to intraspecific variation.

Brachiella regia, new species

FicuREs 68, 69

MateriaAL.—Iwo females and 2 males from the gill arches of
Lampris regius (Bonnet) captured in the Oahu region and examined
by Walter Fujii, at the Honolulu Aquarium. One of the females
(USNM 112936) has been designated as the holotype, 1 of the males
(USNM 113033) as the allotype, and the remaining male and female
(USNM 113034) have been designated as paratypes.

|
}
}
|

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 181

MrASUREMENTS.— (In mm) 1 female and 1 male:
Total length, excluding lateral

posterior processes but female

including median projection 9.08
Length of cephalothorax, anterior

to maxillae 4.70
Width of anterior region of

cephalothorax 0.86
Length from maxillae to trunk 0.78
Width of posterior region of

cephalothorax (at maxilla) 0.99
Length of trunk 3.07
Width of trunk 1.60
Length of median posterior process ee:
Length of dorsal posterior processes 0.52
Length of ventral posterior processes 0.58
Length of egg string 2.48
Total length, excluding posterior male

processes 3.44
Length of prosome 1.08
Width of prosome 0.90
Length of urosome 2.25
Width of urosome 0.81
Width at constriction between prosome

and urosome 0.77

DESCRIPTION OF FEMALE.— Body (fig. 68a) 2-parted, anterior part
consisting of cephalothorax, posterior of trunk. Cephalothorax elon-
gate, at sharp angle to longitudinal axis of trunk, consisting of cepha-
lon and maxilliped-bearing segment; anterior third with heavily
sclerotized tergum. Trunk, excluding posterior processes, approx-
imately three-fourths the length of cephalothorax, separable from
cephalothorax by distinct groove on lateral and ventral surfaces,
groove not as distinct dorsally. Trunk flattened dorsoventrally,
basically rectangular from dorsal viewpoint although with numerous
small irregularities. Posterior end (fig. 68c) with 2 pairs of short,
knoblike processes laterally, with rounded projection on ventral
median surface.

Antennules, antennae, maxillules, maxillae, and maxillipeds covered
with thick layer of cuticular material, material overlying thin, darker
inner layer. Segment division of mentioned appendages not ex-
tending through outer layer although usually discernible and distinct
in inner layer.

Antennule (fig. 68¢) 3-segmented, situated on ventral anterior sur-
face of cephalothorax. First segment large, irregular, approximately
114 times the combined lengths of second and third segments. Second

182 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

segment slightly more than half the length of third; third segment
elongate, rounded distally, distal end with 4 naked setules. Antenna
(fig. 689) biramous, originating from lateral anterior surface of cepha-
lothorax. Protopodite 1-segmented, inner, darker portion of cuticle
forming 3 knob-shaped proximal processes articulating with inden-
tations on knob-shaped processes of cephalothorax. Distal end of
protopodite irregular, indistinct along inner surface, at origin of endop-
odite. Exopodite appearing l-segmented, approximately three-
fourths the length of protopodite. Distal end of exopodite curved
inward slightly, outer margin broadly convex, surface finely dentic-
ulated, inner margin broadly concave. Endopodite small, 2-seg-
mented, first segment slightly longer than second, second with spinule
on distal end.

Mandible (fig. 68h) appearing 3-parted, proximal part broad proxi-
mally, tapered abruptly, remaining parts tapered gradually to bluntly
rounded distal end. Inner surface of distal part with 8-10 primary
denticulations and 2 secondary denticulations, 1 between the first and
second primary denticulations, the other between the second and
third. Maxillule (fig. 682) situated just posterior to mandible base,
consisting of stalk bearing palp on median posterior surface, palp with
2 spinules distally; distal surface of maxillule with 2 nodules and spi-
nule, both nodules with spinule distally. Maxillae (fig. 68a) elongate,
approximately 1% times the length of cephalothorax, situated at
posterior end of cephalothorax, well behind maxillipeds. Maxillae
separate to distal end, attached to small, top-shaped bulla.

Maxilliped (fig. 687) 2-segmented, extending anteriorly from pad-
like ledge slightly posterior to mouth cone. First segment strongly
developed, with smoothly irregular outline, bearing small, spinule-
like projection from medial inner surface. Second segment elongate,
with nodule on medial ventral surface, nodule bearing minute, sub-
conical projection. Distal end of second segment irregular, with step-
like indentation on inner surface, bearing 2 small, spinelike accessory
processes from inner surface and single, large, clawlike terminal
process.

DESCRIPTION OF MALE.—Body (figs. 69a, 6) tapered at both ends,
lateral margins parallel throughout most of length although constric-

Ficure 68.—Brachiella regia, new species, female: a, lateral view; b, anterior portion of
cephalothorax, dorsal view showing irregular cuticular splotches; c, posterior region of
trunk showing posterior processes and portion of egg strings, ventral view; d, same, from
dorsal viewpoint with all except base of egg strings removed; e, left antennule, ventral
view; f, oral region, right side showing antennule, antenna (a-2), mouth cone, mandible,
maxillule (ma-1), and maxilliped (mxpd), ventral view; g, right antenna, ventral view;
h, denticulated portion of mandible, lateral view [drawn by Z. Kabata]; 7, right maxillule,
ventral view; 7, right maxilliped, ventral view.

COPEPOD CRUSTACEANS—LEWIS 183

NO. 3574

\
i
’
!
’
1
1
1
1
1
'
1
'
'
1

wt
STH

atl
ree
Cosi bass

CBE

184 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

tion present immediately posterior to maxillipeds in 1 specimen (not
shown in figures). Posterior region with pair of unarmed subterminal
dorsal flaps (caudal rami?) and pair of slightly projecting, knoblike
terminal processes. Anterior third of body heavily sclerotized dor-
sally; mouth cone projecting anteriorly and ventrally, slightly past
anterior end of cephalothorax.

Antennule (fig. 69c) 3-segmented, situated lateral and _ slightly
posterior to anterior edge of mouth cone base. First segment more
than 1% times the combined lengths of second and third segments,

SS
0.2 mmc"

Ficure 69.—Brachiella regia, new species, male: a, lateral view; }, dorsal view. Lateral
view: c, right antennule; d, right antenna; e, denticulated portion of mandible [drawn
by Z. Kabata]; f, right maxillule; g, right maxilla; 4, right maxilliped.

broad proximally, tapered slightly to distal end. Second segment
short, collar shaped; third segment dactyliform, approximately twice
the length of second, distal end rounded, tipped by 3 naked setules.
Antenna (fig. 69d) biramous, situated slightly posterior and lateral to
antennule base. Protopodite 2-segmented, segments of approxi-
mately equal size, both irregular in outline. Exopodite 1-segmented,
small, less than one-third the length of protopodite, broadly rounded

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 185

distally, without armature. Endopodite 2-segmented, slightly more
than one-third the length of protopodite. First segment approxi-
mately twice the length of second, with small, finely denticulated
knob on inner surface. Distal surface of second segment flattened,
with 2 small spinules on inner half and single, larger spinule on outer
half.

Mandible (fig. 69¢) appearing 3-parted, rodlike; distalmost part
flattened, inner surface with 9 denticulations, the third slightly smaller
and considered a secondary denticulation by Kabata (from correspond-
ence). Maxillule (fig. 69f) adjacent to posterior lateral portion of mouth
cone base, consisting of stalk bearing single palp on median posterior
surface, palp with 2 terminal spinules; 2 elongate, spinule-tipped
dactyliform processes present on distal end of maxillule. Maxilla
(fig. 699) prehensile, 2-segmented, situated well posterior to mouth
cone base. First segment strongly developed, proximal and outer
margins flatly convex, inner surface indented medially, indentation
with pocket-like depression receiving terminal process of second
segment when segment flexed. Second segment short, heavily scle-
rotized, fused with clawlike terminal process, and bearing minute,
spinelike accessory process on inner surface.

Maxilliped (fig. 69h) 2-segmented, originating from irregular pro-
jection immediately posterior to maxillae. First segment strongly
developed, narrow proximally, flared to broad distal half; inner surface
indented distally, indentation with small concavity receiving terminal
process of second segment when segment flexed. Second segment
small, heavily sclerotized, fused with clawlike terminal process,
bearing small, spinelike accessory process from inner surface and
minute, spinule-like process from outer surface.

Discussion.—The species is placed in the genus Brachiella be-
cause the female cephalothorax is elongate and cylindrical, is flexed
backward, and is “covered” by a heavily sclerotized tergum. Addi-
tionally, the trunk of the female is swollen and flattened dorsoven-
trally, there are 4 pairs of posterior processes, an unpaired genital
process is present, and there is no visible abdomen or caudal rami.
The characteristics of the male cast some doubt on the inclusion of
the species in Brachiella. 'The prosome and urosome of the figured
specimen are in an essentially straight line although the urosome of
the second male specimen is at an angle to the prosome. The author
is deeply indebted to Dr. Z. Kabata for a rather exhaustive examina-
tion of the paratype material and a comparison of the male with the
male of Andropoda lampri (Scott, 1901). Based upon the comparison
of the males of B. regia and A. lampri and upon the similarity of the
female of B. regia with other members of the genus Brachiella, the
species is presently placed within this genus.

186 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

The female of Brachiella regia shows some affinities with species
such as B. concava Wilson, 1913, and B. gracilis Wilson, 1908, in the
nature of the cephalothorax and maxillae. Additionally, the short
posterior processes of the female are comparable with those of B.
pteroplateae Yamaguti and Yamasu, 1959, B. bera Yamaguti, 1939c,
and B. mitrata Wilson, 1915. The combination of the long reflexed

cephalothorax, the long maxillae, the short posterior processes, and |

the large genital process is, however, unique.

The male shows some affinities with the male of B. thynni Cuvier,
1829, in the general nature of the body. The appendages of the
cephalothorax, however, are strongly reminiscent of those of Andro-
poda lampri (Scott, 1901), although the mandible has one denticu-
lation that Kabata (correspondence) terms a secondary tooth, while
the denticulations in A. lampri are homogeneous. There is some
variation in the denticulation of the mandible of both the female and
the male, the female possessing 8-10 primary denticulations, the male
“secondary tooth” being slightly larger in the unfigured male than in
the figured male. Further, the posterior end of the male of B. regia
does not protrude as in Andropoda lampri, and the posterior processes
are not as distinct and do not project past the thick, heavily sclero-
tized cuticle.

The species name is derived from the name of the host, Lampris
regius.

Family Naobranchiidae
Naobranchia Hesse, 1863

Draenosis.—Female: Cephalothorax elongate, vermiform; trunk
swollen, anterior end including maxilla-bearing segment. Abdomen
and caudal rami usually distinct although sometimes covered by
egg masses. Eggs enclosed in membranous extension of cuticle,
enveloping posterior and at least part of lateral surface of trunk.
Antennule 2-5 segmented; antenna biramous. Mandible rodlike.
Maxillule bipartite, with or without palp; maxilla foliaceous, usually
with muscle bands extending length of appendage, forming organ
of attachment that envelops gill filaments of host. Mazxilliped
2-segmented, second segment fused with clawlike terminal process.
Oval adhesion pads or cup-shaped structures may be present adjacent
to maxillipeds.

Male: See Yamaguti, 1963, p. 303.

Naobranchia species
FigureE 70

Marertau.—One female (USNM 112937) from gill cavity of
spotted moray eel from Honolulu Aquarium.

NO. 3674 COPEPOD CRUSTACEANS—LEWIS 187

MEASUREMENTS.— (In mm) 1 female:

female
Total length, including egg masses 3. 75
Length of cephalothorax, to maxillae 2. 30
Length of cephalothorax, to maxillipeds 0. 36
Width of anterior region of cephalothorax 0. 37
Width of posterior region of cephalothorax, just anterior to maxillae 0. 94
Length from anterior to posterior end of maxillae 0. 94
Width of base of maxillae 0. 80
Length of trunk 2. 00
Width of trunk 2.33

DESCRIPTION OF FEMALE.— Body (figs. 70a, 6) separable into 2 parts,
vermiform cephalothorax and broad, posteriorly rounded trunk.
Anterior end of cephalothorax slightly expanded, from dorsal view-
point; tapered to narrow, rounded end, in lateral view, ventral sur-
face bearing antennules, antennae, mandibles, maxillules, and
maxillipeds. Margin of anterior end of cephalothorax with heavily
sclerotized band broken by several minute swellings associated with
minute channels through band, similar to nodules bearing hairlike
processes on caligoids. Remaining part of cephalothorax irregular
(irregularities may be due to preservation), with several superficial,
incomplete annuli. Trunk expanded, rounded from dorsal and ven-
tral viewpoints, overlapping dorsal posterior end of cephalothorax;
lateral and posterior surfaces covered by egg cases, cases projecting
dorsally at flat angle giving semi V-shaped appearance in trans-
verse section. No abdomen visible although possibly covered by egg
cases; caudal rami not distinct although 2 minute, knoblike projec-
tions present between egg cases at posterior end of trunk, projections
not visible unless egg cases separated. Outer covering of egg cases
appearing thick, with moderately sclerotized band on lateral region.

Because of the small size of the appendages, with the exception of
the second maxillae and maxillipeds, the presence of only a single
specimen in the collection and the resultant hesitancy to dissect the
appendages, the following description should be used with some
caution.

Antennule (fig. 70d) situated on lateral-median ventral surface of
cephalothorax, behind anterior end of body and slightly lateral to
mouth cone. Appendage 3-segmented, first segment longer than
combined lengths of remaining 2 segments; third segment bearing 4
minute, subconical projections. Antenna (fig. 70e) situated between
antennule and base of labrum, appearing uniramous; 2-segmented,
tipped by clawlike projection. Mouth cone distinct though small,
with distinct space between labrum and labium. Labrum appearing
pointed distally, with heavily sclerotized band medially; labium with
U-shaped opening distally. Mandible (fig. 70e) rodlike, flattened
distally, with approximately 6 falciform denticulations along distal

188 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

inner margin. ®. Maxillule (fig. 70e) nodular, situated adjacent to pos-
terior lateral surface of labium base, with 2 setule-like projections
distally. Maxillae (fig. 70f) fused, foliaceous, situated at posterior

& 0.04 mm.

Ficure 70.—Naobranchia species, female: a, dorsal view; b, lateral view (ma-2= maxillae).
Ventral view: c, anterior end of cephalothorax showing antennules (a-1) (antennae hidden
under antennules and not visible), mouth cone, maxillules, maxillipeds, and cup-shaped
structure lateral to mouth cone; d, left antennule; ¢, oral region showing antenna (a-2),
mouth cone, mandibles, and maxillule (ma-1l); f, maxilla; g, right maxilliped.

end of vermiform cephalothorax and included in anterior portion of
trunk. Fused maxillae forming obcordate ventral projection, fusion

appearing to be at distal ends of maxillae, not with body; maxillae
without distinct muscle bands.

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 189

Maxilliped (fig. 70g) 2-segmented, situated in small concavity
slightly posterior and lateral to labium base. First segment with
shelflike indentation on distal half of inner surface, shelf with pair of
nodules forming depression receiving distal end of second segment
terminal process when segment flexed. Second segment small, sub-
conical, tipped by indistinctly separable, claw-shaped terminal process.

Semicircular cuplike cuticular (?) formation present lateral to max-
illiped base, extending anteriorly to region of antennule base. Pos-
terior surface of formation with pair of membranous, flangelike projec-
tions bearing fine indistinct grooves.

Discussion.—The single specimen has several distinctive features.
The presence of the semicircular cuplike formation (adhesion pads?)
is found in N. aulopi Yamaguti (1939¢), although in this species they
are oriented in a longitudinal direction while in the Hawaiian specimen
they are oriented laterally. The apparent lack of muscle bands in
the maxillae and the fusion of these 2 appendages to each other and
not the body surface may be diagnostic although it may instead suggest
an immature condition as Pearse (1952a, fig. 126) indicates for N.
spinosa Pearse. The possible immature condition of the specimen
may also explain the distinctive cuplike structures. Yamaguti (1939c)
figures a distinct cuplike structure for a 3.2 mm specimen of N. aulopi
while Shiino (1958) states “striated pads just behind 2nd max-
ilipeds [= maxilipeds] rather low and inconspicuous” in a 5.07 mm
specimen of N. aulopi. Both of these authors figure the adult female
with eggs so that if this is a characteristic of immature specimens,
the pad is still present at the time ovulation commences. In adult
female specimens of N. variabilis Brian, deposited by R. Bere in the
U.S. National Museum (79146), there is no distinct process of this
type although there is a slight concavity in the same region and curved,
heavily sclerotized rodlike structures are visible in the cuticle.

The Hawaiian specimen appears to most closely resemble N. varia-
bilis Brian. The similarity is not only in the general shape of the
body in the 3-segmented antennule although Pillai (1962a) describes a
5-segmented appendage for this species. ‘The antenna of the Hawaiian
specimen is here described as uniramous although with some question.
The maxillules and maxillipeds also are similar to those described
for N. variabilis by Bere (1936) and Pillai (1962a).

Host-Parasite List

The list given below provides the scientific and common name
of the host, when available, and the parasitic copepods taken from
The list includes the copepods described

it in Hawaiian waters.

in the present paper and in Lewis, 1964a,b, 1966.
tions, the hosts are arranged in the same sequence used by Gosline
and Brock (1960) in their ‘““Handbook of Hawaiian Fishes.”

host
Carcharodon carcharias (white shark)

Prionace glauca (great blue shark)
Galeocerdo cuvier (tiger shark)

Pterolamiops longimanus (whitetip shark)
Carcharhinus melanopterus? (blacktip shark)
Sphyrna lewint (hammerhead shark)

Hexanchus griseus?
Unidentified sharks

?Aetobatus narinari (eagle ray)
Saurida gracilis (lizard fish)
Muraenidae (moray eels)

Fistularia petimba (cornet fish)
Aulostomus chinensis (trumpet fish)

Holocentrus xantherythrus
Myripristis pralinius
Lampris regius

Sphyraena barracuda
Pranesus insularum?

Seriola dumerilit (yellowtail)
Caranz melampygus?

Coryphaena hippurus (dolphin)

190

With few excep-

copepod parasites
Anthosoma crassum
Dinematura latifolia
Pandarus satyrus
Phyllothyreus cornutus
Nesippus crypturus
Alebion echinatus
Pandarus cranchii
Pandarus smithii
Kroyeria praelongacicula
Pandarus cranchii
Alebion echinatus
Paeon vaissieret
Demoleus heptapus
Alebion gracilis
Dinematura latifolia
Pandarus cranchii
Pandarus smithii
Trebius caudatus
Caligus kala
Pseudotaeniacanthus puhi
Naobranchia species
Dentigryps bifurcatus
Nesippus costatus?
Dentigryps bifurcatus
Caligus ligatus
Caligus flexispina
Caligus ligatus
Nesippus costatus?
Brachiella regia
Midias lobodes
Caligus ligatus
Nesippus costatus?
Dentigryps ulua
Caligus longipedis
Euryphorus nordmanni
Caligus coryphaenae
Caligus quadratus
Charopinopsis quaternia

NO. 3574 COPEPOD CRUSTACEANS—LEWIS 191

host
Mulloidichthys auriflamma
Parupeneus pleurostigma
Chaetodon fremblit

Chaetodon quadrimaculatus
Chaetodon miliaris

Dascyllus albisella

Pomacentrus jenkinst
Bodianus bilunulatus
Unidentified labrid
Scarus species
Zanclus canescens
Acanthurus guttatus
Acanthurus nigroris
Acanthurus olivaceus

Acanthurus dussumiert

Acanthurus xanthopterus
Acanthurus mata
Acanthurus triostegus sandvicensis

Ctenochaetus strigosus
Zebrasoma flavescens

Naso lituratus
Naso hexacanthus

Naso unicornis

Acanthocybium solandri (wahoo, ono)

Auaxis thazard (frigate mackerel)
Katsuwonus pelamis (skipjack)

copepod parasites
Hatschekia breviramus
Lepeophtheirus dissimulatus
Dentigryps bifurcatus
Caligus kapuhili
Lepeophtheirus dissimulatus
Nesippus costatus?
Caligus kapuhili
Caligus kala
Caligus ligatus
Caligus kala
Dentigryps bifurcatus
Caligus flexispina
Nesippus costatus?
Nesippus costatus?
Caligus kalumat
Nesippus costatus?
Dentigryps bifurcatus
Lepeophtheirus dissimulatus
Peniculus calamus?
Caligus ligatus
Lepeophtheirus dissimulatus
Peniculus calamus?
Nesippus costatus?
Peniculus calamus?
Nesippus costatus?
Dentigryps bifurcatus
Lepeophtheirus dissimulatus
Caligus flexispina
Caligus randalli
Peniculus calamus?
Nesippus costatus?
Peniculus calamus?
Lepeophtheirus dissimulatus
Norion expansus
Norion expansus
Dentigryps bifurcatus
Anuretes serratus
Anuretes menehune
Lepeophtheirus dissimulatus
Caligus kala
Caligus ligatus
Peniculus calamus?
Anuretes menehune
Lepeophtheirus? fallolunulus
Peniculus calamus?
Gloiopotes hygomianus
Brachiella thynni
Caligus productus
Caligus coryphaenae
Caligus productus
Caligus bonito

192 PROCEEDINGS OF THE NATIONAL MUSEUM

host
Euthynnus yaito (little tuna)

Neothunnus macropterus (yellowfin tuna)
Parathunnus sibt

Thunnus albacares (in Shiino, 1963a)
Thunnus obesus (in Shiino, 19638a)
Istiompax marlina? (black marlin)
Makaira ampla? (Pacific blue marlin)
Makaitra audax (striped marlin)

Remoropsis brachypterus
Rhombochirus osteochir
Pervagor spilosoma
Ostracion lentiginosus
Diodon holocanthus
Plankton

VOL. 121

copepod parasites

Caligus coryphaenae
Caligus asymmetricus
Caligus productus
Caligus pelamydis
Elytrophora brachyptera
Caligus productus
Elytrophora brachyptera
Brachiella thynni
Brachiella thynni
Brachiella thynni
Gloiopotes huttoni
Gloiopotes huttoni
Pennella histiophoriv?
Gloiopotes huttoni
Pennella species
Lepeophtheirus crassus
Peniculus calamus?
Anchistrotos moa
Nesippus costatus?
Caligus coryphaenae (immature

male)

Literature Cited

(*Reference not seen by author.)

BatrD, W.
1847. Ona new species of Pennella. Ann. Mag. Nat. Hist., vol. 19, p. 280,
1 fig.
BARNARD, K. H.
1955. South African parasitic Copepoda. Ann. South African Mus., vol.
41, no. 4, pp. 223-312, figs. 1-33.
1957. Additions to the fauna-list of South African Crustacea. Ann. Mag.
Nat. Hist., ser. 12, vol. 10, pp. 1-12, figs. 1-8.
BASSETT-SMITH, P. W.
1896a. A list of the parasitic Copepoda of fish obtained at Plymouth. Journ.
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1896b. Notes on the parasitic Copepoda of fish obtained at Plymouth, with
descriptions of new species. Ann. Mag. Nat. Hist., ser. 6, vol. 18,
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1898a. Further new parasitic copepods found on fish in the Indo-Tropical
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1898b. Some new or rare parasitic copepods found on fish in the Indo-
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372, pls. 10-12.
1899. A systematic description of parasitic Copepoda found on fishes, with
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BrERE, RUBY
1936. Parasitic copepods from Gulf of Mexico fish. American Midl. Nat.,
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Bonnet, D.
1948. Some parasitic copepods from Hawaiian fishes. Proc. Hawaiian
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Brapy, G. S.
*1910. Die marinen Copepoden, 1: Ueber die Copepoden der Stiimme
Harpacticoida, Cyclopoida, Notodelphyoida und Caligoida. No.
8 (vol. 5) in Zoology (vol. 11) in Drygalski, Deutsche Siidpolar-
Expedition 1901-1903, pp. 497-593.
BrIANnN, ALESSANDRO
1898. Catologo di copepodi parassiti dei pesci della Liguria. Atti Soc.
Ligustica Sci. Nat. Geogr., vol. 9, pp. 5-31, pls. 1-4.
1899a. Di aleuni crostacei parassiti dei pesci dell’Isola d’Elba. Atti Soc.
Ligustica Sci. Nat. Geogr., vol. 10, pp. 3-10, figs. 1-5.
1899b. Crostacei parassiti dei pesci dell’Isola d’Elbe, 2: Contribuzione.
Atti Soe. Ligustica Sci. Nat. Geogr., vol. 10, pp. 197-207.
1901. Caso di anomalia verificatosi su di una Brachiella del Tonno. Bull.
Mus. Zool. Anat. Comp. Univ. Genova, 3 pp., 1 fig.
1908. Sulla Lophoura edwardsii Kolliker e sopra alcuni altri copepodi del
Golfo di Genova. Atti Soc. Ligustica Sci. Nat. Geogr., vol. 14,
pp. 3-11.

193
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194. PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

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NO. 3574

SHIINO, S.
1952.

1954a

1954b.

1954c.

1954d.

1954e.

1954f.

1955a.

1955b.

1956a.

1956b.

1957a.

1957b

1957¢

1957d

1958.

1959a

COPEPOD CRUSTACEANS—LEWIS 201

M.

Copepods parasitic on Japanese fishes, 1: On the species of Caligus
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. Copepods parasitic on Japanese fishes, 3: On two new species of the
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Copepods parasitic on Japanese fishes, 4: The family Euryphoridae.
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On the parasitic copepod, Lepeophtheirus watanabei n. sp., found on
the fish Myzxocephalus raninus J. & S. Bull. Japanese Soc. Sci.
Fish., vol. 20, no. 2, pp. 96-100, 1 fig.

Note on a new parasitic copepod, Caligus brevis n.sp. Bull. Japa-
nese Soe. Sci. Fish., vol. 20, no. 3, pp. 178-183, figs. 1-2.

On the male form of the copepod, Caligus katuwo Yamaguti, parasitic
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63, pp. 246-249, 1 fig.

A. new piscicola copepod belonging to the genus Caligus from Matu-
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figs. 1-3.

On the male of Caligus cordiventris Shiino. Zool. Mag., vol. 64, no.
3, pp. 65-67, 1 fig.

Copepods parasitic on Japanese fishes, 10: The redescription of three
species of Caligus. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 2,
pp. 233-241, figs. 1-4.

Copepods parasitic on Japanese fishes, 12: Family Lernaeopodidae.
Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 2, pp. 269-311, figs.
1-20.

Copepods parasitic on Japanese fishes, 13: Parasitic copepods col-
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. Copepods parasitic on Japanese fishes, 14: Three species from Sphe-
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Copepods parasitic on Japanese fishes, 15: Eudactylinidae and
Dichelesthiidae. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 3,
pp. 392-410, figs. 1-7.

. Copepods parasitic on Japanese fishes, 16: Bomolochidae and Taeni-
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411-428, figs. 1-6.

Parasitic copepods from fishes collected in the Indian Ocean. Ann.
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. Sammlung der parasitischen Copepoden in der Prafekturuniversitat
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. Revision der auf Goldmakrele, Coryphaena hippurus L., schmarot-
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no. 1, pp. 1-34, figs. 1-9.

. Neuer Artname fiir japanische Exemplare von Caligus bonito. Bull.
Biogeogr. Soc. Japan, vol. 20, no. 11, pp. 51-57, figs. 1-2.

202 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Suino, S. M.

1959d. Ostpazifische parasitierende Copepoden. Rep. Fae. Fish. Pref. Univ.
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1960a. Copepods parasitic on the fishes collected on the coast of Province
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1960b. Parasitic copepods of fishes from the eastern Pacific. Rep. Face.
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1960c. Copepods parasitic on remoras from the Bay of Bengal. Rep. Fac.
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1963a. Parasitic copepods of the eastern Pacific fishes. Rep. Fac. Fish.
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1963b. Parasitic copepods of the eastern Pacifie fishes, 4: On Clavellistes
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STesBInG, T. R. R.

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NO. 3574 COPEPOD CRUSTACEANS—LEWIS 203

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U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum
SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3575

SOME BATHYAL POLYNOIDS
FROM CENTRAL AND NORTHEASTERN PACIFIC
(POLYCHAETA: POLYNOIDAE)

By Marian H. PEtTiIsone

Associate Curator, Division of Worms

Polychaetes obtained from bathyal and abyssal regions are delicate
and, thus, easily mutilated. Among the polynoids, the dorsal scales
or elytra are usually missing; the styles of the antennae and the
tentacular, dorsal, ventral, and anal cirri are often partly or completely
missing; the setae and even the parapodia may be broken off; in
addition, the body may be fragmented.

This study of some deepwater polynoids was prompted by the dis-
covery of a single, somewhat mutilated specimen of polynoid collected
in 900 fathoms off the mouth of the Columbia River, Oregon. It is
herein described as a new species belonging to a new genus, Bathyad-
metella. As the name indicates, it shows certain affinities to Admetella
MelIntosh.

The generic standing of Admetella, based on Polynoe (Admetella)
longipedata McIntosh (1885), was first indicated by Darboux (1900).
Darboux gave a diagnosis for the genus based on the incomplete de-
scription by McIntosh. The description of A. longipedata has been
supplemented subsequently by other polychaete workers, particularly
by Augener (1906), Ehlers (1908), and Ditlevsen (1917). Some speci-

il

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

mens of A. longipedata in the U.S. National Museum, from the
Albatross collections, are in relatively good condition, enabling me to
supplement further the previous descriptions and to give an emended
diagnosis for the genus.

Chamberlin (1919) added 2 new species to Admetella McIntosh,
A. dolichopus and A. hastigerens. 'The type specimens of both species
in the U.S. National Museum have been examined and are herein
referred to as A. longipedata. Hartman (1938) examined the type
specimen of Polynoe(?) renotubulata Moore (1910) in the U.S. National
Museum and referred it to Admetella. The specimen has been re-
examined and is herein designated the type species for the new
genus, Bathymoorea.

This study was aided in part by a grant from the National Science
Foundation (NSF GB-1269).

Family Polynoidae Malmgren
Genus Admetella McIntosh, 1885; emended

Type species: Polynoe (Admetella) longipedata McIntosh, 1885, by
monotypy. Gender: feminine.

Diagnosis: Polynoids with body large, oval, flattened; seements
more than 50 (52-82). Prostomium with 2 long palps and 3 antennae;
median antenna with large ceratophore inserted on middle third of
prostomium; lateral antennae inserted on anterolateral extensions
of prostomium (1.e., lepidonotoid); with paired antennal scales.
Tentacular segment (I) achaetous, with 2 pairs tentacular cirri,
with bulbous facial tubercle between bases of palps. Buccal segment
(II) achaetous, with paired, long, ventral buccal cirri and first pair
elytra; with dorsal, low, transverse nuchal fold. Elytra more than
20 pairs, arranged on segments 2, 4, 5, 7, then on alternate segments
to 23, and then on every third segment to end of body. Parapodia
very long, subbiramous; notopodia small; both rami with elongated
digitiform acicular processes. Notosetae form small bundles. Neu-
rosetae numerous, forming fan-shaped bundles. Both noto- and
neurosetae long, slender, transparent, fragile, inflated, and flattened
distally, with faint spinous rows and tapered tips.

Admetella longipedata (McIntosh)
Fieures 1, 2

Polynoe (Admetella) longipedata McIntosh, 1885, p. 124, pl. 14, fig. 5; pl. 20,
Hew Os spl da hoe lf

Admetella longipedata Darboux, 1900, p. 103.—Augener, 1906, p. 123.—Ehlers,
1908, p. 40, pl. 2, figs. 10, 11; pl. 3, figs. 1-5.—Horst, 1917, pp. 101, 140.—
Ditlevsen, 1917, p. 37.—Fauvel, 1932, p. 27.—Eliason, 1951, p. 133.

Polynoe mirabilis Treadwell, 1906, p. 1149; 1920, p. 590. [Not McIntosh, 1885.]

Se |

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE 3

Admetella hastigerens Chamberlin, 1919, p. 64, pl. 9, figs. 6-8.—Treadwell, 1923,
p. 3.
Admetella dolichopus Chamberlin, 1919, p. 67, pl. 10, fig. 1.

Remarks: The following study is based on 12 specimens in the
U.S. National Museum, collected from seven Albatross stations from
the vicinity of Hawau, Philippine Islands, and Lower California to off
Central America. The specimens were identified by Treadwell as
Polynoe mirabilis McIntosh (Treadwell, 1906, 1920) and Admetella
hastigerens Chamberlin (Treadwell, 1923) and by Chamberlin (1919)
as 2 new species of Admetella, A. hastigerens, and A. dolichopus. All
of the above are herein referred to Admetella longipedata (McIntosh).
Fauvel (1914, p. 39) questioned the identification by Treadwell (1906,
p. 1149) of Polynoe mirabilis from off Hawaii. Uschakov (1950, p.
157) also questioned the identifications in both records of Treadwell
(1906, 1920). Eliason (1951, p. 133) questionably referred A. hasti-
gerens to A. longipedata.

Material examined: From seven Albatross stations, Sta. 2635
(Apr. 18, 1891), off Mexico, 20°47’ N, 106°15’ W, 2022 fms, dark
green mud (USNM 19325, type of Admetella dolichopus Chamberlin) ;
Sta. 4022 (June 21, 1902), vicinity of Kauai, Hawaiian Islands, 374-
399 fms, coral, sand, foraminifera, rock (USNM 5458, identified by
Treadwell as Polynoe mirabilis, 2 spec.); Sta. 4621 (Oct. 21, 1904),
southwest coast of Central America, 6°36’ N, 81°45’ W, 581 fms,
green sand (USNM 19326, type of Admetella hastigerens Chamberlin) ;
Sta. 5114 (Jan. 20, 1908), Balayan Bay and Verde Island Passage,
Philippine Islands, 13°36’ N, 120°45’ E, 340 fms, fine sand (USNM
17507, identified by Treadwell as Polynoe mirabilis, 5 spec.); Sta.
5122 (Feb. 2, 1908), east coast of Mindoro, Philippine Islands, 13°21’
N, 120°30’ E, 220 fms, green mud (USNM 17606, identified by Tread-
well as Polynoe mirabilis, 1 spec.) ; Sta. 5677 (Mar. 17, 1911), between
Ballenas Bay and Santa Maria Bay, Lower California, 25°23’ N,
113°16’ W, 735 fms, green mud, fine sand (USNM 19149, by Tread-
well as Admetella hastigerens, 1 spec.); Sta. 5685 (Apr. 22, 1911),
south of Abrejos Point, Lower California, 25°42’ N, 113°30’ W,
645 fms, black sand, coral (USNM 19150, identified by Treadwell
as Admetella hastigerens, 1 spec.).

Description: Length 50 to 100 mm; width, including parapodia
(without setae), 16 to 35 mm; segments 64 to 82, last 4 to 6 very small.
Body spindle shaped, widest in middle, tapering anteriorly and
posteriorly, arched dorsally, flattened ventrally. Parapodia very long,
as long as width of body, inflated and flattened transversely (fig. 1c).
Elytra (usually missing) on large inflated ely trophores, 25 to 31 pairs,
arranged on segments 2, 4, 5, 7, then alternate segments to 23, and

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

then on every third segment to end of body. Elytra missing on all
specimens examined (according to Treadwell, 1923, elytra are large,
delicate, gray to black in color). Dorsal tubercles on cirrigerous
segments inflated, thin walled, continuous as inflated ridge to base of

Ficure 1.—Admetella longipedata: a, dorsal view anterior end (USNM 19150) with phar-
ynx partially extended, ocular areas not pigmented, styles of median antenna and right
lateral antenna and right antennal scale missing, styles of right tentacular cirri missing (1),
elytra and styles of buccal cirri missing (II), styles of dorsal cirri missing (II); d, lateral
view anterior end (USNM 17606) with pharynx partially extended, ocular area pig-
mented, style of median antenna missing, tentacular cirri missing (I), style of buccal
cirrus and elytron missing (II); c, ventral view of several segments on right half in middle
part of body. (I-II]=segments, anSc=antennal scale, bC=cirrophore buccal cirrus,
elp=elytrophore, fT=facial tubercle, neP=nephridial papilla, nuF=nuchal fold, vC=
ventral cirrus.)

short cylindrical cirrophores of dorsal cirri; styles of dorsal cirri long,
extending beyond the tips of neurosetae, delicate, with tapered tips.
Prostomium (figs. 1@,b) bilobed, much wider than long, with pair of
very large rounded ocular areas, pigmented or colorless. Ceratophore
of median antenna large, occupying middle third of prostomium and
projecting dorsally; style usually missing (delicate, slender, extending

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE 5

to about segment 9, according to Treadwell, 1923). Ceratophores of
lateral antennae formed by continuations of anterolateral borders of
prostomium; styles cylindrical, tapering distally to slender tips, with
slight subterminal enlargements. Paired, thin, flattened, subtriangu-
lar processes or antennal scales attached between bases of ceratophores
of lateral antennae and median antenna; antennal scales broad basally,
tapering to blunt tips, tending to curl around basal parts of lateral
antennae (antennal scales sometimes broken off but scars of attachment
visible). Paired, very long, tapering, smooth palps.

First or tentacular segment (figs. 1a,b) dorsally forms short ring,
projecting lateral to prostomium, achaetous, with 2 pairs tentacular
cirri; cirrophores large, inflated; styles long, smooth, delicate, with
attenuate tips; a bulbous facial tubercle between bases of palps, con-
tinuous with a longitudinal ridge on upper lip. Second or buccal
segment with first pair elytrophores; parapodia rudimentary, achae-
tous; with dorsal, low, slightly bilobed, transverse nuchal fold between
elytrophores; forms posterior lip of mouth ventrally; ventral or buccal
cirri with large inflated cirrophores, with styles long, similar to ten-
tacular cirri. Third segment with setigerous biramous parapodia not
especially elongated. Beginning with segment 4, parapodia elongated.
Ventral cirri on segments 3-5 extend beyond tips of neuropodia. On
remaining segments, ventral cirri smaller, subulate, in middle of
elongated neuropodia (figs. 2a,6).

Parapodia (figs. 2a,b) elongated, inflated, thin walled, flattened
transversely, subbiramous. Notopodia small lobes on anterodorsal
faces of elongated neuropodia, inflated basally with elongated digiti-
form acicular processes, with small bundles of notosetae (easily broken
and may be overlooked). Notosetae somewhat more slender than
neurosetae, transparent, flattened distally, tapered to rounded tips,
with faint spinous rows (fig. 2c). Neuropodia elongated, somewhat
flared distally, postsetal lobes subtriangular, presetal lobes extending
as long digitiform acicular processes. Acicula of both rami yellowish,
tapering distally to fine tips within the acicular processes. Neurosetae
numerous, forming fan-shaped bundles. Neurosetae (fig. 2d) long,
slender, flexible, transparent, iridescent, fragile (may be mostly
broken), with long bare basal regions and flattened distal spinous
regions consisting of transverse rows of low spines; tips very thin, bare,
tapered (when broken, fracture in various ways, as indicated by pl.
12A, fig. 17 in McIntosh, 1885; pl. 3, figs. 1-5 in Ehlers, 1908; pl. 9,
figs. 7, 8, in Chamberlin, 1919).

Anus dorsal, surrounded by small parapodia of last 4 to 5 segments.
Anal cirri not noted. Nephridial papillae (figs. 1¢,26) on ventral bases
of parapodia, begin on segment 6 and continue posteriorly, inflated
basally, tapered distally to short tubes directed between parapodia.

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Pharynx partially extended in most preserved specimens, thin walled, |
with conspicuous dorsal longitudinal ridge on basal part which is con- |
tinuous with bulbous facial tubercle between bases of palps (figs. 1a,6).
Pharynx reddish to blackish; distally with soft papillae and 2 pairs
light-brown interlocking jaws. Color (according to Treadwell, 1920)
livid flesh pink, some cirri showing violet color. .

Remarks: Admetella longipedata is characterized particularly by |
the paired antennal scales on the prostomium. It may be distin- |

aE
<= a SS
ae vC
b neP

Ficure 2.—Admetella longipedata: a, parapodium of elytragerous segment, anterior view,
elytron missing, position of acicula shown in dotted lines; b, parapodium of cirrigerous
segment, posterior view, lateral part of dorsal tubercle indicated, style of dorsal cirrus
missing; ¢, tip of notoseta; d, tip of neuroseta. (dT=dorsal tubercle, neP=nephridial
papilla, vC=ventral cirrus.)

guished from Bathyadmetella and Bathymoorea according to the key |
on page 13.

Distribution: Widely distributed deep-sea form, North and South |
Atlantic, Indian Ocean, North Central Pacific. In 220 to 2779 fms.

1) UY)

Bathyadmetella, new genus

Type species: B. commando, new species. Gender: feminine.
Diagnosis: Polynoids with body large, oval, flattened, segments
more than 50 (58). Prostomium with 2 long palps and 3 antennae;

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE i

median antenna with ceratophore inserted on anterior part of pro-
stomium; lateral antennae inserted on anterolateral extensions of pro-
stomium (i.e., lepidonotoid) ; with paired long antennal sheaths dorsal
to lateral antennae. Tentacular segment (I) achaetous, with 2 pairs
tentacular cirri, with paired long tentacular sheaths between palps
and tentacular cirri, with bulbous facial tubercle between bases of
palps. Buccal segment (II) with parapodia rudimentary, achaetous,
with paired, long, ventral buccal cirri and first pair dorsal ely trophores.
With lamellar papillae on ventral lip and ventral bases of parapodia
on segments 2 and 3. Elytra or elytrophores more than 20 pairs (23),
arranged on segments 2, 4, 5, 7, then on alternate segments to 23,
and then on every third segment to end of body. Parapodia very
long, subbiramous; notopodia small, with digitiform acicular lobes,
achaetous; neuropodia with elongated acicular processes. Neuro-
setae numerous, forming brushlike bundles, transparent, fragile, in-
flated and flattened distally, with faint spinous rows and tapered tips.

Bathyadmetella commando, new species

Ficurss 3, 4

Material examined: The species is represented by a single damaged
specimen, collected in a trawl haul, southwest of the mouth of the
Columbia River, Oregon, 45°45’ N, 125°09’ W, 900 fms, 2.6° C
bottom temperature, 34.372°/oo salinity, May 29, 1964, M/V Com-
mando, M.S. Alton, collector. The holotype is deposited in the
U.S. National Museum (USNM 33419).

Description: Length 55 mm, width of body 6 mm; width including
parapodia 24 mm, including setae 34 mm; segments 58, last few small.
Body widest in middle, tapering gradually anteriorly and more so
posteriorly; flattened dorsoventrally; with extremely long parapodia.
Integument thin, delicate. Body reddish brown, darker on parapodia
and distal halves of elytrophores, lighter ventrally. Elytra all missing;
elytrophores elongated, inflated basally, 23 pairs, arranged on seg-
ments 2, 4, 5, 7, then on alternate segments to 23, and then on every
third segment to end of body. Dorsal tubercles on cirrigerous seg-
ments thin walled, inflated, continuous as low ridges to cirrophores
of dorsal cirri; styles of dorsal cirri all missing.

Prostomium (figs. 3a,b) bilobed, wider than long, with paired very
large bulging eyes occupying most of posterior half of prostomium;
eyes with semicircular, dark purplish pigmented cups basally. Cer-
atophore of median antenna short, cylindrical, on anterior border
prostomium; style missing. Ceratophores of lateral antennae long
cylindrical extensions of anterolateral borders of prostomium; styles
long, slender, with slight subterminal enlargements and slender tips.
Paired elongate antennal sheaths dorsal to the lateral antennae, wider

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

basally, tapering distally; sheaths may completely enclose lateral
antennae. Paired palps very long, tapering, and smooth. Prosto-
mium with triangular pigmented area around base of ceratophore of
median antenna, extending posteriorly between the two large eyes.

Ficure 3.—Bathyadmetella commando, new species: a, dorsal view anterior end, styles of
median antenna and left tentacular cirri missing, ceratophores of lateral antennae hidden
by antennal sheaths (shown in dotted lines on right); b, same, ventral view; c, anterior
view of achaetous parapodium of tentacular segment (I); styles of tentacular cirri missing;
d, posterior view of left achaetous parapodium of buccal or first elytragerous segment
(Il). (anSh=antennal sheath, bC=buccal cirrus, elp=elytrophore, ff! = facial tubercle,
1P=lamellar papilla, tsh=tentacular sheath.)

First or tentacular segment (figs. 3a—c) forms short ring dorsally,
projecting laterally to prostomium, achaetous, with 2 pairs long
slender tentacular cirri similar to lateral antennae, cirrophores short,
cylindrical, with a projecting acicular process. Paired long tentacular

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE 9

_ sheaths attached basally on tentacular segment, wider basally, tapering
distally, as long as tentacular cirri and may partially encircle the
_palps. A bulbous facial tubercle between bases of palps, continuous
with a longitudinal ridge on upper lip. Second or buccal segment
(figs. 3b, d) with first pair elytrophores; parapodia rudimentary, not
elongated, with noto- and neuroacicular achaetous lobes; ventral or
buccal cirri with inflated cirrophores and long styles; a lamellar
papilla on ventral base of parapodium; ventral lip (fig. 36) inflated,
longitudinally grooved, with 11 lamellar papillae in three irregular
rows. Setigerous parapodia elongated from third segment on.
Lamellar papillae at ventral parapodial bases of segments 2 and 3;

dT.
SS -
SS =f E
= oO
|
a vC i
neP
e
=
oO
wl
\
a I eee SES SIL a [2a = See f
— 7 Sea ees aS
a SS
h ——

Ficure 4.—Bathyadmetella commando, new species: a, cirrigerous parapodium, posterior
view, style of dorsal cirrus missing; b, elytragerous parapodium, anterior view, elytron
missing, position of acicula shown in dotted lines; c, tips of neurosetae. (dT'=dorsal
tubercle, neP=nephridial papilla, vC=ventral cirrus.)

ventral cirri on these 2 segments extra long, extending beyond the
distal tips of the neuropodia (fig. 3b). Rest of ventral cirri short,
subulate, in middle of elongated neuropodia (figs. 4a, 6).

Parapodia (figs. 4a,b) elongated, subbiramous. Notopodia small
lobes on anterodorsal faces of neuropodia, inflated basally, tapering
to short digitiform acicular lobes, enclosing the slender tips of the
notoacicula; notosetae lacking. Neuropodia flattened transversely,
inflated subdistally, postsetal lobes subtriangular, presetal lobes with
elongate acicular processes containing the neuroacicula. Neurosetae
numerous, forming brushlike transparent bundles. Neurosetae with
long, bare, basal regions, distally wider, flattened, with somewhat

10 PROCEEDINGS OF THE NATIONAL MUSEUM

spirally arranged fine spinous rows, tapered rather abruptly to very:
thin tips (fig. 4c).

Anus dorsal in position, surrounded by last few small segments
(damaged somewhat in thisarea). Analcirrinot observed. Nephrid-|
ial papillae on ventral bases of parapodia, beginning on segment 4)
and continuing posteriorly, small at first, becoming longer (shorter)
than ventral cirri), cylindrical and directed between the parapodia |
(fig. 4a). Pharynx not extended. The single specimen is ovigerous. |

Remarks: Bathyadmetella commando is unique among the polynoids
in having long antennal and tentacular sheaths and lamellar papillae!
on the ventral lip and ventral bases of the second and third segments. |
It is also unusual for its extremely large bulging eyes. It may be}
distinguished from Admetella and Bathymoorea according to the key |
on page 13.

Distribution: Northeastern Pacific (off Oregon). In 900 fms. {

|

'

Bathymoorea, new genus

Type species: Polynoe(?) renotubulata Moore, 1910. Gender: |
feminine. |

Diagnosis: Polynoids with relatively few segments (33). Pro- |
stomium with 2 long palps and 8 antennae; median antenna with |
ceratophore inserted in middle of prostomium; lateral antenna in- |
serted on anterolateral extensions of prostomium (i.e., lepidonotoid).
Tentacular segment (I) achaetous, with 2 pairs tentacular cirri, |
with bulbous facial tubercle between bases of palps. Buccal segment |
(II) with first pair elytra, setigerous parapodia, and long ventral |
buccal cirri. Elytra 14 pairs, arranged on segments 2, 4, 5, 7, then |
on alternate segments to 23, 26, 28, and absent on last 5 small seg-
ments. Parapodia elongated, subbiramous, with small notopodia;
both rami with elongated, digitiform acicular processes. Notopodia
with small tufts of setae. Notosetae delicate, finely spinous. Neu-
ropodia with dense, brushlike bundles of transparent setae. Neuro-
setae distally expanded and flattened, with faint spinous rows and
tapered tips. Nephridial papillae remarkably elongated, some ex-
tending to tips of neuropodia.

Bathymocrea renotubulata (Moore), new combination
FIGURE 5
Polynoe(?) renotubulata Moore, 1910, p. 368, pl. 31, figs. 59-64.
Admetella renotubulata Hartman, 1938, p. 123.
Material examined: Holotype Polynoe(?) renotubulata (USNM
16878), Albatross Sta. 4397 (Apr. 1, 1904), off Santa Catalina Islands, |
southern California 33° 43’ N, 117° 42’ W, 2196 to 2228 fms, gray
mud.

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE Lib

Description: Length 26 mm, width of body 3.6 mm; width includ-
ing parapodia 13 mm, including setae 22 mm; segments 33, last few
segments very small. Body subfusiform, widest in middle, tapering
anteriorly and posteriorly. Elytra (usually missing) on large inflated
elytrophores, 14 pairs, arranged on segments 2, 4, 5, 7, then on alter-
nate segments to 23, 26, 28. First elytron (observed by Moore,
now missing) suborbicular, of thick, soft, cushiony texture, thickly
covered with soft dome-shaped papillae with single coarse filament
at summit. Dorsal tubercles on cirrigerous segments inflated; cirro-
phores of dorsal cirri small; styles missing.

Prostomium (fig. 5a) bilobed, wider than long, with large pair,
opaque ocular areas. Ceratophore of median antenna in middle of
prostomium; style missing. Lateral antennae with short cerato-
phores formed by anterolateral extensions of prostomium; styles
tapered. Palps paired, thick, smooth, tapering.

First or tentacular segment projecting lateral to prostomium,
achaetous, with 2 pairs tentacular cirri; styles long, slender, tapered.
A bulbous facial tubercle between palps, continuous with longitudinal
ridge on upper lip. Second or buccal segment with first pair elytro-
phores, notopodial acicular lobes, and neuropodial acicular lobes
bearing bundles of neurosetae; cirrophores of ventral cirri large,
styles missing. Ventral cirri on remaining segments short, subulate
(fig. 5c).

Parapodia (figs. 56, c) elongated, subbiramous, with tapering yellow
acicula. Notopodia inflated basally, prolonged into long acicular
processes, with small tufts of notosetae. Notosetae delicate, finely
spinous (fig. 5d). Neuropodia with subconical postsetal lobes;
presetal lobes prolonged into delicate acicular processes, with long,
dense, brushlike bundles of neurosetae. Neurosetae transparent, dis-
tally expanded and flattened, with faint spinous rows, abruptly
tapering to entire tips (may be split, appearing bifid, fig. 5e).

Anus dorsal, surrounded by small posterior segments. Anal cirri
not observed. Nephridial papillae begin on segment 6 and continue
posteriorly, becoming remarkably elongated, some extending beyond
distal tips of neuropodia (figs. 5b, c). Pharynx not extended.

Remarks: Moore (1910) questionably placed this species in the
genus Polynoe. Hartman (1938) referred it to Admetella; however,
there is no trace of the triangular antennal scales characteristic of
Admetella. Other differences are indicated in the key on page 13.
As the specific name indicates, B. renotubulata is remarkable for the
length of the nephridial papillae.

Distribution: Off southern California. In 2196 to 2228 fms.

The 3 bathyal genera, Admetella, Bathyadmetella, and Bathymoorea,
have a number of characters in common, e.g., prostomia with paired

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

neP

Ficure 5.—Bathymoorea renotubulata: a, dorsal view anterior end, pharynx partially
extended, opaque ocular areas stippled, left palp and styles of median and right lateral
antennae missing, lower left and right tentacular cirri missing (I), right elytrophore and
parapodium missing (II), b, elytragerous parapodium, anterior view, elytron missing,
position of acicula shown in dotted lines; ¢, cirrigerous parapodium, posterior view, style
of dorsal cirrus missing; d, tip of notoseta; ¢, tips of neurosetae. (I-II=segments,
fT=facial tubercle, neP=nephridial papilla, vC=ventral cirrus.)

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE 13

large eyes or ocular areas; ceratophores of lateral antennae formed by
anterolateral continuations of prostomia (i.e., lepidonotoid); bulbous
facial tubercle present and continuous with ridges on upper lips; long
ventral cirri (buccal cirri) on segment II; parapodia very long, sub-
biramous, notopodia scarcely separated basally from the neuropodia,
both rami with digitiform acicular processes; neurosetae all of one
kind, numerous, long, transparent, flattened distally, distal tips very
thin, entire (unless broken). The 3 genera may be distinguished
according to the following key.

Key to Admetella, Bathyadmetella, and Bathymoorea

1. Segments 33. Elytra (or elytrophores) 14 pairs, on segments 2, 4, 5, 7, then
on alternate segments to 23, 26, 28, and absent from last 5 segments. Pro-
stomium without antennal scales or sheaths. Ceratophore of median anten-
na in middle of prostomium. Without tentacular sheaths. Without
nuchal fold. Buccal segment (II) with setigerous parapodia. Lower lip
region bare, without papillae. Notopodia with long acicular processes and
small bundles of delicate notosetae. Nephridial papillae extremely long,
may extend beyond neuropodial lobes . . . . Bathymoorea, new genus

B. renotubulata (Moore)

Segments more than 50. Elytra (or elytrophores) more than 20 pairs, on
segments 2, 4, 5, 7, then on alternate segments to 23, and then on every
third segment continuing to ead of body. Prostomium with antennal
scales or sheaths. Buccal segment (II) with parapodia rudimentary,
achaetous, with or without acicula. nee ee papillae not especially
elongated, shorter than ventral cirri. . . SRE a pee

2. Antennal scales short, triangular, attached am een Seeoanores of median and
lateral antennae. Ceratophore of median antenna in middle of prosto-
mium. Without tentacular sheaths. With low transverse nuchal fold on
segment II between first pair elytrophores. Lower lip region bare, without
papillae. Notopodia with long acicular processes and small bundles noto-
RetHeN Ls MoMe a. Gib nasi ce tue orice eee ng it hk AGINetelan Wichntosia

A. longipedata (McIntosh)

Antennal sheaths long, digitiform, attached basally to and enclosing lateral

antennae. Ceratophores of median antenna on anterior part of prosto-

mium. With paired long digitiform tentacular sheaths between palps and

tentacular cirri. Without nuchal fold. Lower lip region with series of
lamellar papillae. Notopodia with short acicular processes, achaetous.

Bathyadmetella, new genus

B. commando, new species

Literature Cited

AUGENER, HERMANN

1906. Westindische Polychaeten. Bull. Mus. Comp. Zool. Harvard, vol.

43, pp. 91-196, 8 pls.
CHAMBERLIN, Rawpu V.

1919. The Annelida Polychaeta. Mem. Mus. Comp. Zool. Harvard, vol.

48, pp. 1-514, 80 pls.
Darsovux, J. GASTON

1900. Recherches sur les Aphroditiens. Bull. Sci. France et Belgique,

vol. 33, pp. 1-274, 83 figs.
DITLEVSEN, HJALMAR

1917. AnnelidesI. Pt. 4 of vol. 4 in The Danish Ingolf expedition, 71 pp.,

6 pls.
EHLERS, ERNST

1908. Die bodensissigen Anneliden aus den Sammlungen der Deutschen
Tiefsee-Expedition. Jn Wissenschaftliche Ergebnisse der Deut-
schen Tiefsee-Expedition, 1898-1899, vol. 16, pp. 1-168, 23 pls.

Extason, ANDERS

1951. Polychaeta. In Report of the Swedish Deep-Sea Expedition, 1947-

1948, Zool., vol. 2, no. 11, pp. 131-148, 2 pls., 5 figs.
FAUVEL, PIERRE

1914. Annélides Polychétes non pélagiques provenant des campagnes de
l’ Hirondelle et de la Princesse-Alice (1885-1910). Res. Camp.
Sei. Monaco, vol. 46, pp. 1-432, 31 pls.

1932. Annelida Polychaeta of the Indian Museum, Calcutta. Mem.
Indian Mus., vol. 12, pp. 1-262, 40 figs.

HARTMAN, OLGA

1938. The types of the polychaete worms of the families Polynoidae and
Polyodontidae in the United States National Museum and the
description of a new genus. Proc. U.S. Nat. Mus., vol. 86, pp.
107-134, 41 figs.

Horst, R.

1917. Polychaeta errantia of the Siboga Expedition, 2: Aphroditidae and
Chrysopetalidae. In Siboga-Expedition, vol. 24b, pp. 1-140, 19
pls., 5 figs.

McInrosu, WILLIAM C.

1885. Annelida Polychaeta. In Challenger reports, vol. 12, pt. 3, pp. 1-554,

94 pls.
Moors, J. Percy

1910. The polychaetous annelids dredged by the U.S.S. Albatross off the
coast of southern California in 1904, 2: Polynoidae, Aphroditidae,
and Segaleonidae. Proc. Acad. Nat. Sci. Philadelphia, vol. 62,
pp. 328-402, pls. 28-33.

TREADWELL, AARON LOUIE

1906. Polychaetous annelids of the Hawaiian Islands collected by the
steamer Albatross in 1902. Bull. U.S. Fish Comm. Washington,
vol. 23, pp. 1145-1181, 81 figs.

14

a tt a eS ti a a A Ta

a

NO. 3575 BATHYAL POLYNOIDS—PETTIBONE 15

1920. Polychaetous annelids collected by the United States Fisheries
steamer Albatross in the waters adjacent to the Philippine Islands
in 1907-1910. Bull. U.S. Nat. Mus., no. 100, pp. 589-602, 8 figs.

1923. Polychaetous annelids from Lower California with descriptions of
new species. Amer. Mus. Novitat. New York, no. 74, pp. 1-11,
8 figs.

‘Uscuakovy, P.

1950. [Polychaeta from the Sea of Okhotsk.] Issled. dalnevost morei

SSSR, no. 2, pp. 140-234, 2 pls., 39 figs. [In Russian.]

U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum
SMITHSONIAN INSTITUTION «© WASHINGTON, D.C.
Volume 121 1967 Number 3576

NEW SPECIES AND RECORDS
OF PACIFIC AMPELISCIDAE
(CRUSTACEA: AMPHIPODA)

By J. Laurens BarnarpD

Associate Curator, Division of Crustacea

Ampeliscidae from the east tropical and warm-temperate Pacific
Ocean and from warm-temperate Japan collected by the “Albatross,”
“Velero III,” and individual collectors are reported upon in this paper.
These records and descriptions complete an inventory of all Pacific
Ocean materials of this family collected by the two expeditionary
vessels, on which previous reports have been made by Holmes (1908),
Shoemaker (1925), and J. L. Barnard (1954a, 1960, 1964a). Ampe-
liscidae are among the most dominantly occurring families of sub-
littoral and bathyal depths. Of particular interest are three new
species of Byblis from Japan which demonstrate a remarkable diversity
in that genus.

This study was supported by the Richard Rathbun Fund of the
Smithsonian Institution and was part of a project organized by
Dr. Fenner A. Chace, Jr., to complete the analysis of important ex-
peditionary collections deposited in the U.S. National Museum. The
writer is grateful also to the Beaudette Foundation of California for
assistance in the completion of the work in 1960.

Station data are presented in the list below. Depths are quoted in
fathoms as in the original data, but in the sections on distribution of

1

2, PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

species they are converted to meters to conform to international
practice.

List of Localities

Albatross Station Records

2787, Golfo de Penas, Chile, 46°47'30’’S, 75°15'00’’W, 61 fms, green mud,
Feb. 9, 1888

2798, Pearl Islands, Panama, 08°10’30’’N, 78°50’30’’W, 18 fms, gray sand, bro-
ken shells, Mar. 5, 1888

2799, Bay of Panama, 08°44'00’’N, 79°09’00’’ W, 29.5 fms, green mud, Mar. 6, 1888

2824, Greater Bay of La Paz, Gulf of California, 24°22’30’’N, 110°19’30’’ W,
8 fms, broken shells, Apr. 30, 1888

2831, Magdalena Bay, Baja California, 24°32’00’’N, 111°59’00’’W, 12 fms,
fine gray sand, May 2, 1888

2834, near San Juanico Bay, Baja California, 26°14’00’’N, 113°13’00’’ W, 48 fms,
yellow mud, May 3, 1888

2835, near Punta Abreojos, Baja California, 26°42’30’’N, 113°34’15’’W, 5.5 fms,
green mud, May 4, 1888

2837, Cedros Island, Baja California, 28°10’00’’N, 115°09’45’’W, 23 fms, fine
sand, May 5, 1888

2911, Cortes Bank, Calif., 32°27'30’’N, 119°05’00’’ W, 60 fms, rock, sand, Jan. 16,
1889

2922, Cortes Bank, Calif., 32°27'15’’N, 119°05’15’’W, 47 fms, fine gray sand,
Jan. 17, 1889

2969, Santa Barbara Channel, Calif., 34°20’40’’N, 119°37'45’’W, 26 fms, gray
sand, pebbles, stones, Feb. 11, 1889

3138, Monterey Bay, Calif., 36°55’30’’N, 122°02’00’’W, 19 fms, fine sand,
mud, stones, Mar. 15, 1890

3187, near Pt. Sur, Calif., 36°14’00’’N, 121°58’40’’W, 298 fms, yellow sand,
mud, Apr. 3, 1890

3195, near Morro Bay, Calif., 35°14’00’’N, 121°07’00’’W, 252 fms, green mud,
Apr. 5, 1890

3566, San Diego Bay, Calif., 3 fms, fine sand, broken shells, Mar. 19, 1894

3698, Manazuru Zaki, N 8°, W 4.5 m, off Honshu Island, Japan, 153 fms, green
mud, vol. a. s.; May 5, 1900

3702, Seno Umi, N 13°, W 1.5 m, off Honshu, Japan, 31-41 fms, volcanic mud,
sand, rock, May 7, 1900

3708, Ose Zaki, S 55°, W 2.25 m, off Honshu Island, Japan, 60-70 fms, green
mud, volcanic sand, May 8, 1900

3716, Ose Zaki, S 36°, W 0.8 m, off Honshu Island, Japan, 65-125 fms, volcanic
sand, shells, rock, May 11, 1900

3738, Ent. Port Heda, N 84°, E 1.2 m, off Honshu Island, Japan, 167 fms, stiff
blue mud, May 17, 1900

3739, Ose Zaki, S$ 25°, W 0.25 m, off Honshu Island, Japan, 55-65 fms, volcanic
sands, shells, rock, May 17, 1900

3767, Oboro Saki, N 67°, E 2.3 m, off Honshu Island, Japan, 14-18 fms, gray
sand, June 5, 1900

3769, Nagane Saki, N 55°, E 5.3 m, off Honshu Island, Japan, 40-42 fms, green
mud, sand, June 5, 1900

3771, Doumiki Saki, N 19°, W 4.5 m, off Honshu Island, Japan, 61 fms, green
mud, sand, June 5, 1900

NO. 3576 AMPELISCIDAE—BARNARD 3

3809, Honolulu Light, N 28°, E 2’, Hawaii, 51-125 fms, fine coral sand, black

specks, Mar. 27, 1902

_ 4322, Soledad Hill, Point La Jolla, Calif., S 34°, E 3.2 miles, 110-199 fms, green

mud, shells, Mar. 7, 1904

5091, Joga Shima Lt., N 15°, W 4.2 miles, 35°04’10’’N, 139°38’12’’E, Uraga
Strait, Japan, entrance to Gulf of Tokoyo, 197 fms, green mud, coarse black
sand, pebbles, Oct. 26, 1906

Velero IIT Station Recoids

| 221, off Gorgona Island, Colombia, 03°01’25’’N, 78°10’00’’W, 20 fms, rock,

shell, Feb. 12, 1934

: 224, off Gorgona Island, Colombia, 02°59’45’’N, 78°13’20’’W, 10 fms, gravel,

shell, Feb. 12, 1934

| 257, off South Viradores Islands, Port Culebra, Costa Rica, 10°35’00’’N,

85°43/15’’ W, 10 fms, sand, shells, Feb. 25, 1934

| 268, north of White Friars, Mexico, 17°31’30’’N, 101°29’27’’W, 25 fms, coarse

sand, Mar. 3, 1934

420, off Port Utria, Colombia, 05°58’00’’N, 77°21’15’’W, 35 fms, mud, sand,
Jan. 25, 1935

460, Playa Blanca, Costa Rica, 10°56’00’’N, 85°52’50’’ W, 3-5 fms, sand, shells,
Feb. 8, 1935

562, east of San Esteban Island, Gulf of California, 28°41’25’’N, 112°32’15’’W,
20-70 fms, sand, rock, Mar. 10, 1936

Collected by Dr. Waldo L. Schmitt

1, Salinas, Ecuador, littoral, Sept. 12, 1926
10, just north of Paita, Peru, dredged off coast, Oct. 7, 1926

24, off passage between San Lorenzo and adjacent island, Callao Bay, Peru,

dredged with scallop trawl, Nov. 7, 1926

Collected by Dr. E. F. Ricketts, Pacific Biological Laboratories

24-10, Pacific Grove, Calif., channel, rock, 70 fms, Apr. 13 1928
28-5, Pacific Grove, Calif., channel, rock, 80 fms Apr. 18, 1928
125-1, Monterey Bay, Calif., mud, 65 fms, June 22, 1930

125-2, Monterey Bay, Calif., rocks, 80 fms, June 27, 1930

Miscellaneous

Hemphill 5733, San Diego, Calif.
Hilton A—4, Laguna Beach, Calif., Aug. 5, 1918

Ampeliscidae

Ampelisca brevisimulata J. L. Barnard
Ampelisca brevisimulata J. L. Barnard, 1954a, pp. 33-35, pls. 23-24; 1954b, p. 7;
1964a, p. 212.
Material: Albatross 2837 (2).
Record: Cedros Island, Baja California, 23 fms.
Distribution: Eastern Pacific Ocean from Monterey Bay, Calif.,

to Panama, 11-172 m; Caribbean Sea off Colombia and Venezuela,

9-38 m.

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121 |

Ampelisca compressa Holmes

Ampelisca compressa Holmes, 1905, pp. 480-481, 1 fig —Kunkel, 1918, p. 66.—
J. L. Barnard, 1960, pp. 31-32; 1964b, p. 101, chart 4; 1964a, p. 213.
Ampelisca vera J. L. Barnard, 1954a, pp. 23-26, pls. 14-16; 1954b, p. 3, pl. 1,
figs. k-1.
Material: Albatross 2834 (3), 2835 (5), 2837 (6).
Records: Near San Juanico Bay, Baja California, 48 fms; near
Punta Abreojos, Baja California, 5.5 fms; Cedros Island, Baja
California, 23 fms.

Distribution: Western Atlantic Ocean; Caribbean Sea; Pacific |

Ocean from Panama to Puget Sound, Wash., 1-266 m.

Ampelisca cristata Holmes

Ampelisca cristata Holmes, 1908, pp. 507-508, figs. 16-17.—J. L. Barnard, 1954a, |
pp. 26-29, pls. 17-18 (includes forma microdentata); 1954b, pp. 3-4, pl. 1, |
figs. a-g (includes forma microdentata); 1959, p. 18 (forma microdentata) ;

1964a, p. 213.

Material: Albatross 2835 (5), 2911 (3), 2922 (1), 3566 (1). Velero |

420 (1).

Records: Near Punta Abreojos, Baja California, 5.5 fms; Cortes
Bank, California, 47-60 fms; San Diego Bay, Calif., 3 fms; off Port
Utria, Colombia, 35 fms.

Distribution: Eastern Pacific Ocean from Tomales Bay, Calif.,
to La Plata Island, Ecuador, 6-152 m; Caribbean Sea, off Venezuela
and Colombia, ca. 9-42 m.

Ampelisca cristoides J. L. Barnard

Ampelisca cristoides J. L. Barnard, 1954a, pp. 29-31, pls. 19-20; 1954b, p. 4,
pl. 1, figs. h-j.

Material: Albatross 2798 (1). Velero 224 (1), 257 (3), 268 (1).

Records: Pearl Islands, Panama, 18 fms; Gorgona Island, Colom-
bia, 10 fms; South Viradores Islands, Costa Rica, 10 fms; N of White
Friars, Mexico, 25 fms.

Distribution: Eastern Pacific Ocean from Thurloe Head, Baja
California, to Gorgona Island, Colombia, ca. 9-73 m; Caribbean Sea
off north coast of Colombia, ca. 24-40 m.

Ampelisca eoa Gurjanova

Ampelisca eoa Gurjanova, 1951, pp. 313-314, fig. 178.—J. L. Barnard, 1960, p. 25.
Ampelisca catalinensis J. L. Barnard, 1954a, pp. 7-9, pls. 1-2.

Material: Albatross 3698 (8), 3738 (1).

Records: Honshu Island, Japan 153-167 fms.

Distribution: Bering Sea, 1000 m; Honshu Island, Japan, 280-
330 m; offshore basins of southern California, 421-1, 833 m.

NO. 2576 AMPELISCIDAE—BARNARD 5

Ampelisca eschrichtii Krgyer

FicureE lo

Ampelisca Eschrichtii Krdéyer, 1842, p. 155.—Sars, 1895, pp. 174-176, pl. 61,
fig. 1.—Stephensen, 1925, pp. 139-141.

Ampelisca eschrichtii—Stebbing, 1906, p. 100.—Shoemaker, 1930, pp. 27-28;
1931, pp. 9-10.

Ampelisca eschrichti.—Stephensen, 1933, pp. 23-24, fig. 9 (map); 1935, pp. 121-
123.

Ampelisca eschrichti eschrichtt.—Gurjanova, 1955, p. 170, fig. 169.

The shape of the head and article 2 of pereopod 5 distinguish this
species from A. maorocephala Liljeborg. The obliquely truncated
ventral margin of pereopod 5 is a contrast to the transversely trun-
cated margin in A. macrocephala (see Stephensen, 1935). In A.
eschrichtia the ventrolateral corneal lens occurs posterior to the anter-
oventral cephalic corner, and the ventral cephalic margin posterior
to the lens is straight or convex, not excavate as in A. macrocephala.
The ventrolateral lens of A. macrocephala forms the corner of the head.
The presence or absence of a long distal spine on the outer ramus of
uropod 2 is an invalid character in distinguishing the two species,
as both species have the spine (contra J. L. Barnard, 1954a, key).

The Pacific specimens at hand seemed so unlike Sars’ (1895) draw-
ings of A. eschrichtit that the materials were compared with similar
large specimens (20-27 mm) from the western Atlantic Ocean in
Smithsonian collections. These large individuals differ materially
from Sars’ drawings in several conspicuous characters: the shortened
uropod 1; the deeper recessment of the ventral pair of corneal lenses;
and the greater convexity of the ventral cephalic margin posterior to
the lenses. Pacific and some western Atlantic materials are com-
parable in these characters. A specimen from Bedford Basin, Halifax,
Nova Scotia (reported in Shoemaker, 1931), has uropod 1 reaching only
halfway along the rami of uropod 2; specimens from Hebron, Labrador,
1908, 75 fms, have uropod 1 reaching two-thirds along the rami of
uropod 2; and specimens from Ungava Bay (1883, USNM 9413) have
uropod 1 extending to the end of the rami of uropod 2 as shown by
Sars (1895). The specimens from Ungava Bay are remarkable also
for the development of a high dorsal crest on the posterior pereonal
and anterior pleonal segments.

Gurjanova (1955) has figures of A. eschrichtit eschrichtit and her
A. e. pacifica with uropod 1 extending about two-thirds along the rami
of uropod 2; A. e. pacifica has the indented ventral corneal lens and
slightly tumid ventral cephalic margin. Specimens at hand have
these characters more exaggerated than previously shown. A repre-
sentative head is given in figure lo. The Pacific specimens are justi-
fiably identified as A. eschrichtii, sensu lato, and this extends the range

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

of the species in the Pacific as far south as Monterey Bay, California.
That may be the southern limit because in the time since my (1954a)
paper I have examined more than 600 samples and tens of thousands
of specimens of Ampelisca from southern California, 200 miles south
of Monterey, and have never found a specimen of A. eschrichtii (paper
in preparation). Samples extended from depths of 4 to 1500 m.

There can be no confusion of A. eschrichtit with the southern
Californian A. indentata which has a similar head, because the latter
has a large lobe on article 4 of pereopod 5, a strongly saddle-shaped
dorsal carina on urosomite 4 and virtually no tooth on pleonal epimeron
3, all in contradistinction to A. eschrichtii.

Material: Albatross 3187 (9), 3195 (29).

Records: Near Pt. Sur and near Morro Bay, Calif., 252-298 fms.

Distribution: Circumpolar in the northern hemisphere, with known
southern limits as follows: The deep sea (1375 m) off Atlantic France
(SW of Ireland); sublittoral depths of Georges Bank, east of Massa-
chusetts (41°25’N); bathyal depths off Morro Bay, California; and
“Japan Sea” (Gurjanova, 1951), ca. 10-800 m.

Ampelisca furcigera Bulycheva

Ampelisca furcigera Bulycheva, 1936, pp. 242-244, figs. 1-3.—Gurjanova, 1938,
p. 256, fig. 4; 1951, pp. 314-316, fig. 180.—J. L. Barnard, 1960, pp. 26-27,
fig. 6.
Material: Albatross 3698 (1), 3739 (1), 5091 (1).
Records: Honshu Island, Japan 65-153 fms; Uraga Strait, Gulf of
Tokyo, Japan, 197 fms.
Distribution: Bering and Okhotsk Seas, Japan, 60-361 m; eastern
Pacific Ocean in offshore basins of southern California, 212-386 m.

Ampelisca hancocki J. L. Barnard
Ampelisca hancocki J. L. Barnard, 1954a, pp. 37-38, pl. 26; 1964a, p. 213.

Material: Pacific Biological Laboratories 125-1 (1).

Records: Monterey Bay, Calif., 65 fms.

Distribution: Eastern Pacific Ocean from Monterey Bay, Calif.,
to Costa Rica, 9-157 m.

Ampelisca indentata J. L. Barnard

Ampelisca indentata J. L. Barnard, 1954a, pp. 43-45, pl. 30; 1964a, pp. 213-214.

Material: Albatross 2837 (1).

Record: Cedros Island, Baja California, 23 fms.

Distribution: Eastern Pacific Ocean from about Pt. Conception,
Calif., into the Gulf of California, 33-98 m.

NO. 3576 AMPELISCIDAE—BARNARD 7

Ampelisca lobata Holmes

Ampelisca lobata Holmes, 1908, pp. 517-518, fig. 25.—Shoemaker, 1921, p. 99;
1941, p. 187; 1942, p. 7.—J. L. Barnard, 1954a, pp. 11-14, pls. 5-6; 1954b,
p. 2; 1964a, p. 214.

Ampelisca articulata Stout, 1913, pp. 639-640.

Material: Velero 562 (1). Schmitt 1 (5). Pacific Biological
Laboratories 125-2 (4). Hemphill 5733 (6). Hilton A-4 (1).

Records: San Esteban Island, Baja California, 20-70 fms; Salinas,
Ecuador, littoral; Monterey Bay, Calif., rocks, 80 fms.; San Diego
Bay, Calif.; Laguna Beach, Calif.

Distribution: Eastern Pacific Ocean from Monterey Bay, Calif.,
to Lobos de Afueras Islands, Peru, and the Galapagos Islands, 0-183 m
but rare in depths exceeding 30 m; Caribbean Sea, off Colombia,
Aruba, and Barbados Islands, 9-70 m.

Ampelisca mexicana J. L. Barnard

-Ampelisca mexicana J. L. Barnard, 1954a, pp. 45-46, pls. 31-32; 1954b, p. 7;
1964a, p. 214.
Material: Albatross 2799 (1).
Record: Bay of Panama, 29.5 fms.
Distribution: Eastern Pacific Ocean from Punta Canoas, Baja
California, to Independencia Bay, Peru, 9-73 m; Caribbean Sea:
Caledonia, Panama, ca. 18 m.

Ampelisca miharaensis Nagata

Ampelisca miharaensis Nagata, 1959, pp. 70-73, figs. 3-5; 1960, p. 168; 1965,
pp. 152-153, fig. 4 (subfigs. 2-4).
Material: Albatross 3698 (1), 3702 (1), 3708 (2).
Record: Honshu Island, Japan, 41-153 fms.
Distribution: Japan, 2—280 m.

Ampelisca milleri J. L. Barnard
Ampelisca milleri J. L. Barnard, 1954a, pp. 9-11, pls. 3-4.—Jones, 1961, pp.
253-254.—J. L. Barnard, 1964a, p. 215.
Material: Pacific Biological Laboratories 125-1 (1).
Record: Monterey Bay, Calif., 65 fms.
Distribution: Eastern Pacific Ocean from San Francisco Bay,
Calif., to Ecuador, and the Galapagos Islands, 0-187 m.
Ampelisca misakiensis Dahl
Ampelisca misakiensis Dahl, 1944, pp. 6-9, figs. 4-6—Nagata, 1965, p. 152.
fig. 5 (subfigs. 3-6).

Material: Albatross 3702 (1), 3708 (1).
Record: Honshu Island, Japan, 31-60 fms.
Distribution: Warm-temperate Japan, 30-110 m.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ampelisca pugetica Stimpson

Ampelisca pugetica Stimpson, 1864, pp. 158-159.—J. L. Barnard, 1954a, pp.
49-51, pls. 35-36 (includes forma macrodentata); 1954b, p. 7; 1960, p. 31,
fig. 9; 1964a, p. 215.
Ampelisca californica Holmes, 1908, pp. 513-515, fig. 23.
Ampelisca gnathia J. L. Barnard, 1954a, pp. 46-48, pls. 33-34.
Material: Albatross 2831 (1), 2837 (5), 4322 (1).
Records: Magdalena Bay, Baja California, 12 fms; Cedros Island
Baja California, 23 fms; La Jolla, Calif., 110-199 fms.
Distribution: Eastern Pacific Ocean from Puget Sound, Wash.,
to Todos Santos Island, Baja California (pugetica pugetica), from Pt.
Fermin, Calif., to Peru and the Galapagos Islands (pugetica macro-
dentata), 0-183-++ m; Caribbean Sea off Venezuela, Colombia, and
Aruba, 24-42 m.

?

Ampelisca romigi J. L. Barnard

Ampelisca romigi J. L. Barnard, 1954a, pp. 18-20, pls. 10-11; 1954b, p. 3; 1960,
p. 34; 1964a, pp. 215-216.
Ampelisca isocornea J. L. Barnard, 1954a, pp. 20-21, pl. 12.

Material: Albatross 2824 (1). Pacific Biological Laboratories
24-10 (1), 28-5 (1).

Records: Greater Bay of La Paz, Gulf of California, 8 fms; Pacific
Grove, Monterey Bay, Calif., 70-80 fms.

Distribution: Eastern Pacific Ocean from Monterey Bay, Callif.,
to Santa Elena Bay, Ecuador, 3-504 m, probably inhabiting only
very coarse sediments (rubble, gravel) and thus rarely occurring in
shallow water of high latitudes because of absence of substrate;
Caribbean Sea, north shore of Colombia and Aruba, 24-42 m.

Ampelisca schellenbergi Shoemaker

Figures la—m

Ampelisca schellenbergi Shoemaker, 1933, pp. 3-5, fig. 2; 1942, p. 9—J. L. Barnard,
1954a, pp. 14-16, pls. 7-8; 1954b, p. 2.

Material: Albatross 3809 (86). Velero 221 (10), 224 (1), 562 (1).
Schmitt 1 (5). Pacific Biological Laboratories 24 (8).

Records: Off Honolulu, Hawaii, 51-125 fms; Gorgona Island,
Colombia, 20 fms (2 records); San Esteban Island, Baja California,
20-70 fms; Salinas, Ecuador, littoral; Pacific Grove, Calif., 70 fms,
channel.

Notes on Hawaiian material: Antenna 1 variable, in the largest
specimen scarcely exceeding article 4 of antenna 2 peduncle, in other
smaller females reaching nearly three-fourths along article 5, in most
young males reaching halfway or fully along article 5 and occasionally
exceeding it slightly; antenna 2 in female approximately three-fourths
as long as body, in young males as long as body; young male antennae

NO. 3576 AMPELISCIDAE—BARNARD 9

stouter than in female, with stouter flagellar articles; base of flagellum
of antenna 1 slightly more setose, but young males lacking setal

Ficure 1.—Ampelisca schellenbergi Shoemaker, Albatross 3809, female, 7.6 mm: a, head;
b-f, pereopods 1-5; g, telson; h-7, uropods 1-3; k, /, coxae 1, 2; m, continuation of antenna
2 from fig. a; n, pleonites 2-6, left lateral (5-6 coalesced). Ampelisca eschrichtit Krgyer,
female, 20.0 mm, Albatross 3195: 0, head.

tufts typical of adult ampeliscas (no fully adult males at hand);
dactyl of pereopod 3 with 3 accessory spines but only 1 on pereopod
225-669 672

10 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

4; submarginal medial row of setae on posteroventral edge of article
2 on pereopod 5 more densely packed than in American population;
notch and distal end of article 5 with slight ochraceous bulbosities
appearing almost lenslike; posterior edges of articles 6-7 with row of
small pits similar to stridulating ridges; coxae 1-3 each with 1 small
posteroventral slit; peduncle of uropods 1 and 2 without ventro-
lateral setules occurring on continental American specimens.

Remarks: The members of this one population are consistent in
the condition of their third uropods as reflected by the accompanying
illustration, unlike the variation shown for A. schellenbergi by J. L.
Barnard (1954a), and none of the young males has any sexual
differentiation on uropod 3.

The Hawaiian population of this species has been compared with
Caribbean samples used by Shoemaker in his description of the
species and with samples from the eastern Pacific Ocean described
by J. L. Barnard (1954a). The scabrous condition of pereopods
3-4 noted in the Hawaiian specimens and the ornamental pits of
pereopod 5 also occur in the others. Individuals from all 3 areas
have a few serrate spines in the distal groups on article 5 of pereopods
34.

Distribution: Hawaiian Islands, 93-229” m; eastern Pacific Ocean
from Laguna Beach, Calif., to Lorenzo Island, Peru, 0—‘‘128”’ m, but
probably rare in depths esceeding 40 m; western Atlantic Ocean from
Florida, Yucatan, and Panama, 0-46 m.

Ampelisca shoemakeri J. L. Barnard
Ampelisca shoemakeri J. L. Barnard, 1954a, pp. 39-40, pls. 27-28; 1964a, p. 216.

Material: Albatross 2837 (29). Schmitt 10 (2).

Records: Cedros Island, Baja California, 23 fms; north of Paita,
Peru.

Distribution: Eastern Pacific Ocean from Cedros Island, Baja
California, to Paita, Peru, 7-76 m.

Byblis albatrossae, new species
FIGURE 2

Diagnosis of female: Head with distinct rostrum about one-third
as long as article 1 of antenna 1, anterior cephalic margin with strong
angular protrusion, anteroventral cephalic margin deeply excavate
for reception of antenna 2; ventral pair of corneal lenses situated at
and forming rounded anteroventral cephalic angle, arranged so as to
point obliquely anteroventrally, dorsal pair of lenses enormous, about
1.5 times as large as ventral pair, protruding strongly; antenna 1
nearly as long as peduncle of antenna 2, article 2 more than twice as
long as article 1 and more than half as long as article 4 of antenna 2;

I EDEIEEE

NO. 3576 AMPELISCIDAE—BARNARD 11

Ficure 2.—Byblis albatrossae, new species, female, holotype, 15.0 mm, Albatross 3738:
a, head; b, c, coxae 1, 2; d-h, pereopods 1-5; 7 ,telson; 7, outline of maxilla 2, minus setae;
k-m, uropods 1-3, latter enlarged.

(2 PROCEEDINGS OF THE NATIONAL MUSEUM von. 121 |

article 4 of antenna 2 longer than peduncle of antenna 1 and much |
longer than article 5 of antenna 2, flagellum 4 times as long as article |
5 of peduncle; anterior coxae unserrate ventrally, shallow, pointing
very strongly anteroventrad, coxae 3 and 4 progressively more trun-
cated ventrally, coxa 4 with long, flat ventral margin and narrow
posterior tooth; pereopods 1-2 scarcely disproportionate in size,
dactyls slightly longer or subequal in length to their respective sixth
articles, pereopod 2 more strongly setose than pereopod 1; pereopods
3-4 without special modifications, posterior bilobation of article 2
on pereopod 3 especially symmetrical for genus; article 2 of pereopod
5 with typical overall shape but ventral margin evenly rounded,
article 6 about 80 percent as long as article 5, article 7 slightly more
than half as long as article 6; uropod 1 extending to end of uropod 2
but with disproportionately short peduncle and long rami, distolateral
apex of peduncle with long, slender spine, outer ramus longer than
inner and lacking spines, inner ramus with basal ridge for locking
outer ramus; uropod 2 with hooked distolateral cusp on peduncle,
outer ramus with more than twice as many marginal spines as inner
ramus; uropod 3 typically forcipiform, apposing margins of rami
strongly serrate; telson of medium length, cleft about one-third.

Remarks: Pleonal epimera are similar to those of Byblis affinis (in
Sars, 1895, pl. 65). Mouthparts are similar to those of B. gavmardi
(in Sars, 1895, pl. 65), but article 2 of the mandibular palp is expanded
proximally in asymmetrical fashion, each inner plate of the maxillipeds
has 2 spine-teeth, and the outer plate of maxilla 2 is slightly broader
and distally expanded.

Male: Two specimens, 15 and 13 mm long, have article 5 of antenna
2 nearly as long as article 4, and antenna 1 is equal to or exceeds the
peduncle of antenna 2 by the length of 2 flagellar articles.

Holotype: USNM 111268, female, 15.0 mm.

Type locality: Albatross 3738, entrance to Port Heda, off Honshu
Island, Japan, 167 fms, May 17, 1900.

Material: Albatross 3698 (3), 3738 (3).

Records: Manazuru Zaki and Port Heda, Honshu Island, Japan,
153-167 fms.

Relationship: In some ways this species resembles B. veleronis
J. L. Barnard (1954a), but the disproportionate corneal lenses, the
angular protrusion on the anterior margin of the head, and the shape
of coxa 4 distinguish B. albatrossae. The shape of the anterior 4 pairs
of coxae and the first antennal length relate this species to one being
described by M. Imbach from Viet Nam, but numerous other char-
acters of that species distinguish it: serrate anterior coxae, equal
ocular lenses, straight anterior cephalic margin, more deeply cleft
telson, and the form of article 2 on pereopod 3.

NO. 3576 AMPELISCIDAE—BARNARD 13

The distinct rostrum of B. albatrossae prompts comparison with
Haploops securiger K. H. Barnard (1932) which I transfer to Byblis.
That species also has a distinct but much longer rostrum, and its
ventral pair of corneal lenses is situated beneath the head and con-
cealed from lateral view.

Distribution: Honshu Island, Japan, 280-329 m.

Byblis ampelisciformis, new species
FIGURE 3

Diagnosis of male: Rostrum vestigial, anterior cephalic margin
lacking protrusion, anteroventral cephalic margin sinuously excavate
for reception of antenna 2, strongly oblique; two pairs of corneal
lenses of medium and subequal size, ventral pair situated at and
forming rounded anteroventral angle, pointing anterolaterally;
antenna 2 as long as body, antenna 1 reaching nearly three-fourths
along full length of antenna 2, article 2 of peduncle 2.5 times as long
as article 1 and nearly as long as article 4 of peduncle on antenna 2;
anterior coxae serrate ventrally, coxa 4 of medium width, slightly
truncate ventrally, posterior tooth of medium size; pereopods 1-2
scarcely disproportionate in size, dactyls slightly longer than their
respective sixth articles; pereopods 3-4 without special modifications;
article 2 of pereopod 5 with typical overall shape, article 6 about 67
percent as long as article 5, article 7 about 40 percent as long as
article 6; uropods 1 and 2 of normal proportions, distolateral end of
uropod 1 peduncle with stout spine, of uropod 2 with hooked cusp,
spines of peduncles and rami remarkably short and stout; rami of
uropod 8 furciform, short, asetose, medial margin of inner ramus
lined with short spines, ventromedial margin of outer ramus bearing
several scales, typical serrations absent; telson short, very broad,
cleft halfway; urosome slightly miniaturized in comparison to other
species of genus.

Remarks: Mouthparts are similar to those of Byblis gaimardi
(Krgyer) (in Sars, 1895, pl. 64), but the outer plate of maxilla 2 is
expanded distally very slightly and article 2 of the mandibular palp
is asymmetrically expanded. Pleonal epimera are rounded posteriorly
and resemble those of B. crassicornis Metzger (in Sars, 1895, pl. 66).
The medial surface of article 2 of pereopod 5 is densely setose.

Female: Similar to male.

Holotype: USNM 111271, male, 12.0 mm.

Type locality: Albatross 3708, Ose Zaki, off Honshu Island, Japan,
60-70 fms, May 8, 1900.

Material: Albatross 3708 (3), 3716 (1).

Records: Ose Zaki, Honshu Island, Japan, 70-125 fms.

Relationship: This species resembles B. veleronis J. L. Barnard,

14 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

B. japonicus Dahl, and B. erythrops Sars in the following character-
istics: Corneal lenses present, lower pair visible laterally, pereopod
2 similar to pereopod 1, telson cleft a third or more, antenna 1 subequal

Ficure 3.—Byblis ampelisciformis, new species, male, holotype, 12.0 mm, Albatross 3708:
a, head; b-f, pereopods 1-5; g, h, coxae 1, 2; 7, telson; 7-1, uropods 1-3, latter enlarged.

to antenna 2 or greatly exceeding peduncle of antenna 2, pereopod 4
lacking a cusp on article 2. The new species differs from the others
in its unusual third uropods with short blunt spines and furciform
appearance. Indeed, few other species of Byblis are known to have

NO. 3576 AMPELISCIDAE—BARNARD t5

these uropods which are suggestive of the following species of Am-
pelisca: lobata Holmes (females only), bidentata Schellenberg, sca-
bripes Walker, and excavata K. H. Barnard. From the literature,
those species appearing to resemble B. ampelisciformis in their third
uropods are B. antarctica Schellenberg (1931) which lacks corneal
lenses and has a large distal protrusion on article 5 of pereopod 3,
and B. subantarctica Schellenberg (1931) which has a very distinctive
pereopod 5, no ventral lenses, and which is transferred to Ampelisca.
Distribution: Honshu Island, Japan, 128-228 m.

Byblis orientalis, new species
Figure 4

Diagnosis of male: Rostrum vestigial, anterior cephalic margin with
small protrusion, anteroventral cephalic margin with short, regular
excavation for reception of antenna 2; two pairs of corneal lenses of
medium and subequal size, ventral pair pointing anterolaterally,
situated at and forming rounded anteroventral cephalic angle; antenna
2 as long as body, antenna 1 reaching about three-fourths along article
5 of antenna 2, article 2 of peduncle almost twice as long as article 1
and slightly more than one-third as long as peduncular article 4 of
antenna 2; anterior coxae unserrate ventrally, coxa 4 of medium
width, not truncate ventrally but posteroventral margin beveled and
posterior tooth therefore very slender; pereopods 1-2 scarcely dis-
proportionate in size, dactyls shorter than their respective sixth
articles; pereopods 3-4 without special modifications, article 2 of
pereopod 5 with typical overall shape, but neither as strongly expanded
nor as ventrally extended as in many species of Byblis, articles 5 and
6 equal in length, article 7 about half as long as article 6; uropods 1
and 2 of normal proportions, distolateral end of uropod 1 peduncle
with stout spine, of uropod 2 with weakly hooked cusp, spines of
peduncles and rami of normal size; uropod 3 typically forcipiform,
apposing margins of rami weakly serrate; telson of medium length,
cleft about one-third.

Remarks: Mouthparts are like those of B. gaimardi (Krgyer) (in
Sars, 1895, pl. 64) except for the expanded article 2 of the mandibular
palp and the stouter spines of the outer plates of the maxillipeds.
Article 2 of pereopod 5 is covered medially with setae. Male antennae
lack setal tufts on the peduncles. The holotype has the best developed
dorsal subcrest on urosomite 1 of any of the specimens. The anterior
cephalic protrusion is also best developed on the figured holotype.

Female: Antenna 1 extending along antenna 2 only about 2 flagellar
articles beyond article 4.

Holotype: USNM 111274, male, 11.0 mm.

16 PROCEEDINGS OF THE NATIONAL MUSEUM Vou. 121

Type locality: Albatross 3769, Nagane Saki, off Honshu Island,
Japan, 40-42 fms, June 5, 1900.

Ficure 4.—Byblis orientalis, new species, male, holotype, 11.0 mm, Albatross 3769: a, head;
b-f, pereopods 1-5; g, h, coxae 1, 2; i, telson; j-I, uropods 1-3, latter enlarged; m, pleonites
2-6, left to right (5-6 coalesced).

Material: Albatross 3767 (2), 3769 (9), 3771 (1 specimen, the

largest, 16 mm).

:

NO. 3576 AMPELISCIDAE—BARNARD V7

Records: Oboro Saki; Nagane Saki; and Doumiki Saki, Honshu
Island, Japan, 18-61 fms.

Relationship: Differing from B. rhinoceros Pirlot (1936) to which
it is closely related in view of antenna 1, head, telson, uropod 3,
pereopod 5 articles 3-7, and uropods 1 and 2, by the slightly more
quadriform posterodistal corner of article 2 of pereopod 5, and by the
considerably shorter first antenna of the female. In many respects
this species also resembles B. crenulata Pirlot (1936) which is close to
if not identical with B. daleyi Giles (Pirlot’s, 1936, identification of
daleyi did not account for the extremely long second article of antenna
1), but the head and eyes of B. crenulata differ, the lower lens pointing
more ventrally, the anterior edge lacking a small protrusion, and the
ventral margin of the head apparently not being strongly excavate
for the attachment of antenna 2. Article 2 of antenna 1 is subequal
to article 1 in contrast to B. orientalis and the anterior coxae are
serrate ventrally. In addition, article 7 of pereopod 5 is much shorter
in B. crenulata than it is in B. orientalis.

The new species is also related to Pirlot’s (1936) identification of
B. daleyi, but article 2 of antenna 1 is much shorter than in that
individual; however, one aberrant specimen of B. orientalis in sample
3138 has left and right antennae dissimilar, the left with article 2
more than twice as long as article 1, the right with article 2 scarcely
longer than article 1.

This species also resembles a new species to be described by M.
Imbach from the South China Sea but differs from it by the sparsity
of spines on the ‘‘posterior”’ edge of article 6 on pereopod 3, by the
less attenuated and asymmetrical lobe of article 2 on pereopod 5, by
the slightly shorter first antennae, the slightly deeper cleft of the
telson, the stronger posterior cusp of coxa 4, and the thinness of the
setae on the outer ramus of uropod 3. Those two species may be
races of a common stem.

Distribution: Honshu Island, Japan, 33-112 m.

Haploops spinosa Shoemaker

Haploops spinosa Shoemaker, 1931, pp. 13-18, figs. 5, 6.
Haploops tubicola.—J. L. Barnard, 1960, p. 35. [Not Liljeborg.]

Barnard overlooked the row of spines on article 3 of pereopod 5 in
his misidentification of 1960.

Material: Albatross 3698 (1).

Record: Honshu Island, Japan, 153 fms.

Distribution: Western Atlantic Ocean, Bay of Fundy, and Nova
Scotia, 22-2300 m; eastern Pacific Ocean, near Pt. Conception, Calif.,
117-171 m; Honshu Island, Japan, 250 m.

18 PROCEEDINGS OF THE NATIONAL MUSEUM VoL, 121

Literature Cited

BARNARD, J. L.

1954a. Amphipoda of the family Ampeliscidae collected in the eastern Pacific
Ocean by the Velero III and Velero IV. Allan Hancock Pacific
Expeditions, vol. 18, pp. 1-137, pls. 1-38.

1954b. Amphipoda of the family Ampeliscidae collected by the Velero III
in the Caribbean Sea. Allan Hancock Atlantic Expedition, Rept. 7,
pp. 1-13, pls. 1-2.

1959. Estuarine Amphipoda. Jn Barnard and Reish, Ecology of Amphi-
poda and Polychaeta of Newport Bay, California. Allan Hancock
Found. Publ. Occ. Pap. 21, pp. 13-69, pls. 1-14.

1960. New bathyal and sublittoral ampeliscid amphipods from California,
with an illustrated key to Ampelisca. Pacific Nat., vol. 1, no. 16,
pp. 1-386, figs. 1-11.

1964a. Los anfipodos bentonicos marinos de la costa occidental de Baja
California. Rev. Soc. Mexicana Hist. Nat., vol. 24, pp. 205-274,
figs. 1-11.

1964b. Marine Amphipoda of Bahia de San Quintin, Baja California. Pa-
cific Nat., vol. 4, pp. 55-139, figs. 1-21.

BarNarp, K. H.

1932. Amphipoda. Jn Discovery Reports, vol. 5, pp. 1-326, figs. 1-174,

plod:
BuLycuHeva, A.

1936. New species of Amphipoda from the Japan Sea. Ann. Mag. Nat.

Hist., ser. 10, vol. 18, pp. 242-256, figs. 1-35.
Daut, E.

1944. Amphipoda of the family Ampeliscidae from Professor Sixten Bock’s

expedition to Japan1914. Arkiv Zool., vol. 354A, pp. 1-18, figs. 1-10,
GuRJANOVA, E.

1938. Amphipoda, Gammaroidea of Siaukhu Bay and Sudzukhe Bay
(Japan Sea). Reports of the Japan Sea Hydrobiological Expedi-
tion of the Zoological Institute of the Academy of Sciences of the
USSR in 1934, pt. 1, pp. 241-404, figs. 1-59.

1951. Bokoplavy morei SSSR i sopredel’nyx vod (Amphipoda-Gammaridea).
Opred. po Faune SSSR, Izd. Zool. Inst. Akad. Nauk SSSR, vol. 41,
pp. 1-1,031, figs. 1-705. [In Russian.]

1955. Novye vidy bokoplavov (Amphipoda, Gammaridea) iz severnoi
chasti Tixogo Okeana. Trudy Zool. Inst. Akad. Nauk SSSR,
vol. 18, pp. 166-218, figs. 1-23. [In Russian.]

Houmgs, S. J.

1905. The Amphipoda of southern New England. Bull. U.S. Bur. Fisheries,
vol. 24, pp. 459-529, pls. 1-13, numerous text figs. [unnumbered].

1908. The Amphipoda collected by the U.S. Bureau of Fisheries Steamer,
“Albatross,” off the west coast of North America, in 1903 and 1904,
with descriptions of a new family and several new genera and
species. Proc. U.S. Nat. Mus., vol. 35, pp. 489-5438, figs. 1-46.

Jones, M. L.

1961. <A quantitative evaluation of the benthic fauna off Point Richmond,
California. Univ. California Publ. Zodél., vol. 67, pp. 219-320,
figs. 1-30.

NO. 3576 AMPELISCIDAE—BARNARD 19

Kroyer, H.
1842. Une nordiske Slaegter og Arter af Amfipodernes Orden, henhgrende
til Familien Gammarina (Forelgbigt Uddrag af et Stgrre Arbejde).
| Naturh. Tidsskr., vol. 4, pp. 141-166.
Kun«xKEL, B. W.
| 1918. The Arthrostraca of Connecticut. Connecticut Geol. Nat. Hist.
Surv., vol. 6, bull. 26 (Amphipoda), pp. 15-181, figs. 1-55.
| Naaara, K.
1959. Notes on five species of the amphipod genus Ampelisca from the
stomach contents of the triglid fishes. Publ. Seto Mar. Biol. Lab.,
vol. 7, pp. 67-82, pls. 2-11.
1960. Preliminary notes on benthic gammaridean Amphipoda from the
Zostera region of Mihara Bay, Seto Inland Sea, Japan. Publ.
Seto Mar. Biol. Lab., vol. 8, pp. 163-182, figs. 1-2, pls. 13-17.
1965. Studies on marine gammaridean Amphipoda of the Seto Inland Sea, 1.
| Publ. Seto Mar. Biol. Lab., vol. 18, pp. 131-170, figs. 1-15.
Pirtot, J. M.

1936. Les amphipodes de l’expedition du Siboga, Deuxiéme partie: Les
amphipodes gammarides, II. Les amphipodes de la mer profonde, 3:
Addendum et partie générale, III: Les amphipodes littoraux, 1:
Lysianassidae . . . Gammaridae. Monogr. 33e (vol. 127) in
Weber, Siboga-Expeditie, pp. 237-328, figs. 102-146.

Sars, G. O.

1895. Amphipoda. Jn An account of the Crustacea of Norway with short
descriptions and figures of all the species, vol. 1, pp. i-viii + 1-711,
pls. 1-240, suppl. pls. 1-8.

ScHELLENBERG, A.

1931. Gammariden und Caprelliden des Magellangebietes, Siidgeorgiens und
der Westantarktis. Further Zoological Results of the Swedish Ant-
arctic Expedition 1901-1903, vol. 2, no. 6, pp. 1-290, pl. 1, figs.
1-136.

SHOEMAKER, C. R.

1921. Report on the amphipods collected by the Barbados-Antigua Expedi-
tion from the University of Iowa in 1918. Univ. Iowa Studies
Nat. Hist., vol. 9, pp. 99-102.

1925. The Amphipoda collected by the United States Fisheries Steamer
“Albatross” in 1911, chiefly in the Gulf of California. Bull.
American Mus. Nat. Hist., vol. 52, pp. 21-61, figs. 1-26.

1930. The Amphipoda of the Cheticamp Expedition of 1917. Contr.
Canadian Biol. Fish., n.s., vol. 5, no. 10, pp. 1-141, figs. 1-54.

1931. The stegocephalid and ampeliscid amphipod crustaceans of New-
foundland, Nova Scotia, and New Brunswick in the United States
National Museum. Proc. U.S. Nat. Mus., vol. 79, no. 2888, pp.
1-18, figs. 1-6.

1933. Amphipoda from Florida and the West Indies. American Mus.
Nov., no. 598, pp. 1-24, figs. 1-13.

1941. On the names of certain California amphipods. Proc. Biol. Soc.
Washington [D.C.], vol. 54, pp. 187-188.

1942. Amphipod crustaceans collected on the Presidential Cruise of 1938.
Smithsonian Misc. Coll., vol. 101, no. 11, pp. 1-52, figs. 1-17.

Stepsine, T. R. R.

1906. Amphipoda, 1: Gammaridea. Vol. 21 in Das Tierreich, pp. 1-806,

figs. 1-127.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

STEPHENSEN, K.
1925. Crustacea Malacostraca, VI (Amphipoda, II). Jn Danish Ingolf-

Expedition, vol. 3, pp. 101-178, figs. 23-53.
1933. Amphipoda. Jn The Godthaab Expedition, 1928. Medd. om Grgn-
land, vol. 79, no. 7, pp. 1-88, figs. 1-31.
The Amphipoda of N. Norway and Spitsbergen with adjacent waters.

Troms¢ Mus. Skr., vol. 3, pp. 1-140, figs. 1-19.

1935.

Stimpson, W.
1864. Descriptions of new species of marine Invertebrata from Puget Sound,

collected by the naturalists of the North-west Boundary Com-
mission, A. H. Campbell, Esq., Commissioner. Proc. Acad. Nat.
Sci. Philadelphia [vol. 16], 1864, pp. 153-165.
STout,) Vile
1913. Studies in Laguna Amphipoda.
633-659, figs. 1-3.

Zool. Jahrb., vol. 34 (Syst.), pp.

U.S. GOVERNMENT PRINTING OFFICE: 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1966 Number 3577

BREDIN-ARCHBOLD-SMITHSONIAN
BIOLOGICAL SURVEY OF DOMINICA

1. The Echinoids of Dominica!

By Porter M. Kier

Associate Curator, Division of Invertebrate Paleontology

Although the echinoids have been described from most of the larger
islands of the Caribbean, no one has ever reported the echinoids of
Dominica. As a member of the Bredin-Archbold-Smithsonian Bio-
logical Survey of Dominica, I spent April 1964 studying and collecting
echinoids off the island. Underwater diving apparatus was used to
make traverses down to 85-foot depths off most of the promotories
and in most of the bays. Live individuals were found in 8 of the 10
species collected and their living positions are described. Most of the
diving was done in the Caribbean because of the lack of suitable boats
on the Atlantic side of the island.

I thank Miss Maureen Downey of the U.S. National Museum for
identifying the basket stars and crinoids and Mr. Louis R. Purnell
for making the drawings. I wish also to thank William P. Campbell,
who assisted in the collecting.

1 This paper is the first of a series on the faunal studies from the survey that
will appear in the ‘“‘Proceedings of the United States National Museum.”” A com-
panion series on the flora will appear in the ‘‘Contributions of the United States
National Museum.”

E

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

General Description of Coast

The island is of great relief with a high gradient to the shore. Off
the promotories the gradient is so steep that in some places there is
only a narrow belt less than 100 feet wide with depths of less than 85
feet. The lowest gradients were in some of the sandy bays such as
that at Mero, where a depth greater than 85 feet was not encountered
within a half mile of the shore. Two main types of environments were
apparent on the Caribbean side of the island: the rocky-coral areas
and the sand areas.

Rocky-Coral Environment

Off the promontories the bottom is generally rocky with many corals
and sponges (fig. 1; pl. 1: fig. 1). Small sandy patches of approxi-
mately 100 square feet occur within this rock-coral. Usually this

ROCKY—CORAL ENVIRONMENT

Echinometra lucunter (Linnoeus)
Diodema antillarum Philippi
Tripneustes ventricosus (Lamarck)
Eucidoris tribuloides (Lomorck)

\ abundont

| rore

Clypeoster rosoceus (Linnaeus)
Clypeaster subdepressus (Gray) - 10 ft.
Meomo ventricoso (Lamarck)

& 4
in sandy patches — down -2
2 S — 10 85 ft re
Ss Diadema antillarum Philippi 30
down to 65 ft.
sO -40
Coral, sponges
o” encrusting rock -50
- 60
70
80
> Schizoster (Paroster)
“== > _floridiensis Kier ond
Se Gront
“SAND. =
Ficure 1

rocky-coral area continues down to approximately 75 feet, where the
rock ceases, and sand continues downward at a steep grade. Al-
though some of the coral masses are large, most of them are not very
thick and barely cover the rocky substrate. The crinoid Comactinia
echinoptera (Muller) lives commonly in crevices (pl. 1: fig. 1) in the
coral from depths of 15-85 feet and probably deeper.

In the shallow water of the rocky-coral areas, from low tide to 10
feet, the most common echinoid is Diadema antillarum Philippi (pl. 2:
figs. 4, 5) except in areas of strong current, where Echinometra lucunter
(Linnaeus) is more abundant. I saw a few specimens of Hucidaris

naar eae seers feat

eerie

i ta i a

i
j

NO. 3577 ECHINOIDS—KIER 3

tribuloides (Lamarck) and Tripneustes ventricosus (Lamarck) (pl. 2:
fig. 4).

Below 10 feet, Diadema antillarum was the only echinoid seen in
any numbers. The spatangoid Meoma ventricosa (Lamarck) lives in
the sandy patches together with a few individuals of Clypeaster sub-
depressus (Gray) and C. rosaceus (Linnaeus).

Sand Environment

The bottom of the bays are mostly sand or silt except for a narrow
band of gravel at the shoreline (fig. 2). Only one echinoid, Leodza
sexiesperforata (Leske), was found in the shallower parts of these
sandy areas. Between 25 and 65 feet holothurians were encountered

SAND ENVIRONMENT

Seas

Gravel

Leodio sexiesperforata (Leske)
‘ abundant— buried - 10 ft,

<a

Holothurians

- 40
Storfish-— OS - 50
- 60
Patches of sponges with - 70
Meoma ventricosa (Lamarck)
Ficure 2 - 80

frequently and, at one site north of Tarreau Point at 50 feet depth, the
starfish Oreaster reticulatus (Linneaus) was present in great numbers.
In a few of these sandy areas small patches of sponges and a few
specimens of Meoma ventricosa (Lamarck) were present between the
depths of 25 and 50 feet.

Echinoids and Their Localities

species locality
Eucidaris tribuloides (Lamarck)
Tripneustes ventricosus (Lamarck)
Echinometra lucunter (Linnaeus)
Echinometra viridis Agassiz
Diadema antillarum Philippi
Clypeaster subdepressus (Gray)
Clypeaster rosaceus (Linnaeus)
Leodia sexiesperforata (Leske)
Meoma ventricosa (Lamarck)
Schizaster (Paraster) floridiensis Kier and Grant

00 CO

~

0, 18

1
14-16, 18

, 6, 8, 10, 13-17

|
a WwW
me OV

or
IP so
ae
So
ia s
So i"
ew
—_
bo

me DI NT be 00 00

PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

|

|

Woodfor:
Hill Bay

(3}Savane

S

stil

BY mii
CS
:

Massacre

61°30" 61°15!

Ficure 3

NO. 3577 ECHINOIDS—KIER 5

Locality Data

numerals=location of stations on map (fig. 3)

_

Rocky, few corals, sponges, many Diadema antillarum Philippi.

2 Abundant corals, extensive flat area of Porpites, large brain corals,
dropping rapidly from depth of 25 feet. Abundant crinoids, many
Diadema antillarum Philippi, Meoma ventricosa (Lamarck), few (4
dead tests) Clypeaster subdepressus (Gray).

3 Rocky area with few corals, Diadema antillarum Philippi in great numbers,

one Meoma ventricosa (Lamarck) in sandy patch at 50-foot depth.

Two basket stars.

4 Rocky area with few corals down to 60 feet, sand at greater depth.
At 65 to 85 feet many Meoma ventricosa (Lamarck); one Schizaster
(Paraster) floridiensis Kier and Grant at 85 feet.

5 Rocky area with few corals, abundant Diadema antillarum Philippi.

6 Rocky area with many corals, large sandy patches in shallower region

(10’-20’); large sponges, many crinoids and basket stars, abundant

Diadema antillarum Philippi, Meoma ventricosa (Lamarck), one living,

many dead tests of Clypeaster subdepressus (Gray).

7 Sandy area, no coral, many holothurians, small amount of grass, many
conchs, many Leodia sexiesperforata (Leske).
8 Rocky area, many corals, sponges, crinoids, on shallow ledge, 1—2 feet

deep, many Echinometra lucunter (Linnaeus), Diadema_ antillarum
Philippi, and few Tripneustes ventricosus (Lamarck) in deeper waters,
10-60 feet, Diadema antillarum Philippi, Echinometra viridis Agassiz,
and Meoma ventricosa (Lamarck). No rock below 75 feet.

9 Sandy area, no corals, many holothurians, conchs, small amount of grass,
Leodia sexiesperforata (Leske).
10 Rocky area first 25 feet with few corals, many crinoids, Diadema antillarum

Philippi, one specimen Tripneustes ventricosus (Lamarck) (2 feet),
Meoma ventricosa (Lamarck) in sandy patches, few dead test of Clypeaster
rosaceus (Linnaeus), Clypeaster subdepressus (Gray) at 25 feet in sandy
patch. At depths greater than 25 feet, no rock, sandy bottom, small
amount of grass, one T'ripneustes ventricosus at 30 feet.

11 Sandy area, small amount of grass, many Oreaster reticulatus (Linneaus),
two basket stars.
12 Sandy area, Leodia sexiesperferata (Leske) in great numbers at 5-20

feet; at 25 feet, rocky ledge with corals, sponges, crinoids, and Meoma
ventricosa (Lamarck), Diadema antillarum Philippi, at 60 feet sand
again,
13 Rocky area heavy currents, few corals, sea fans, sea whips, Diadema
antillarum Philippi, many dead tests of Clypeaster subdepressus (Gray).
14-16 Echinometra lucunter (Linnaeus), Diadema antillarum Philippi in shallow

water.

IN Rocky area down to 35 feet with corals, sponges, sea whips, Diadema
antillarum Philippi. Below 35 feet sandy with conches and
holothurians.

18 Rocky area, heavy currents, in shallow water many Echinometra lucunter

(Linnaeus), in 10-15 feet on rock platform abundant Tripneustes
ventricosus (Lamarck).

225-667—66——_2

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Discussion of Species

Descriptions, synonyms, and illustrations of the species discussed

below can be found in Mortensen (1928-1951), and Kier and Grant |

(1965).
Eucidaris tribuloides (Lamarck)

Only one specimen of this species was seen on the Caribbean side:
in two feet of water in a crevice between a sponge and rock. Living
with it were many individuals of Echinometra lucunter (Linnaeus)
and Diadema antillarum Philippi.

Tripneustes ventricosus (Lamarck)

PLATES 1 (FIG. 2), 2 (FIG. 4)

Only three specimens were seen in the Caribbean. Two of them
(pl. 2: fig. 4) occurred in just two feet of water on a rock outcrop,
one of them up under a rock. The third specimen was in 30 feet of
water on sandy, grassy bottom. None of the specimens had any weed
or fragments held over its test. Although a thorough search was
made for more specimens, none was found.

On the Atlantic side this species was abundant at Woodford Hill
Bay, where they were found in 10 feet of water (pl. 1: fig. 2). Here
a few fragments of weed were held over each test. In some places
the echinoids occurred in such abundance that their tests were literally
touching each other. Lewis (1958, p. 614) reports that in the Bar-
bados individuals of this species crowd together on the upper surface
of rocks during March and April in order to spawn. Presumably,
since it was April when I saw this concentration in Dominica, these
individuals also were spawning.

Echinometra lucunter (Linnaeus)

This species was found only at a few sites on the Caribbean side,
usually restricted to intertidal zones. It was found only where the
normal steep gradient of the sea floor was interrupted by a rock out-
crop that formed a platform at intertidal depths. Such sites occur
at Massacre, Crabiere Point, and Point Ronde. Here the echinoid
occurs in great abundance with approximately two to four specimens
in every square foot. Normally, the echinoid lives in a hollow that
he presumably has formed in the rock. He is associated with many
sea anemones and much algae. Many of the echinoids are exposed
at low tide to the air. Deeper than eight feet they are rare, with
their greatest concentration at one to two feet below low tide. In

Ficure 1.—Coral mass in 50-foot depth at locality 8; the crinoid Comactinia echinoptera
(Muller) can be seen living in crevices in the coral. Ficure 2.—Tripneustes ventricosus
(Lamarck) in 10 feet of water in dense aggregations, presumably to spawn.

anne

PROC. U.S. NAT. MUS. VOL. 121

KIER—PLATE 1

PROC. U.S. NAT. MUS. VOL. 121 KIER—PLATE 2

NO. 3677 ECHINOIDS—KIER 7

every case they were living in areas of high energy with considerable
wave motion. Two color types were present, the most common
being dark brown but some specimens were much redder. Color
difference is distinct, with no transition between the two types.
Many individuals of Diadema antillarum Philippi and a few of Evuct-
daris tribuloides (Lamarck) and Tripneustes ventricosus (Lamarck)
live with this species.

Echinometra viridis Agassiz

Only one specimen was seen and this one was found living in a
niche in the coral at a depth of 50 feet. A long search was made for
more specimens but none was found.

Diadema antillarum Philippi
Puate 2 (Ftas. 4, 5)

This species is the most abundant of the echinoids living on the
Caribbean side of the island. It occurs most commonly on the
rocky-coral areas and is usually absent in the sandy bays. It was
found in great numbers from low tide level to 65 feet and presumably
lives even deeper. Most specimens live in hollows in the coral or
rock, but some individuals were seen on the sandy patches within
the rocky-coral areas. Although no individuals were ever seen under
attack by fish, many disassociated spines were seen, indicating that
successful attacks had been made. Both small fish and shrimp
were frequently seen living among the spines.

Clypeaster subdepressus (Gray)

Although many dead tests of this species were seen, only one living
individual was found. The species lives buried three or four inches
in the sandy patches in the rocky-coral areas in association with
Meoma ventricosa (Lamarck). It was most common in depths of
10 to 25 feet. The living specimen was found in 50 feet of water
completely buried. Its location was indicated by the presence of
whiter sand (which is usually underneath) on top of the specimen.

pope wea pyre © EST INE bees AE
Ficure 1.—Meoma ventricosa (Lamarck) with fragments of coral and shell pulled up over
the test in 35-foot depth at locality 2; note coarser fragments over madreporite. FIGURE
2.—Meoma ventricosa (Lamarck) in 85-foot depth at locality 4; test covered only by bits
of algae instead of the usual thick covering of shell fragments and pieces of coral; this is
the greatest depth reached during the study and perhaps the echinoid is not covered very
much because of the decreased amount of light at this depth. Ficure 3.—Meoma
ventricosa (Lamarck) in 35-foot depth at locality 6, showing lack of detrital material over
petals and trail made by the echinoid. Ficure 4.—Tripneustes ventricosus (Lamarck)
clinging to rock in 2 feet of water at locality 8; Diadema antillarum Philippi can be seen
to the right. Ficure 5.—Diadema antillarum Philippi in 5 feet of water at locality 13.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Clypeaster rosaceus (Linnaeus)

Three dead tests were found at a depth of 25 feet in one of the sandy
patches in the rocky-coral areas in association with Clypeaster sub-
depressus and Meoma ventricosa (Lamarck).

Leodia sexiesperforata (Leske)

This species was found living on the sandy areas in the bays in
depths from 5 to 20 feet. The animal lives completely buried under
2 to 4 inches of silty sand. The echinoid makes no track in the sand
visible on the upper surface of the sea bottom. In most cases the
ripple marks on the bottom continue uninterrupted directly over
the spot where the echinoid is buried. The only clue to the presence
of living specimens is the occurrence of dead tests at the surface.
In most places it occurs with no vegetation although in some areas
a small amount of eelgrass is present.

Meoma ventricosa (Lamarck)

PLaTE 2 (Fias. 1-3)

Although not abundant, this species was found in sandy patches
in every coral-rock environment from depths of 15 to 85 feet. Pre-
sumably it occurs at greater depth off Dominica but no dives were
made below 85 feet. In other areas in the West Indies it has been
dredged from depths down to 600 feet. It is not conspicuous on the
bottom because it covers itself with sand and fragments of coral or
weed. It would not be correct to say that it buries itself in the
sediment, for only approximately one-fifth of its test is below the
general level of the surrounding substratum. Rather than pushing
itself under the sediment, the animal pulls the sediment over itself
with its spines and tubefeet. Generally, larger fragments of coral
are concentrated around the apical system (pl. 2: fig.1). Presumably,
the echinoid avoids placing small objects in the area of his madreporite.
Although usually covered with sediment, the echinoid can be found
because of the sorting of this sediment over the test, the large mound
it makes on the bottom, and the lighter color of the sediment on the
test. Furthermore, the echinoids leave a conspicuous track (pl. 2:
fig. 3), usually two to three feet long, consisting of a furrow with a
small ridge of sand on each side.

The echinoids do not occur in great density. On the average,
four to six specimens occur in an area of 100 square ft. These sandy
patches are usually 1-300 square feet in area and occur quite com-
monly in the coral-rock areas. Although similar sediment occurs
in the noncoral tracts, this echinoid was never found there. At
every station except one no weed or large algae was living in the sandy
patches with the echinoids. The one exception occurred at a locality

| NO. 3577 ECHINOIDS—KIER 9

85 feet deep (pl. 2: fig. 2), in which a small amount of algae was present.
It may be of significance that, in this deepest area studied, the echi-
noids were almost completely naked with only a small amount of

algae over their tests and no sediment. Perhaps it is because of the
decreased amount of light at these depths that the test is not as
completely covered.

[

Schizaster (Paraster) floridiensis Kier and Grant
One dead test was found at a depth of 85 feet on silt bottom devoid

of vegetation off Point Guignard. Although no living specimens

were found, the great fragility of the paper-thin test suggests that
it could not have been carried far from where it once lived. Pre-
sumably, this species lives buried as is the case with all schizasterids
with known living habits.

Comparison with Echinoids of the Florida Keys
It is of interest to compare the living habits of the Dominican

-echinoids with individuals of the same species in the Florida Keys,

as recently described by Kier and Grant (1965). The echinoid
fauna of Dominica was much less varied with only 10 species found,

only 5 of which occurred in large numbers, whereas, in the Keys,
17 species were reported, most of which were common. Probably

this difference is caused by the small number of different environ-
ments in the Caribbean off Dominica.

Meoma ventricosa was never found as deeply buried in Dominica
as in the Florida Keys. The Dominican specimens only partially
bury with approximately one-fifth of their tests below the general
level of the substratum, whereas in the Keys most of the individuals
keep their tests almost completely buried, many with an inch or so
of sand over the upper surface. In Dominica, M. ventricosa lives
very much like Clypeaster rosaceus does in the Keys, with sand and/or
plant material pulled up over its test.

Lecdia sexiesperforata lives deeper in the sand in Dominica than
in the Keys perhaps because it occurs in shallower water there. In
contrast, Clypaester subdepressus lives much deeper in Dominica,
where it is found three to four inches below the surface, but in the
Keys it normally walks along the surface with sand pulled over it
or is buried under only an inch of sand. Because of this, they are
very difficult to find in Dominica but easy in the Keys.

Diadema antillarum, Echinometra lucunter, Echinometra viridis,
Tripneustes ventricosus, and Eucidaris tribuloides live in similar habi-
tats in Dominica and the Florida Keys. It is of interest that the two
color types of Echinometra lucunter found in Dominica are also present
in the Keys.

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Literature Cited
Krier, P. M., and Grant, R. E.

1965. Echinoid distribution and habits, Key Largo Coral Reef Preserve, |

Florida. Smithsonian Misc. Coll., vol. 149, no. 6, 68 pp., 16 pls., |
15 figs.
Lewis, J. B.

1958. The biology of the tropical sea urchin T'ripneustes esculentus Leske |
in Barbados, British West Indies. Canadian Journ. Zool., vol. 36, |
no. 4, p. 607-621, pls. 8-9, 7 text-figs.

MortTENSEN, TH.
1928-1951. A monograph of the Echinoidea, 5 vols. and index.

Copen- |
hagen: C, A. Reitzel.

U.S. GOVERNMENT PRINTING OFFICE: i066

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1966 Number 3578

BREDIN-ARCHBOLD-SMITHSONIAN
BIOLOGICAL SURVEY OF DOMINICA

2. New Species of Diptera from Dominica
(Anisopodidae and Bibionidae)'

By ALAN STONE ”

Family Anisopodidae

Recent collecting by J. F. G. Clarke, H. E. Evans, O. S. Flint, Jr.,
R. J. Gagné, and W. W. Wirth for the Bredin-Archbold-Smithsonian
Survey of Dominica resulted in 34 specimens of two apparently
undescribed species in two genera of the family Anisopodidae. The
Mycetobia species is the third one described from the New World and
the second from the Neotropical Region. The abundant Oldiogaster
species has male terminalia apparently identical with those of 0.
antillarum Lane and Andretta from St. Croix, but the color differences
are too great to permit determination as that species. A second
Olbiogaster is represented by a single male with very distinctive

terminalia.
Mycetobia limanda, new species

Female: Vertex black, the ocelli yellow; eyes separated by slightly

less than width of anterior ocellus; occiput laterally yellow; antenna
black, the scape and pedical slightly brownish; face yellow; palpi and

1 Other faunal studies in this series are: 1, Kier, Proc. U.S. Nat. Mus., 1966, vol.
121, no. 3577, pp. 1-9. A companion series on the flora appears in the “Contri-
butions of the United States National Herbarium.”

2 Entomology Research Division, U.S. Department of Agriculture.

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121}

mouthparts darkened. Mesonotum subshining, dark reddish brown |
medially, somewhat paler laterally; pronotum yellowish brown but |

not contrasting with sides of mesonotum; hairs and setae of meso-

notum brown; scutellum yellowish brown with four setae; pleuron, |
postnotum, and halter yellow brown. Wing hyaline, slightly tinged |
with yellow; veins dark; subcosta ending just before level of radial |
sector fork; under side of subcosta with row of long hairs. Foreleg |
yellow brown, the tarsus slightly darkened; midleg the same but tibia |
darkened except at extreme base; hindleg mostly yellowish, the coxa |
posteriorly and femur near apex darkened. Dorsum of abdomen |
nearly black except for tergum I, tergum II laterally, tergum VIII, |
and terminalia, which are brownish yellow; venter yellowish with |
some darkening laterally before apex. Length 2.5-3.5 mm. Wing |

3—4 mm.

Male: Coloration as in female. Eyes nearly touching. Genital |
capsule not as long as terga VI and VII combined whereas this is |

longer in M. divergens. Slightly smaller than female.
Distribution: Dominica, Lesser Antilles; probably Costa Rica.
Holotype: 9, Clarke Hall, Jan. 18, 1965 (Wirth).

Paratypes: Clarke Hall, Mar. 21-31, 1965, 1 9; Clarke Hall, Feb. |

11-20, 1965, 1 ° (all Wirth). (USNM nos. 68, 116.)

A single specimen bearing the data ‘“‘La Suiza de Turrialba, Costa
Rica, Aug. Pablo Schild” in the U.S. National Museum is apparently
this species. It differs only in having the midtibia paler.

Because of the contrasting yellowish tip of the abdomen, I use as
a specific name the generic name of the flounder known as the Yellow
Tail.

The three New World species of Mycetobia may be differentiated
by the following key:

i eSubcostaibare (Nearctic)) 116) 2 ove. ees letaete tae e eeeeee divergens Walker
Subcosta with numerous long setae ventrally (Neotropical). ...... 2

2. Thorax, abdomen, and legs strongly patterned with dark markings; wing
more ‘than 5mm songs 5, iyi (9s ee ay wee = ke stonei Lane and Andretta

No distinctive pattern except for yellowish at base and tip of abdomen;
wine not more: than 4mm long) 0) eee limanda, new species

Olbiogaster danista, new species

Female: Frons and occiput dark, grayish pruinose, only the ocellar
triangle shining black; scape and pedicel yellow, flagellum black; face
yellow, in some slightly darkened above; palpus and mouthparts
yellow. Thorax mostly yellow with yellowish and some brown hairs,
with or without a pair of elongate darker markings on mesonotum
anteriorly, and a darkened spot on upper mesopleuron; scutellum
dark except at base, with a row of many dark hairs; postnotum some-
what infuscated medially; halter with yellow stem, black knob.

|
|
|

NO. 3578 DIPTERA—STONE 3

Wing hyaline except for dark stigmal spot on anterior inner basal
portion of coxa and tarsus; midleg yellow except for dark coxa, apical
two-thirds of femur, and tarsus; hindleg yellow except for dark coxa,
apical two-thirds of femur, and tarsus. Abdomen with three dark
spots on tergum J, usually clearly separated; terga II-V black on
basal half, yellow on apical half; terga VI and VII dark except very
narrow basally; venter yellow with sterna I1I-V more or less darkened
basally, VI and VII apically. Body and wing length each 6—6.5 mm.

Male: Coloration as in female except abdomen beyond tergum I
with wider black areas. Terminalia indistinguishable from those of
the type of 0. antillarum. Lane and Andretta (1958; p. 517, figs 19)
figured those for antillarum.

Distribution: Dominica, Lesser Antilles.

Holotype: 9, Clarke Hall, Jan. 11-20, 1965 (Wirth).

Paratypes: same data, 1 9, 3077; Clarke Hall, Mar. 1-10, 1965, 1 9,
1, and Mar. 21-31, 1965, 1 @ (Wirth); 5 miles south of Pont Casse,
Apr. 11, 1964 (Flint), 1 9; Clarke Hall, Apr. 12, 1964 (Flint), 1 9;
Hillsborough Estate, Mar. 15, 1965 (Wirth), 6 0"; Cabrits Swamp,
Feb. 23, 1965 (Wirth), 1 #; Layou R. mouth, Mar. 8, 1965 (Wirth),
1 @; Antrim, 1000 ft. Mar. 12, 1956 (Clarke), 1 o; Newfoundland,
on road to Rosalie, 1000 ft. Apr. 13, 1966 (Gagné), 1 of. (USNM
Nos. 68, 117.)

This species would run in the key of Lane and Andretta (1958)
to 0. similans Lane and Andretta, but similans has all coxae entirely
dark (not implied in key), the base of tergum I broadly dark, the
mesonotum and pleura mostly dark, and the stigma not crossing
vein R,. From antillarum Lane and Andretta, danista differs in
having the knob of the halter dark, the sternopleuron entirely yellow,
and terga IV and V with broad apical yellow bands rather than with
only lateral spots.

The name of this species (from the Greek word “danista,” meaning
“money-lender”) was suggested by the three dark spots on the first
tergum reminiscent of a pawnbroker’s sign.

Olbiogaster evansi, new species

Male: Frons and occiput dark, thinly grayish pruinose; scape
and pedicel yellow, flagellum black; face yellow; palpus and mouth-
parts yellow. Thorax mostly blackish with brown hairs; pronotum
yellow; mesonotum faintly reddish anterolaterally and before scutel-
lum; area around anterior spiracle and mesepimeron yellowish;
mesopleuron shiny black with a few hairs above and a broad trans-
verse band of gray pollinosity. Halter with yellow stem, black
knob. Wing hyaline except for narrow stigmal spot crossing cell Ry.
Legs mostly dark but with the following yellow: A large patch

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

anteriorly on forecoxa; fore- and midfemora; fore- and midtibiae,
but slightly darkened. Abdomen with terga shiny black except
for narrow yellow hind margins on I-V, slightly widened laterally;
sterna I-V mostly yellow, a dark area on side of sternum IJ. Wing
leneth 5.5 mm. Terminalia as figured (fig. 1).

Holotype: o&, South Chiltern, 1600 feet, Feb. 19, 1965 (Evans),
(USNM 69076).

This species would run in the key of Lane and Andretta (1958)
to O. chavantesi Lane and Andretta, but the color pattern of the head,
thorax, coxae, and sterna, and the structures of the terminalia are
different. The terminalia are closest to those of similans Lane and
Andretta, but the appendages lying dorsad of the cerci are broad

Ficure 1.—Olbiogaster evansi, new species, male terminalia, dorsal view: a, cerci and at-
tached appendages; b, aedeagus and attached appendages (drawn by Arthur D. Cushman).

and flat, pot narrow and rodlike, and in similans the tergal yellow
bands are much broader and the sterna are mostly black.
This species is dedicated to the collector, Howard E. Evans.

Family Bibionidae

The only Bibionidae collected on Dominica by the Bredin-Archbold
Smithsonian Survey consisted of 21 specimens of an apparently un-
described species of the genus Plecia Wiedemann. I take this oppor-
tunity to describe it.

Plecia porca, new species

An entirely dark brownish-black species with four shiny stripes on

notum and fumose wings. Jt runs to couplet 5 of Hardy’s key (1945,

No. 3578 DIPTERA—STONE 5

| p. 391) but with very different distimeres (harpagones) from the two
species of that couplet.

- Male: Head: Antenna black with 7 cylindrical flagellomeres, the
first about as long as scape and pedicel combined, the others subquad-
rate except last, which is reduced in size, hemispherical. Rostrum
about equal to antenna in length, shorter than head and sharply
turned backward under head, reddish except near apex. Thorax
| brownish pruinose, the notum dull reddish before scutellum and on
sides, the three narrow dorsal stripes somewhat grayish; a submedian
pair of raised, narrow, polished black stripes on median half of notum
, and a pair of similar but broader stripes from just behind the antero-
lateral depression to corner of scutellum, often slightly reddish poste-

VIII T

|
|
|
|
|

;
!

VIII T

Ventral Dorsal

Lateral

| Ficure 2.—Plecia porca, new species, male terminalia, ventral, dorsal, and lateral views
(D=distimere, S=sternum, T=tergum).

-riorly; scutellum and postnotum dull; anterior pronotum and upper
portion of sternopleuron with hairs; hairs of mesonotal furrows very
small. Wing fumose, the stigma and costal cell darker; veins dark
with short fine hairs; vein R; curved toward apex, shorter than basal
portion of radial sector; r-m crossvein about half way between medial
crossvein and fork of M. Halter, legs and abdomen nearly black.
Terminalia (fig. 2): Ninth tergum with a broad triangular emargina-
tion posteriorly with an inwardly turning pair of somewhat shiny pro-
tuberances in the median angle; a low protuberance medially near
_ base; ninth sternum slightly concave medially near apex with a pair
of marginal ridges each ending in a slender submedian process; disti-
mere rather shiny black but covered with hairs except on the concave
dorsal surface; each distimere bilobed, the apico-median lobe with a

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

transverse depression in the apex and the lateral lobe curving dorsally
to a point. Length of wing 6.5-7.0 mm.

Female: Coloration and other nonsexual characters as in male |
except that the antenna has 9 flagellomeres. Frons subshining |
medially, the sides and occiput with brownish pollinosity; distance |
between eyes about equal to smallest transverse dimension of eye; |!
rostrum slightly shorter than antenna. Wing length 8.0 mm.

Holotype: Sylvania, Dominica, Jan. 28, 1965 (Clarke and Clarke),
o', (USNM 68228).

Paratypes: 1800 ft., Pont Casse, Jan. 12, 1965 (Clarke and Clarke), |
1 o@; mouth of Layou River, Jan. 20, 1965 (Clarke and Clarke), 1 7; |
light trap, Sylvania, Jan. 23, 1965 (Wirth), 1 o;grassy marsh, Sylvania,
Jan. 25, 1965 (Wirth), 6 o, 1 9; 2.5 miles west of Pont Casse, Jan. 27,
1965 (Wirth), 1 o, 1 9; 2 miles west of Ponte Casse, Jan. 27, 1965
(Wirth), 1 &; Sylvania, Jan. 28, 1965 (Clarke and Clarke), 2 <;0.5 |
miles west of Point Lolo, Jan. 28, 1965 (Wirth), 2 o&; 1 mile east of |
Pont Casse, Jan. 29, 1965 (Wirth), 1 9; Clarke Hall, Apr. 5, 1965, |
(Flint), 1 o, 19.

Literature Cited
Harpy, D. Eumo
1945. Revision of Nearctic Bibionidae including Neotropical Plecia and
Penthetria (Diptera). Bull. Univ. Kansas, vol. 30, pp. 367-547,
13 pls.
LANE, J., and p’ANpDRETTA, C., JR.
1958. Neotropical Anisopodidae (Diptera, Nematocera). Stud. Entom.,
vol. 1, pp. 497-528, illustr.

U.S. GOVERNMENT PRINTING OFFICE: 1966

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION « WASHINGTON, D.C.
Volume 121 1967 Number 3579
VALID ZOOLOGICAL NAMES
OF THE PORTLAND CATALOGUE

By Haratp A. REHDER

Research Curator, Division of Mollusks

Introduction

An outstanding patroness of the arts and sciences in eighteenth-

century England was Lady Margaret Cavendish Bentinck, Duchess
of Portland, wife of William, Second Duke of Portland. At Bulstrode
in Buckinghamshire, magnificent summer residence of the Dukes of

Portland, and in her London house in Whitehall, Lady Margaret—a

widow for the last 23 years of her life—entertained gentlemen in-
_ terested in her extensive collection of natural history and objets d’art.

Among these visitors were Sir Joseph Banks and Daniel Solander,
pupil of Linnaeus. As her own particular interest was in conchology,
she received from both of these men many specimens of shells gathered
on Captain Cook’s voyages. Apparently Solander spent considerable
time working on the conchological collection, for his manuscript
on descriptions of new shells was based largely on the ‘Portland

~ Museum.’’

When Lady Margaret died in 1785, her ‘“Museum” was sold at

auction. The task of preparing the collection for sale and compiling
_ the sales catalogue fell to the Reverend John Lightfoot (1735-1788).

For many years librarian and chaplain to the Duchess and scientif-
1

2 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

ically inclined with a special leaning toward botany and conchology,

he was well acquainted with the collection. It is not surprising he |

went to considerable trouble to give names and figure references to so

many of the mollusks and other invertebrates that he listed. It is |
interesting to note also that, although the sale of the collection was |
supposed to begin on April 24 of that year, the auction did not take |
place until a month later—a delay due probably to the time required :

to compile and print the catalogue.

Although the ‘Portland Catalogue’ has been used as a reference |
source of scientific names almost since its publication—witness its |
citation by Dillwyn in 1817, Swainson in 1822, and Gray in 1824-28
(Kay, 1965, p. 13) and the use of Solander’s names by such a conserva- |
tive worker as G. W. Tryon in his “Manual of Conchology’’—its |
modern use in malacological nomenclature dates from the appearance |
of papers by Iredale (1916) and Dall (1921). In spite of some orig- |
inal opposition to the resurrection of these names, the “Portland |
Catalogue,” as Kay has stated (1965, p. 10), is now generally con- |

sidered a valid source of scientific names.
That the Reverend John Lightfoot was the anonymous compiler

of the ‘‘Catalogue’’—and thus the author of the new names proposed |

therein—has recently been stated clearly by Dance (1962). The re-
searches of Kay (1965) have confirmed a long-held view of the present
writer that the authority to be cited for all new names in the “Cata-
logue”’ is the compiler and not Solander. Lightfoot used the names
found in Solander’s manuscript and possibly also on labels in the
collection, which may explain the absence in the manuscripts of some
names credited to Solander in the “Catalogue.”’

The names derived from Solander’s manuscripts and labels were
designated by Lightfoot in most instances by adding the letter “S” to
the scientific name. It must be emphasized, however, that only the
names were taken from the manuscripts and rarely did the compiler
use a bibliographic reference to a published figure that was given by
Solander. In my view, Solander cannot in any way be considered
responsible for the principle condition that made the names available—

1.e., publication (see Article 50 of the “International Code of Zoolog- |

ical Nomenclature,’ 2nd edition, 1964)—although he can be consid-
ered to have been the source for some of the names in the “Catalogue.”
Since he was not the compiler (and therefore not the author of the
“Catalogue,” either solely or in part), he did not validly introduce any
of the names in the “Catalogue” into the literature.

Although many names first proposed in the “Portland Catalogue”
are now in general use—some have been accepted for over a hundred
years— others have been ignored or passed by until fairly recently.
In a work that has been accepted as a valid source of scientific names,

|
|
:

NO. 3579 PORTLAND CATALOGUE—REHDER 3

we cannot, in my estimation, choose to use some of the validly pro-
posed names and not use the others, unless, of course, the latter names
are truly nomina dubia.

I, therefore, publish this list of names in the conviction that they
have been validly proposed and should enter into the ranks of zoolog-

A

CorAni Awl) OnG \iUek

OF THE

PORTLAND MUSEUM,

LATELY THE PROPERTY OF

he Duchefs Dowager of Portland,
DECEASED:
Which will be SOLD by AUCTION,

B Y

Mr. SEINNER azd Co:

Ona MONDAY the 24th of APRIL, 1786,
AND THE = Lig
THIRTY-SEVEN forrowine DAYS,,~ :

- GAM IT oes,

Ar LWELVE O7 CLOCK,

Sunpays, and the sth of June, (the Day his Mayesty’s Birtu-Day
is kept) excepted ;

At her lae DWELLING-HOUSE,

In PRIVY-GARDEN, WHITEHALL;
By ORDER or tue ACTING EXECUTRIX.

To be viewed Ten Days preceding the Sale.

CATLOGUES may now be had on the Premises, and of Mr. SKINNER
and C°, ALDERSGATE-STREETy Price Fiye SHILLINGS, which will admit
the Bearer during the Time of Exhibition and Sale.

N°S2

Figure 1.—Title page of the “(Catalogue of the Portland Museum.”

ical nomenclature, to stand or fall on their own merits. Some of
these may supplant long-familiar names. If, however, we are to main-
tain the classification of mollusks and other marine invertebrates on a
firm basis, we must adhere as closely as possible to the rule of strict
priority and avoid exceptions and suspensions of the Rules.

+ PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Dall (1916) and, to a lesser extent, Iredale (1921) attempted to
identify the names proposed in the ‘‘Portland Catalogue,” some identi-
fications of which I comment on in the present paper. Many of the
“Catalogue”? names listed by Dall are nomina nuda and thus invalid.
These I have omitted. Others actually were taken over from Lin-
naeus or Born in their original sense. In some entries Lightfoot has
referred to the second volume of Martyn’s ‘Universal Conchologist,”’
usually dated from 1789, but without citing a figure number. It is
probable that he saw the plates of this as yet unpublished volume.

Each entry in the present paper is arranged in the following order:
the lot number of the ‘‘Portland Catalogue,’ a verbatim citation of the
relevant textual material, the name to be used for the taxon (in bold-
face type), the principal synonyms (in italics), including the most re-
cent ones in which the name is credited to either Solander, Humphrey,
or Lightfoot, and, where necessary, my own remarks.

Lightfoot’s name sometimes is enclosed in brackets to follow Recom-
mendation 51A of the International Code of Zoological Nomenclature,
which states that the name of the author of a taxon published anony-
mously “should be enclosed in square brackets to show the original
anonymity.” Where the trivial name has been transferred to another
genus I have replaced the square brackets around the author’s name
with the required parentheses alone, to avoid the use of both square
brackets and parentheses.

In all, 2 genera and 120 trivial names are considered to have been
first validly proposed in the “Catalogue.’’ The 2 genera, Isog-
nomon and Placuna, have been in general use now for some time
although usually credited to Solander.

Of the 120 trivial names, 111 pertain to mollusks, 5 to the Cirripedia
(Crustacea), 2 to the Echinodermata, and 1 each to the Madreporaria
(Coelenterata) and Brachiopoda.

Of these 120 names, I consider 9 to be nomina dubia, 3 junior
homonyms, and46synonymsof earlier names. This leaves 62 available
names that should be credited to Lightfoot as author. Forty-six of
these have been used in recent malacological literature, credited
either to Solander of Humphrey or, in a few cases, to Lightfoot. Only
16 (or 13%) of all the new names in the “Catalogue” have apparently
not come into use. Six of these I have been unable to locate as
accepted senior synonyms in the literature of the past 140 years and,
thus, they should be considered nomina oblita. The remaining 10,
in all cases but one, have been used as senior synonyms in the past 50
years and, thus, are available.

NO. 3579 PORTLAND CATALOGUE—REHDER 5

NinereentH Day's SAtLe..

MONDAY tua 15th or MAY, 1786,

SHELLS, CORALS, &.

3924 IFTEEN cards, containing various fpecies of: Nerita, among which are
~ Canrcna, albumen, Mamilla, pelorontha, &c,

1g2g Four.fine Oltrez, viz. Eburnea, fquamofa, and 2 lurida, S—all rare

1926 Eight fine Petrifactions, viz. 2 Strombus luctator,.Brander’s Fo/s. Haxson. pl. 5+
fig: 64. an Anomia Gryphites, Z. an Echinites from Verona, &cs

3927 Fourteen Bivalves of various genera, among which are 2 Tellina radiata, ¢
Oftrea maxima, L. fine, 3 Arcas, &c.

1928 Nine cards of various fpecies of Englifh Crabs, viz. Cancer pifiim, minutusy.
phalangium male and female, Dorfettenfis; Aftacus Bernhardus, all defcribed
by Pennant, and the eared Crab, nox defcript.

1929 A very large and fine Buccinum Galea, Z. or tun fhell from-the Mediterranean

1930 Six different curious Ovaries of marine fhell-fih, and 4 eggs of Helix ovipara,
Lifter, 23. 21. fome pearls, &ce.

1931 Seven cards containing various fpecies of Cardium,. viz. one valve of hami-
cardium, 2.odd.valves of.retufum, medium, Z. muricatum, Ciliare Se varie-
gatum, S.

1932 Fourteen.cards of different fpecies of Needle Buccina, viz. Hecticum, ftrigila-
tum, lanceatum, 5S. torolofum, articulatum, pertufum, capreolus, cinercum,
& fufus,. S$.

1933 A {mall Echinus rofaceus and y.varieties of Echinus orbiculus, L.—/ome rare

1934 Six fine f;ecies of Venus, viz. Deflorata, literata, merctrix, &c,

1935 Two pair of very rare fpecies of Voluta, viz» Sanguifuga, ZL. and turricula,,
S.—Jine

1936 ‘Two fine varicties-of Chama Gigas, L. or furbelowed clamp

1937 A beautiful Oftrea nodofa, or Duck’s-foot Pecten, and Oftrea plica, L. 2orb rare

1938 Three fine varieties of Murex ramofus, viz. the Stag’s horn, and 2 others

1939 Three-kinds of Pholas, one imbedded ix svood, a fmall rare Solen, a Mya, &c.

1940 A fine pair of Voluta calata and another of Pepo, S. Martyn, Vol. III.
Fg. 788—770

1941 A pair of very fine Trochus Solaris, L. cr Gun fhell, from the I. Iudies—rare

3942 Six fine fpecimens of Trochus, viz. 2 of Granofus from New-Zealand, Mary,
Fol. 1. figs 37+ t..2 maculatus, LZ. and 2 others

Farious teftaceous and other ANIMALS, VEGETABLES, &¢. in SPIRITS.

3943 A fine clufter of Lepas fignita, S. found in the Briti/o Chanuel, the animal
Terebella, with it’s augur, of the Teredo navalis, or ship-worm, and fundry
Ovaria of Buccinum undatum, L, ix 3 dotiles 1944 A

Ficure 2.—A sample page (87) of the “Catalogue of the Portland Museum.”

The Valid Names

15 “Strombus Fusus, L. Persicus, or Persian Spindle, a pair,
fine, Lister. 854. 12.”
Tibia insulachorab Réding, 1798
Strombus Fusus L. Persicus (Lightfoot, 1786], p. 3.
Rostellaria curvirostris Lamarck, 1816, p. 4.

Because Lightfoot used Persicus as a geographic adjectival trinomial
and placed it in italics, I consider this name as invalidly proposed.
It should be noted that in almost all cases the geographic origin of the
specimens listed is given in italics, and in those cases, relatively few

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

in the “Catalogue,” where the scientific names are in italics, the
entire name is so printed.

31 “A fine pair of Trochus Onustus, or Carrier Trochus, rare,
Favanne. Labs 120 Cyr ken2?
Xenophora conchyliophora (Born, 1780)

Trochus onustus [Lightfoot, 1786], p. 4.
Phorus onustus Reeve, 1842, p. 160.

Not Trochus onustus Nilsson, 1827 (=Xenophora sp., Creataceous,
Sweden).
50 “ . « 2e0trombus “tricornis: \WMartin. 84345. Disk 87327
Strombus tricornis [Lightfoot, 1786]
Strombus tricornis [Lightfoot, 1786], p. 5.
Strombus tricornis Fischer, 1807, p. 188.
Strombus tricornis Lamarck, 1816, p. 3.
Strombus tricornis Humphrey, 1786.—Abbott, 1960, p. 53.
89 “Voluta Nobilis, S. an extremely rare species of the Music
kind, from the E. Indies. List. 799.6.”
Volutocorona nobilis [Lightfoot, 1786]
Voluta nobilis [Lightfoot, 1786], p. 6.
Voluta scapha Gmelin, 1791, p. 3468.—not Voluta scafa Lightfoot,
1786.
Aulica nobilis Solander.—Abbott, 1962, p. 98.
Volutocorona nobilis (Solander, 1786).—Weaver, 1964, p. 6.
Troschel in 1866 had already accepted this trivial name, citing it
as ‘‘Voluta (Aulica) nobilis (Solander).”
The group Volutocorona Pilsbry and Olsson, 1954, is being retained
as a distinct genus, until its radular characters are known, although
it appears to be very close to Aulica Gray, 1847.

97 “A young specimen of Serpula gigantea or great Oriental
Worm Shell, with its Operculum, very rare—Seba, vol. III.
Tab. 94.”

Kuphus polythalamia (Linnaeus, 1766)
Serpula gigantea (Lightfoot, 1786], p. 6.
Septaria arenaria Lamarck, 1818, p. 437 (not Serpula arenaria L.,

1758).
Not Serpula gigantea Pallas, 1766, as Dall erroneously states (1921,
p. 129).
176 “Ostrea isognomon L. called Isognoma lignea by S. very rare,

with an Oyster adhering.”’
Isognomon [Lightfoot, 1786]

This is the first use of the generic name Jsognoma (or Isognomon).
See lots 968 and 3041, wherein the now generally accepted form
Isognomon is used. Ostrea isognomon Li. (=O. isognomum L.) becomes
the type of the genus by tautonymy.

NO. 3579 PORTLAND CATALOGUE—REHDER 7

187 “Three fine specimens of Tellina, viz. ... cruentae, S.
Knorr VI. 12. 1. all very rare.”
Sanguinolaria cruenta (Lightfoot, 1786)

Tellina cruentae (Lightfoot, 1786], p. 10.
Tellina operculata Gmelin, 1791, p. 3235.
Tellina semiplanata RGding, 1798, p. 186.

Abbott (1954, p. 439), following Dall (1921, p. 99), erroneously used
this name for 7. sanguinolenta Gmelin, in which he has been followed
by Shikama (1964, pl. 50, f. 3).

An examination of Knorr’s figure shows that the name cruenta
should be applied to the large, more elongate and unicolored species
formerly known as S. operculata Gmelin, inhabiting the southern
shores of the Caribbean, from Nicaragua south to Brazil.

It is not the Eastern Pacific S. bertini Pilsbry and Lowe, as Keen
(1958, p. 188) thinks possible.

See lot 1360, wherein Lightfoot spells the trivial name correctly:
cruentda.

264 “Voluta incrassata, S. Martyn, 499, 500... . very rare.”
Oliva incrassata (Lightfoot, 1786)

Voluta incrassata (Lightfoot, 1786], p. 13.

Oliva angulata Lamarck, 1811, p. 310.

Voluta incrassata Dillwyn, 1817, p. 516.

Oliva incrassata (Solander, 1786).—Keen, 1958, p. 420.

295 “Kchinus anemonoides, or purple Anemony Echinus, extremely
rare, and one ditto with its spines on Favanne 56. H.H.”
Podophora atrata (Linnaeus, 1758)

Echinus anemonoides (Lightfoot, 1786], p. 14.

301 “Buccinum Iris, S. Martyn, vol. I, fig. 2.b. the Epidermis of
this singular species when wet is of various colors, and it is
exceedingly scarce.”

Latirus iris (Lightfoot, 1786)
Buccinum prismaticum Martyn, 1784, fig. 2 (nonbinomial).
Buccinum iris (Lightfoot, 1786], p. 14.
Murex prismaticus Gmelin, 1791, p. 3559.
Latirus prismaticus Martyn.—Tryon, 1881, p. 93.
Latirus iris Humphrey.—Kaicher, 1957 [sect. 7], pl. 8.

353 “Two fine species of Placuna, S. viz. placenta, and Ephippium,
the Chinese Window, and Polish Saddle Shells, both very
rare.”

Placuna [Lightfoot, 1786]

Placenta Retzius, 1788.
Placuna Bruguiere, 1792.

This is the first use of the generic name.

8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

372 “Two fine specimens of Buccinum monodon, S. or Unicorn
shell, from Terra del Fuego, Martyn, vol. I, fig. 10e.”
Acanthina monodon (Pallas, 1774)

Buccinum monodon Pallas, 1774, p. 33.
Calcar Martyn, 1784, fig. 10 (nonbinomial).
Buccinum monodon [Lightfoot, 1786], p. 17.
Buccinum monoceros Bruguiere, 1789, p. 253.

400 “A large and fine Helix plicata, undescribed, Favanne, pl. 61,
De ae ce rares”
Pila urceus (Miieller, 1774)
Helix plicata (Lightfoot, 1786], p. 18.
Not Helix plicata Born, 1778.
408 “A large and fine Turbo undulatus, from New Holland,
extremely scarce, Martyn, vol. I. 29. p.”
Subninella undulata (Lightfoot, 1786)
Limaz undulatus Martyn, 1784, fig. 29 (nonbinomial).
Turbo undulatus [Lightfoot, 1786], p. 18.

Turbo undulatus Gmelin, 1791, p. 3597.
Subninella undulata Solander.—Allan, 1959, pp. 68, 486.

566 “A very large Voluta ponderosa, S. or heavy Volute, with
a fine mouth, Martyn 916.”
Turbinella ponderosa (Lightfoot, 1786)
Voluta ponderosa [Lightfoot, 1786], p. 25.
Xancus pyrum Réding, 1798, p. 1384 (not Linnaeus, 1758).
Turbinella rapa Lamarck, 1816, p. 7.
A critical study of large series of this species and Xancus pyrum
(L., 1758) may show that the former is merely a subspecies of the
latter (see Dodge, 1955, pp. 132-134).

601 “Two beautiful specimens of Helix Vitellus Rum. 22.1... .”
Polymita picta (Born, 1780)

Helix vitellus (Lightfoot, 1786], p. 26.

610 “‘A pair of Voluta virescens, S. Martyn, 932. 933 from Guinea.”
Voluta virescens [Lightfoot, 1786]

Voluta virescens [Lightfoot, 1786], p. 26.

Plejona muta Réding, 1798, p. 60.

Voluta polyzonalis Lamarck, 1811, p. 68.

Voluta virescens Dillwyn, 1817, p. 562.

Voluta virescens Solander.—Abbott, 1954, p. 250.

611 “Voluta arausiaca, S. or Prince of Orange’s Flag Music shell,
from Amboyna .. . extremely rare, Rumph. 37. 2.”
Harpulina arausiaca (Lightfoot, 1786)

Voluta arausiaca [Lightfoot, 1786], p. 26.

Voluta arausiaca Shaw, 1790, pl. 52.

Voluta vexillum Gmelin, 1791, p. 3464.

Harpulina arausaica [sic] Solander.—Shikama, 1968, pl. 81.

NO. 3579 PORTLAND CATALOGUE—REHDER 9

626 “Mya gigas, S. List. conch. 414, with two Gorgoniae adhering
to it.”

Panope glycimeris (Born, 1778)
Mya gigas (Lightfoot, 1786], p. 27.

707 “Voluta elongata, S. or Midas’s ear Land Snail, a very rare
terrestrial shell from New Caledonia, Martyn, vol. I, fig. 25.
leet
Placostylus elongatus (Lightfoot, 1786)

Limaz fibratus Martyn, 1785, fig. 24 (nonbinominal).
Voluta elongata [Lightfoot, 1786], p. 30.

Bulimus bovinus Bruguiere, 1792, p. 345.

Ellobium australe Réding, 1798, p. 106.

Not Voluta elongata Schroeter, 1804 (=?Aurinia sp.); not Voluta
elongata Swainson, 1821 (= Alcithoe swainsoni Marwick, 1926).

It is strange that both Franc (1957, p. 152) and Solem (1961,
p. 477) use Martyn’s name, even though in 1957 the ‘Universal
Conchologist”’ was officially rejected for nomenclatorial purposes.
Neither do they list the three subsequent, earliest names, those of
Lightfoot, Bruguiere, and Réding. Since the name fibratus is un-
available, the adoption of Lightfoot’s name should prove acceptable.

708 “A fine specime of Voluta amphora, S. or clouded Persian
Crown, from China, rare . . . Martyn, 780.”
Cymbium (Melo) amphera (Lightfoot, 1786)
Voluta amphora [Lightfoot, 1786], p. 30.
Cymbium flammeum Réding, 1798, p. 151.
Melo diadema Lamarck, 1811, p. 57.
Cymbium (Cymbium) amphorus (Solander).—Shikama, 1963, pl. 85.

915 “Helix Otis an extremely rare species, figured by Favanne, tab.
63. fig.11.”
Lampadion otis (Lightfoot, 1786)
Helix otis (Lightfoot, 1786], p. 38.
Helix labyrinthus Lamarck, 1792, p. 352.
Lampadion labyrinthus Réding, 1798, p. 78.
Helix (Labyrinthus) otis Soldr. [sic], Beck, 1837.
Pleurodonte (Labyrinthus) otis (Solander).—Pilsbry, 1910, p. 504
The generic name Labyrinthus Beck, 1837, must yield to the earlier
name Lampadion Réding, 1798. Winckworth (1945, p. 141) lists as
type Lampadion labyrinthus Réding, citing Gray (1847, p. 173) as
authority for this type designation. Although Gray gave Heliz otis
as the type of the genus, which was not one of the originally included
species, Winckworth’s action is valid (‘International Code of Zoologi-
cal Nomenclature;” art. 69(a)(iv)). Wurtz (1955, pp. 106-107),
apparently overlooking Winckworth’s statement, designated as type
the same species of Réding, Lampadion labyrinthus, fixing this name
on the figure in Favanne (1780), cited by Réding. Wurtz then goes

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

on to say that this figure represents an unrecognizable species and

thus disposes of the name Lampadion. Wurtz was unaware of the

fact that Réding cited the wrong figure in Favanne for his species.

Réding apparently meant to cite F 11 on plate 63 but referred instead

to “fig. 2,”’ possibly reading the 11 asa Roman II. That Réding did

make an erroneous citation for the species to which he gave the

popular name ‘Das Labyrinth,” is proven by the fact that Favanne,
in his own later anonymous work (1784, p. 6), cites pl. 63, F 11, of
his “Conchylogie” for this species, “Le Labyrinthe.”’

968 “Three varieties of Isognomon perna S. (Ostrea, L.)
enveloped with Sponge, and another curious species of the
same genus.”

Isognomon
Here Isognomon is used, as it is later (lot 3041) with the same
species: Ostrea perna L. The type of the genus is, however, Ostrea

isognomum by tautonymy (see lot 176).

969 “Two fine species of Voluta, viz. Melo, Martyn, vol. III.
TEAR TOs ue
Cymbium (Melo) melo (Lightfoot, 1786)
Voluta melo [Lightfoot, 1786], p. 41.
Voluta anguria Lightfoot, 1786, p. 64.
Voluta indica Gmelin, 1791, p. 3467.
Cymbium maculatum Réding, 1798, p. 152.
Cymbium melo Riding, 1798, p. 152 (in part).
Voluta melo ’Soland.—Lamarck, 1811, p. 59.
Melo melo Solander.—Abbott, 1962, p. 102.

969 “... and Scafa b. S. Adans. Seneg. 3. 2.”
Cymbium cymbium (Linnaeus, 1785)
Voluta scafa (Lightfoot, 1786], p. 41.

Adanson’s figure is cited by Lightfoot for Solander’s ‘‘variety b”’
of his Voluta scafa, which is validated in lot 3039 by a reference to a
figure in Martini. Both of these figures represent Voluta cymbium
Linnaeus.

990 “Two fine Patella Mytiliformis, Humphrey’s Conch. pl. III.
Sig. 9 from Falkland’s Islands . .. .”
Nacella mytilina (Helbling, 1779)
Patella mytiliformis (Lightfoot, 1786], p. 42.
Patella mytilina Gmelin, 1791, p. 3698.
Not Patella mytiliformis Gmelin, 1791 (=Crepidula sp.).
1001 ‘Two fine specimens of Arca fusca, S. Gualt. 87. G. one of
them affixed to a pinna.”’
Nomen dubium (figure indeterminable)

Not Arca fusca Bruguiere, 1789 (=Barbatia amygdalumtostum
(Réding, 1798).

|
|
|

No. 3579 PORTLAND CATALOGUE—REHDER ial

Solander’s name was assigned by earlier authors to various species.
Donovan (1803, p. 158) gave Solander’s name to specimens he believed
to be what is now called Arca imbricata Bruguiere. In this he was
followed by Montagu (1808, p. 51), who assigned certain British
specimens to this species, obviously confusing Arca tetragona Poli with
imbricata, since he, like Donovan, cites Lister’s figure Chie2.. pl sore
fig. 207) of imbricata for Arca fusca. Dillwyn (1817, p. 227) actually
places A. fusca Solander (mss.) in the synonymy of Arca imbricata
Bruguiere, in which he is followed by Iredale (1961, p. 92). Lamy
(1907, p. 41), on the other hand, places the “Portland Catalogue”
name in the synonymy of A. tetragona Poli, 1795, and Dall (192.55p.
98) equates the species with fusca Bruguiere, apparently on the sole
and uncritical basis of the similar name. An examination of the
figure in Gualtieri, the only reference cited by both Solander and
Lightfoot (Kay, 1965, p. 15) leaves us in doubt as to exactly what
species is represented. The three figures are rather stylized and
could represent any of several species of Arca. For this reason I
feel that I am justified in regarding this name as a doubtful one.

1005 “Five species of Solen . . . pallidus, S. (List. conch. t. 412
WIUCLUOT) wt.
Nomen dubium (figure indeterminable)
Dillwyn (1823, p. 22) says that Lister’s figure represents Solen
truncatus Wood, 1815, but this must be an error, for there is no
resemblance between the figures of Wood and Lister.

1005 “Solen. . . plebeius, S. (Lister, 421. f. 265).”
Tagelus plebeius (Lightfoot, 1786)
Solen plebeius (Lightfoot, 1786], p. 42.
Solen gibbus Spengler, 1794, p. 104.
Solen plebetus ‘Solander.’—Dillwyn, 1823, p. 22.
Tagelus plebecws Solander.—Abbott, 1954, p. 440.
1046 “A pair of fine Conus Augur, S. or dotted Cone, Knorr. VI.
HbR AS fig. Gn
Conus augur [Lightfoot, 1786]
Conus augur [Lightfoot, 1786], p. 44.
Conus augur Hwass, 1792, p. 685.
Conus augur Solander.—Tomlin, 1937, p. 216.

”)

1055 “‘Argonauta navicula, Rum. 18. 4 .
Nomen dubium
This may be the same as Argonauta gondola Dillwyn, 1817, but the
figure in Rumphius is too crude to identify with certainty the species
intended.

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

1055.) “21. ,. andshians.S:.2hid.18:Bxboth vanes?
Argonauta hians [Lightfoot, 1786]
Argonauta hians |Lightfoot, 1786], p. 44.
Argonauta hians Dillwyn, 1817, p. 334.
Argonauta hians Solander.—Tryon, 1879, p. 136.

1206 ‘“T'wo Cypraea pantherina, S. List. 681.28... .”
Cypraea pantherina [Lightfoot, 1786]
Cypraea pantherina (Lightfoot, 1786], p. 50
Cypraea vinosa Gmelin, 1791, p. 3421.
Cypraea pantherina Solander.—Schilder, 1939, p. 185.
This and the previous species, with Solander given as the author
of each, have been in general use for a long time.

1240 ‘Two scarce varieties of Trochus alveolatus, or Beehive
snail Mist. G2: iG0r. <. cave?
Sagda epistylium (Miiller, 1774)
Trochus alveolatus (Lightfoot, 1786], p. 52.
1242 “Four varieties of Ostrea perna, Z. (Isognomon, 8S.) ;
Isognomon [Lightfoot, 1786]
See lot 968.

1266 “Three fine species of Balanus. . . Pyramidalis, S. Ellis,

jigs Oe?
Balanus tintinnabulum Linnaeus, 1758
Balanus pyramidalis (Lightfoot, 1786], p. 53.
It seems certain that the above reference refers to the plate illustrat-
ing Ellis’ paper on barnacles published in the ‘Philosophical Trans-
actions” (1758, vol. 50, part 2, pp. 845-855, pl. 34).

1301 ‘Two very large specimens of Patella fungoides or Mushroom
limpet, from the Cape of Good Hope. Humphrey’s Conch. pl.
DVeajig. 16.”
Patella (Scutellastra) barbara Linnaeus, 1758
Patella fungoides [Lightfoot, 1786], p. 55.

Not Patella fungoides Réding, 1798 (=Acmaea jamaicensis Gmelin,
1791).

1315 “Four fine spectmens of Placuna, S. (Anomia, L.) viz.

Placenta and Ephippium, both from China, and rare.”
Placuna [Lightfoot, 1786]

See lot 353.
1358 ‘A large and fine Cardium robustum, S. from Florida, very
SCOTCE.. bt oASTer ove oan |

Dinocardium robustum [Lightfoot, 1786]
Cardium robustum [Lightfoot, 1786], p. 58.
Cardium magnum Born, 1778, p. 34 (not Linné, 1758).
Cardium ventricosum Bruguiere, 1789, p. 228.
Dinocardium robustum Solander.—Abbott, 1954, p. 401.

SSS

No. 3579 PORTLAND CATALOGUE—REHDER te

1359 ‘Three fine and rare species of Patella, viz. two of Mytili-
formis, Humphrey’s Conchology, 3.9 . i
Nacella mytilina (Helbling, 1779)
See lot 990.

1360 ‘Four curious and uncommon species of Tellina .
Cruenta, S. Knorr VI. 12. 1.”
Sanguinolaria cruenta (Lightfoot, 1786)
See lot 187.
It should be noted that Lightfoot gives here the correct spelling
for this specific name.

1448 “Voluta. . . Anguria, S. or great brown African Melon,
Vol. Martyn UI. 767.”
Cymbium (Melo) melo (Lightfoot, 1786)
The figure cited for this species represents the adult stage of the
shell on which melo is based.
See lot 969.

1455 ‘“Buccinum Iris, S. Martyn vol. I. fig. 2. b. very fine and
extremely scarce.”
Latirus iris (Lightfoot, 1786)
See lot 301.

1479 ‘‘A pair of Murex elongatus, a nondescript species, extremely
scarce. Favanne pl. 79. H.”
Pterynotus (Marchia) elongatus (Lightfoot 1786)
Murex elongatus (Lightfoot, 1786], p. 65.
Murex clavus Kiener, 1841, p. 111 (not Gmelin, 1791).
Murex (Marchia) elongatus (Solander).—Kira, 1959, p. 61.
Not Murex elongatus Lamarck, 1816 (=Chicoreus asianus Kuroda,
1942).

1501 ‘Conus quercinus S. Martyn, Vol. II. 657... .”
Conus quercinus (Lightfoot, 1786], p. 67.
Conus quercinus Hwass, 1792, p. 681.
Conus quercinus Solander.—Tomlin, 1937, p. 300.

1560 “Mytilus Castaneus S. List. 1055. 9. extremely scarce.’
Modiolus vagina Lamarck 1819
Mytilus castaneus (Lightfoot, 1786], p. 69 [a nomen oblitum].
Modiola vagina Lamarck, 1819, p. 112.
Modiola castanea Gray, 1825, p. 138.
Mytilus castaneus Solander.—Dillwyn, 1823, p. 47.
Modiolus vagina Lamarck; Lamy, 1937, p. 329.
Not Modiola castanea Say, 1822 (= Lioberus castaneus Say, 1822).

| Dall (1921 p. 128) equates this name with Elliptio complanatus
(Lightfoot) because in copying Lightfoot’s reference to Lister, he
changed ‘'1055,9” to ‘'154,9.”

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

1601 ‘An extremely fine pair of a species of perforated Patella, the
only two that are known, named Macroschisma, Humphrey’s
Conch. Plate 7. fig. 3. 3.”
Nomen dubium
Not Patella macroschisma Holten, 1802, nor Dillwyn, 1817 (= Macro-
schisma maxima A. Ad., 1850).

An examination of the figures in DaCosta’s ‘Conchology or Natural
History of Shells” [for an explanation of the authorship of this work,
see the “Catalogue of the Library of the British Museum (Natural
History), 1922, vol. 6, p. 218] shows a species of the genus Macro-
schisma with a strongly sinuate posterior end. At first I felt that
Lightfoot’s name should replace Dolichoschisma munita Iredale (1940,
p. 431) restricted to Western Australia (Cotton, 1959, p. 79) since this
is the only species known to me with such a sinuate posterior end.

Because, however, this part of Australia was not scientifically
explored until Baudin’s expeditions at the beginning of the nineteenth
century, it seems unlikely that specimens from these coasts could have
reached London in the 1770s. As these figures are unaccompanied
by any textual material that might furnish us with a clue, I am led to
the conclusion that either an abnormal specimen was figured or that
the drawing is a poor and misleading one. For this reason I feel this
name should be regarded as a doubtful one.

1620 ‘Three rare species of Conus in pairs . . . Leoninus, ibid.
KOT AD CO tas Wogat e
Conus striatus Linnaeus, 1758
Conus leoninus [Lightfoot, 1786], p. 72.
See Kohn, 1963, pp. 161-162.

1705 ‘Four cards of rare species of Voluta, viz... . filosa, S.
Martyn, vol. I. fig. 22. 1.”
Mitra (Cancilla) filaris (Linnaeus, 1771)
Voluta filosa Born, 1778, p. 212.
Voluta filosa (Lightfoot, 1786], p. 76.

1709 “Patella Calyptra, vol. I. Martyn, fig. 18, k... .”
Nomen dubium
This figure seems to resemble most nearly a specifically indeter-
minable specimen of Capulus which has assumed the nodulose sculpture
of the shell that formed its substratum.

1711 ‘“Voluta ... angulata, S. Martyn, vol. IV, 1325.”
Turbinella angulata (Lightfoot, 1786)
Voluta angulata [Lightfoot, 1786], p. 76.
Voluta scolymus Gmelin, 1791, p. 3553.
Xancus angulatus Solander.—Abbott, 1950, p. 204.
Not Voluta angulata Swainson, 1821 = Zidona dufresnet (Donovan),
1823).

NO. 3579 PORTLAND CATALOGUE—REHDER 15

I have elsewhere (Rehder, 1967, in press) discussed at some length
the unfortunate official restitution of Turbinella Lamarck 1799 over
Aancus Réding, 1798. For the sake of stability and to prevent no-
menclatorial anarchy, however, I follow the rulings of the Interna-
tional Commission of Zoological Nomenclature even though I may not
agree with some of these decisions.

1714 “A fine pair of Conus araneosus, S. or Spider’s Web Cones,
from China . . . rare, Martyn, vol. II. 676.”
Conus araneosus [Lightfoot, 1786]
Conus araneosus [Lightfoot, 1786], p. 76.
Conus arachnoideus Gmelin, 1791, p. 3388.
Conus araneosus Solander.—Tomlin, 1937, p. 213.

1718 “A large and fine, and 2 small specimens of Patella Unguis, L.
(Mytilus Lingua, or green Duck’s-Bill Limpet, S.) Humphrey’s
Conchology, Pl. 2. fig. 2. from Amboyna . . . extremely scarce.”’
Lingula lingua (Lightfoot, 1786)

Mytilus lingua (Lightfoot, 1786], p. 77.

Pharetra monoculoides Réding, 1798, p. 159.

Lingula anatina Lamarck, 1801, p. 141.

Mytilus lingua Dillwyn, 1817, p. 322.

Lingula unguis Linnaeus.—Dall, 1920, p. 262 (not Patella unguis
Linnaeus, 1758).

Lingula lingua (Solander).—Uchinomi, 1956, p. 35.

The use of Solander’s name, as initiated by Uchinomi, in place of
L. anatina Lamarck, is made more acceptable by the knowledge that
Lamarck’s name would have been otherwise replaced by monoculoides
Réding, 1798.

Later (see lot 3717) Lightfoot lists the same shell as Patella unguis
L., a reminder of the long-standing but erroneous application of
Linnaeus’ name to this species of Lingula.

1873 “A neat small pair of Voluta Ancilla, S. from the Straits of
Magellan, very rare, D’ Avila’s Catalogue, vol. I. pl. 8. fig. 5.”
Adelomelon ancilla (Lightfoot, 1786)

Voluta ancilla (Lightfoot, 1786], p. 84.

Voluta spectabilis Gmelin, 1791, p. 3468.

Voluta ancilla Lamarck, 1811, p. 69.

Adelomelon ancilla Solander.—Clench and Turner, 1964, p. 152.

1914 “A fine and large variety of Trochus Solaris, S. from New
Zealand, very rare . . . Martyn, vol. I. fig. 30. 9.”
Astraea imperialis (Gmelin, 1791)
Trochus solaris (Lightfoot, 1786], p. 86.
Not Trochus solaris Linnaeus, 1766.
1930 “— 4 eggs of Helix ovipara, Lister, 23. 21.”

Strophocheilus oblongus (Muller, 1774)
Helix ovipara (Lightfoot, 1786], p. 87.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

1940 “A fine pair of Voluta ... Pepo, S. Martyn, vol. III. fig.
768-770.”
Cymbium pepo (Lightfoot, 1786)
Voluta pepo [Lightfoot, 1786], p. 87.
Voluta neptunt Gmelin, 1791, p. 3467.
Voluta navicula Gmelin, 1791, p. 3467.
Cymbium (Cymba) pepo Solander.—Abbott, 1962, p. 103.
1942 “. .. Trochus ... granosus from New Zealand, Martyn, vol.
LG OTR ED Te oct
Modelia granosa (Martyn, 1784)
Trochus granosus [Lightfoot, 1786], p. 87.
Turbo granosus Holten, 1802, p. 69.
Turbo rubicundus Reeve, 1842, p. 168.
Not Trochus granosus Lam., 1822 (=T. ochroleucus Gmelin, 1791)
Martyn’s name, although appearing in a nonbinomial work, was
validated in 1957 by the International Commission of Zoological
Nomenclature under its Plenary Powers (Opinion 479).

1947 ‘. ..2 of Lepas vittata, S. Seba IV. tab. 16. no. 5. middle
EGU Cont des
Conchoderma virgata (Spengler, 1790)
Lepas vittata [Lightfoot, 1786], p. 88 [momen oblitum].
Lepas virgata Spengler, 1790, p. 207.
Lepas vittata Wood, 1815, p. 69.
Cineras vittata Leach, 1824, p. 170.
Conchoderma virgata (Spengler).—Pilsbry, 1907, p. 99.

The “TV” in the Seba reference is an error for “III’’.. Wood cites
Solander’s manuscript as the source for his use of this name, which
was used also by certain later authors, among them Brown (1827,
pl. 5, fig. 16). Darwin (1851, p. 146) used Spengler’s name virgata,
a designation that has been followed by all subsequent workers.

1960 ‘“. .. a fine Buccinum pustulosum, S. Rum. 49. B. rare.”
Argobuccinum pustulosum (Lightfoot, 1786)
Buccinum pustulosum [Lightfoot, 1786], p. 88.
Murex argus Gmelin, 1791, p. 3547.
Buccinum pustulosum S.—Dall, 1921, p. 125.

The figure given by Rumphius, which is the first reference cited
by Gmelin for his Murer argus, represents the form with the
markedly ocellated bands, typical of the South African form. Light-
foot’s earlier name must, therefore, replace that of Gmelin.

1963 “.......Balanus, .S.\ Gepas'4S:) ~ osponderesusysS. Inster:
442, 284.”
Tetraclita squamosa stalactifera Lamarck

Balanus ponderosus (Lightfoot, 1786], p. 89 [momen oblitum].
Balanus stalactiferus Lamarck, 1818, p. 394.
Tetraclita squamosa stalactifera Lamarck.—Pilsbry, 1916, p. 254.

No. 3579 PORTLAND CATALOGUE—-REHDER 17

The form, sculpture, and locality cited under Lister’s figure seem to
point to this tropical Western Atlantic barnacle, but because Light-

foot’s name has never been used for this subspecies, it should be
- considered a nomen oblitum.

2116 # “A large and a small specimen of Voluta incompta, S. from
the South Seas, extremely scarce . . . Martyn, vol. fig. 19. 1.”
Mitra terebralis Lamarck, 1811

Voluta incompta [Lightfoot], 1786, p. 96 [momen oblitum].
Voluta tesselata Martyn, 1784 (nonbinominal).
Mitra terebralis Lamarck, 1811, p. 201.

| 2120 ‘A very fine specimen of Argonauta nodosa, S. or tuberculated
| Paper Nautilus, from the Cape of Good Hope, rare—Rum.
Sate?
Argonauta nodosa [Lightfoot, 1786]

Argonauta nodosa (Lightfoot, 1786], p. 96.
Argonauta tuberculata Réding, 1798, p. 71.
Argonauta tuberculata Shaw, 1812, pl. 995.
Argonauta tuberculosa Schumacher, 1817, p. 260.
| Argonauta nodosa Solander.—Tryon, 1879, p. 140.
2122 “A small but fine specimen of Voluta Cithara, S. or painted
| Aethiopian Crown, from Japan, very rare... Seba Mus.
Voi 3etv Obs yoiee.
Cymbium (Melo) amphora (Lightfoot, 1786)
Voluta cithara [Lightfoot], 1786, p. 96.
Voluta armata Lamarck, 1811, p. 57.
Not Voluta cithara Lamarck, 1803 =Athleta (Neoathleta) citharoedus
Holten, 1802).
I follow Weaver (1964, p. 2) in considering this long-spined form
to be a synonym of amphora Lightfoot (diadema Lamarck).

2132 “A very perfect and finely colored specimen of Chama Laz-
arus, L. var. a@ pannosus, S. Rum. 483... very rare.”
Chama lazarus_ Linnaeus, 1758
Chama lazarus pannosus [Lightfoot, 1786], p. 96.

The figure in Rumphius depicts merely a variant of the common
C. lazarus.

2148 “. . . two Buccinum testudo, S. Seba III. 70. fig. 2, 3, and
4. of the Bucc. Harpa L.”
Nomen dubium
Not Harpa testudo Donovan, 1822 (=H. ventricusa Lamarck 1816).
The figures referred to by Lightfoot in the ‘Catalogue’ are the last
three figures (unnumbered) in the third row from the top on plate
LXX (figure 1 of Harpa would be the second figure from the right in
the second row). Since these three figures represent three different
226-756—67—_2

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121 |

species of Harpa (articularis Lamarck, 1822; major Réding, 1798;
ligata Menke, 1828, respectively), I suggest that this name be con-
sidered a nomen dubium, since we do not know to what species
Lightfoot intended his name to be applied. Donovan later applied
the name H. testudo to still another species (Harpa ventricosa Lamarck,
1816).

inflata Shaw, 1790, based his determination on the numbered figures
on Seba’s plate, ignoring the last four words of the cited reference.

2150 “Venus Punctata, S.. Rum, 43¢Gy. 2 2V’
Meretrix meretrix (Linnaeus, 1758)

Venus punctata (Lightfoot, 1786], p. 98.
Not Venus punctata Linnaeus, 1758,

2158 ‘A fine Arca nodulosa, S. or studded Ark, Gualt. 87.
TG yeah as
Anadara granosa (Linnaeus, 1758)

Arca nodulosa [Lightfoot, 1786], p. 98.
Not Arca nodulosa Miller, 1776 (= Acar scabra (Poli, 1795), fide Lamy,
1907, p. 95).

2190 ‘Two curious and rare species of Mya, viz. Complanata,
Basta oLG0e oman vue wa ese
Elliptio complanatus (Lightfoot, 1786).
Mya complanata (Lightfoot, 1786], p. 100.
Unio violaceus Spengler, 1793, p. 55.
Mya complanata Dillwyn, 1817, p. 51.
Elliptio complanatus (Solander).—Clark and Berg, 1959, p. 21.

2194 ‘Two very large and fine species of Arca, viz. Nodulosa from
China, Gualt. 87. E. and... .”
Anadara granosa (Linnaeus, 1758)
See lot 2158.

2213 “A fine specimen of Mya Gigas, S. from the Mediterranean.
; Last, 414: 2582”
Panope glycymeris (Born, 1778)
See lot 626.

2216 ‘A very fine cluster of Lepas Cornu Copiae, S. D’Argenville,
26: Dim equeraness
Mitella cornucopiae (Lightfoot, 1786).

Lepas cornucopiae (Lightfoot, 1786], p. 101.
Lepas pollicipes Gmelin, 1791, p. 3213.
Pollicipes cornucopia Leach, 1824, p. 171.
Pollicipes cornucopia: Darwin, 1851, p. 298.
Pollicipes cornucopia: Gruvel, 1902, p. 223.
Mitella pollicipes: Pilsbry, 1907, p. 5.

Dall (1921, p. 98), when he identified Solander’s name with Cassis

|
|
|

No. 3579 PORTLAND CATALOGUE—-REHDER 19

As will be seen by the above synonymy, the trivial name cornucopiae
has been used fairly recently by some authors. Dall (1921, p. 127)
had already called attention to the fact that this name had precedence
over Gmelin’s pollicipes.

2220 “A fine Trochus Annulatus, or purple-edged Trochus, Martyn,

Vol. I. fig. 33. r.. . . from K. George’s Sound on the
N.W. Coast of America . . . rare.”
Calliostoma annulatum (Lightfoot, 1786)

Trochus annulatus (Lightfoot, 1786], p. 101.

Trochus annulatus Martyn, 1784 (nonbinomial).

Trochus virgineus Holten, 1802, p. 67.

Calliostoma annulatum Solander.—Abbott, 1954, p. 115.

2220 “. . . and two [Trochus] Canaliculatus, or bronzed Trochus
ibid 33. r. all from K. George’s Sound, on the N.W. coast of
AMCTICH . sca TARO
Calliostoma canaliculatum (Lightfoot, 1786)

Trochus canaliculatus (Lightfoot, 1786], p. 101.

Trochus canaliculatus Martyn, 1784 (nonbinomial).
Trochus doliarius Holten, 1802, p. 67.

Calliostoma canaliculatum (Solander).—Abbott, 1961, p. 63.

The figure cited by Lightfoot for this species is obviously an error
for “32.”’ It should be noted that whereas Martyn gave New Zealand
as the habitat for this and the previous species, Lightfoot cites the
correct locality.

2222 ‘Three curious and rare species of Solen, viz... . Anti-
quatus, Pennant, 46.25 . Hi
Solecurtus (Azorinus) chamasolen (Da Costa, 1778)
Solen cultellus Pennant, 1777, p. 85 (not Linnaeus, 1758).
Solen antiquatus [Lightfoot, 1786], p. 101.
Solen antiquatus Pulteney, 1799, p. 28.
Since Pulteney gives the Solander manuscript as the source of his
name, Lightfoot must have derived his use of antiquatus from the same
source although he does not signify this by the usual letter ‘‘S.”

2274 ‘An exceedingly fine specimen of Voluta gravis, S. with its
Epidermis on, from the Straits of Malacca, Martyn, Vol.
O77?
Turbinella pyrum (Linnaeus, 1758)
Voluta gravis (Lightfoot, 1786], p. 103.
Voluta gravis Wood, 1818, p. 101.
This is not J. rapa Lamarck, 1822 (=ponderosa Lightfoot, 1786)
as Dall states (1921, p. 131).

2284 “. . . Murex plicatus, or plicated Murex, from Falkland’s
Islands, undescribed. Favanne, tab. 79. I... .”
226-756—67_3

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Trophon plicatus (Lightfoot, 1786)
Murex plicatus [Lightfoot, 1786], p. 104.
Buccinum laciniatum Martyn, 1784, fig. 42 (nonbinomial).
Murex lamellosus Gmelin, 1791, p. 3536.
Trophon laciniatus Martyn.—Strebel, 1904, p. 199.
Murex plicatus 8.—Dall, 1921, p. 99.
Not Murex plicatus Gmelin, 1791 (=‘‘Thais”’ sp.).
Not Murex plicatus Dillwyn, 1817 (=Coralliophila erosa (Réding),
1798).
2296 «“‘A fine pair of Buccinum muricatum, Favanne 33, x. 3...
all rare.”’
Colubraria muricata (Lightfoot, 1786)

Buccinum muricatum (Lightfoot, 1786], p. 104.
Murex maculosus Gmelin, 1791, p. 3458.
Not Buccinum muricatum Schroter, 1805 (= Hebra horrida Dunker,
1847).
Shikama (1963, pl. 49, fig. 3) follows Dall (1921, p. 125) in consider-
ing this to be the same as Colubraria obscura (Reeve, 1844); the latter
species, however, is more slender and less strongly sculptured.

2297 +‘... Cardium spinosum, S.... figured in Favanne 52. A. 2.”
Acanthocardia echinatum (Linnaeus, 1758)
Cardium spinosum [Lightfoot, 1786], p. 105.
Cardium spinosum Dillwyn, 1817, p. 115.
Cardium erinaceum Lamarck, 1819, p. 8.
Not Cardium spinosum Sowerby, 1805.—Link, 1807 (= Acanthocardia
aculeatum L., 1758).
2302 ‘Four fine species of Patella, viz. Gorgonica, Humphrey’s
COE a. Seeger at
Patella barbara Linnaeus, 1758
Patella gorgonica (Lightfoot, 1786], p. 105.

2302 “*. © >) [Patella] Pulchra,vor Beautys2bid: 2S. 2”
Patella miniata Born, 1778
Patella pulchra (Lightfoot, 1786], p. 105.

2302 =“. . . and a young one of [Patella] Oculus hirci, ibid. 2. 6.”
Patella oculus Born, 1778
Patella oculushirci (Lightfoot, 1786], p. 105.

2315 “A fine specimen of Voluta incrassa, S. Martyn, Vol. II—
very rare, £.499. 500.”
Oliva incrassata (Lightfoot, 1786)
See lot 264.
It will be noticed that the trivial name is here misspelled ‘“‘incrassa.”’

2327 “A small but extremely fine pair of Murex fimbriatus, or
furbelowed Murex, from Falkland’s Island, Martyn, Vol. 1.
fig. Cav, Paranne; Siew.’

NO. 3579 PORTLAND CATALOGUE—REHDER 21

Trophon geversianus (Pallas, 1774)

Murex fimbriatus [Lightfoot, 1786], p. 106.

Not Murex fimbriatus Brocchi, 1814 (=Fasciolaria (Pleuroploca)
sp.—probably needs a new name).

Not Murex fimbriatus Lamarck, 1822 (=?Ocinebra planilirata Reeve,

1845).

2328 ‘A large and fine Conus Araneosus, S. or Spider’s Web Cone
from Coromandel, very rare. Martyn. Vol. II. 676.”
Conus araneosus [Lightfoot, 1786]

See lot 174.

2230 “Two extremely curious and rare species of Cypraea, viz.

error Pustulata, or orange warted Cowry, from China, Lister,

wer 710:62.......”

2330] #$Jenneria pustulata (Lightfoot, 1786)

Cypraea pustulata (Lightfoot, 1786], p. 106.
Cypraea pustulata Lamarck, 1810, p. 101.
Jenneria pustulata (Solander, 1786).—Keen, 1958, p. 333.

2331 ‘“Serpula attrahens or furbelowed Watering-pot. Humphrey’s
Conch. pl. VII [Error for XIJ] fig. 15. from Madagascar .. .
very rare.’

Penicillus (Warnea) attrahens (Lightfoot, 1786)
Serpula attrahens (Lightfoot, 1786], p. 106.
Aspergillum vaginiferun Lamarck, 1818, p. 480.
Serpula attrahens S.—Dall, 1921, p. 129.

2481 “A fine Trochus sulcatus, from New Zealand, Naturforscher,
Vol. IX. tab. III. fig. 5. 6. Martyn, Vol. I. fig. 33. [error
LOR DER Res
Cookia suleata (Gmelin, 1791)

Trochus sulcatus Martyn, 1784, fig. 35.

Trochus sulcatus [Lightfoot, 1786], p. 13.

Trochus cookiti Gmelin, 1791, p. 3582.

Turbo sulcatus Gmelin, 1791, p. 3592.

Cookia sulcata (Martyn, 1784).—Powell, 1957, p. 90.

Not Trochus sulcatus Lamarck, 1804 (= Astele sp.—needs new name—
Eocene, Paris Basin).

When the name Turbo sulcatus Gmelin, 1791 was placed on the
“Official List of Specific Names in Zoology” in Opinion 479 (Inter-
national Commission of Zoological Nomenclature, 1957: p. 372),
it was not realized that Lightfoot’s name was the first valid taxon for
this species after Martyn. It was also overlooked that Zrochus
cookii Gmelin, 1791 is a synonym of Turbo sulcatus Gmelin.

2516 ‘“Isognoma rigida, S. from Pulo Condore, ist. 227, 62. very
rare.”
Nomen dubium

22 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121 ||

Because Lister’s figure is only an internal view, this name must be»
considered a nomen dubium.

2550 ‘Two fine species of Cardium, viz... . and Hystrix, S.
Gualt. 72. B. both rare.”
Trachycardium aculeatum (Linnaeus, 1767)
Cardium hystrix (Lightfoot, 1786], p. 116.
Not Cardium hystrix Reeve, 1844 (= Fragum (Ctenocardia) symboli-
cum Iredale, 1929).
2554 “A fine pair of Conus mappa, S. from China, very rare. |
Knorr. I. tab. 8. 4.”
Conus ammiralis Linnaeus, 1758
Conus mappa [Lightfoot, 1786], p. 116.
Not Conus mappa Crosse, 1858 (=C. eldredi Morrison, 1935).
2961 ‘A large and very fine variety of Buccinum calcaratum, |
SGU Sia 1 eee TOT en !
Melongena (Volema) hippocastanum (Linnaeus, 1758)
Buccinum calcaratum [Lightfoot, 1786], p. 133.
Volema nuxmoschata Réding, 1798, p. 57.
Pyrula hippocastanum Lamarck, 1816, p. 7.
Murex calcaratus Dillwyn, 1817, p. 710.
Pyrula galeodes Lamarck, 1822, p. 144. |
Dodge (1957, pp. 137-139) has exhaustively detailed the reasons
for the allocation of Linnaeus’ name to this species rather than to the |
species of Muricidae (Thaisinae) to which the Linnean name was |
given by many later authors.

2965 ‘...Cypraea seriata, (rather a young one of Tigris L.)....” |
Cypraea tigris Linnaeus, 1758
Cypraea seriata [Lightfoot, 1786], p. 133.

2965 ‘...[Cypraea] nebulosa, S. (a young one of Mauritiana, L.).” |
Cypraea mauritiana Linnaeus, 1758

Cypraea nebulosa (Lightfoot, 1786], p. 133.
Not C. nebulosa Gmelin, 1791 (=C. stercoraria Linnaeus, 1758).

2967 ‘Two fine specimens of Strombus truncatus, an undescribed |

species of L. showing the different stages of growth, from the
H. Indies, rare s.. DP’ Avila, vol. Viab: 12. 14.”
Lambis truncata truncata (Lightfoot, 1786)

Strombus truncatus [Lightfoot, 1786], p. 133.

Strombus bryonia Gmelin, 1791, p. 3520.

Lambis davilae Réding, 1798, p. 66.

Strombus truncatus Dillwyn, 1817, p. 659.

Lambis truncata truncata (Humphrey, 1786).—Abbott, 1961, p. 155.

2983 “Three curious species of Mya, viz... . ovalis, Lister 146.1...”
Unio ovalis ovalis (Lightfoot, 1786)

NO. 3579 PORTLAND CATALOGUE—REHDER 23

Mya ovalis (Lightfoot, 1786], p. 134.

Mya batava Dillwyn, 1823, p. 13 (not Maton and Rackett, 1807.)
Unio cytherea Kiister, 1833.

Unio crassa cytherea Kiister.—Haas, 1940, p. 129.

Mya ovalis 8.—Dall, 1921, p. 128 (in part).

Not Mya ovalis Pulteney, 1799 (~Unio pictorum Linnaeus, 1758).

A comparison of the figure in Lister, which is accompanied by the
words “a Danubio,” with various figures and descriptions of the
_ polymorphic species and subspecies of Unio occurring in Europe leads
me to the conclusion that Lightfoot’s name must be used for the
subspecies that Haas (1940) called Unio crassus cytherea Kiister, 1833.
Because the name ovalis is two years older than crassus, the central
European, Danubian form becomes the nominate form, and the
western European subspecies becomes Unio ovalis crassus Retzius.
I leave the judgment of the correctness of this assignment to my
European colleagues more familiar with this difficult freshwater fauna.

2995 ‘Patella pulchra, or beauty Limpet, Humphrey’s Conch. pl.
2. fig. 8. from the Cape of Good Hope... .”
Patella miniata (Born, 1778)
See lot 2302.

3020 ‘‘Voluta virescens, S. an extreme scarce species of the Music
kind, from Guinea—Martyn, vol. III. fig. 932. 933.”
Voluta virescens (Lightfoot, 1786]
See lot 610.

3028 ‘‘A large and fine specimen of Placuna Ephippium, S. (Anomia,
L.) from China—very rare.”
Placuna [Lightfoot, 1786]
The ‘‘S” signifies that Solander is the author of the combination,
but not of the trivial name, whose author is Linnaeus.

3030 “. .. Bulla vesicaria, S. from the W. Indies, rbid [Seba III.
tab. 38.], fig. 46. 48 .. . both rare.”
Hydatina vesicaria (Lightfoot, 1786)

Bulla physis Linnaeus, 1758 (in part).
Bulla vesicaria (Lightfoot, 1786], p. 136.
Hydatina vesicaria Solander.—Abbott, 1954, p. 276.

For a discussion of the status of this species, see Bartsch, 1940,
Proc. Biol. Soc. Washington, vol. 53, p. 92.
3039 ‘A large specimen of Voluta Scafa, S. in the utmost perfection,
from Guinea... Martyn, vol. III tab. 70. 764.”
Cymbium cymbium (Linnaeus, 1758)
See lot 969.

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121)

3040 ©‘‘A large and fine Pinna rigida, S. Knorr II. 26,1... very rare.’’
Atrina rigida (Lightfoot, 1786) J
Pinna rigida (Lightfoot, 1786], p. 136.

Pinna rigida Dillwyn, 1817, p. 327.
Atrina rigida Solander.—Turner and Rosewater, 1958, p. 312.
9941 “A cluster of Isognoma Perna, S. (ostrea, L.) enveloped in |
Sponge, from China, rare.” |
Isognomon [Lightfoot, 1786]

See lot 968.

3049 ‘A large and fine specimen of Tellina marginalis, S. List. 387.” |

Tellina laevigata Linnaeus, 1758 |
Tellina marginalis [Lightfoot, 1786], p. 137.

9054 “Voluta Haustrum, S. or banded coronated Melon, very |
rare, from China, Martyn, III. 781.”
Cymbium amphora (Lightfoot, 1786)

See lot 768.

3061 ‘A very fine specimen of Voluta ancilla, S. from the Straits of |
Magellan, extremely scarce, D’ Avila, vol. I. pl 8. fig. s.” |
Adelomelon ancilla (Lightfoot, 1786)

See lot 1873.

9074 1‘. . . a stone with various specimens of Venus arctica
Saaniity Hash, 226M On Baa ed
Hiatella arctica (Linnaeus, 1758)

Venus arctica [Lightfoot, 1786], p. 138.

9991 “A cluster of three, and a single specimen of Ostrea purpurea
S. from New Holland, very rare—Born. mus. caes. Tab. 6. ig.
1022.2

Ostrea cucullata Born, 1778 |
Ostrea purpurea (Lightfoot, 1786], p. 139.

9993 ‘A large and fine Buccinum monodon, S. or Unicorn, from
Terra del Fuego, Martyn, vol. I. fig. 10e....”
Acanthina monodon (Pallas, 1774)

See lot 372.

3117 “A large fine pair of Patella oculus hirci, or Goat’s-eye Limpet
one of them in its native state, the other polished, from the Cape
of Good Hope . . . Humphrey’s Conch. 2. 6.”

Patella oculus Born, 1778
See lot 2302.

3119 “A large and fine specimen of Placuna placenta, S. (Anomia
L.) or the Window Shell, from China . . . rare.”
Placuna (Lightfoot, 1786]

See lot 3028.

No. 3679 PORTLAND CATALOGUE—REHDER 25

3142 ‘A fine pair of Voluta muricata, S. one with the epidermis on,
Inster, 810. 19. from the W. Indies . . . rare.”
Vasum muricatum (Born, 1778)
Voluta muricata (Lightfoot, 1786], p. 142.

3158 “A pair of large and fine Buccinum taurinum, 8S. Lister,
841. 69. from China. . . very rare.”
Terebra taurinum (Lightfoot, 1786)
Buccinum taurinum (Lightfoot, 1786], p. 142.
Epitonium feldmanni Réding, 1798, p. 94.
Terebra flammea Lamarck, 1822, p. 284.
Terebra taurinum Solander.—Abbott, 1954, p. 265.

3161 “A fine specimen of Voluta elongata, S. or Midas’s ear Land

Snail, from New Caledonia, very rare. . . Martyn, vol. I.
Jig. 25. n.”
Placostylus elongatus (Lightfoot, 1786)
See lot 707.
3235 “. . . a fine specimen of Turbo cornutus, a new species allied
to Chrysostomus, L. but with a silver mouth, D’ Avila. Vol.
Ble Septge le

Turbo (Batillus) cornutus [Lightfoot], 1786

Turbo cornutus [Lightfoot, 1786], p. 147.
Turbo cornutus Gmelin, 1791, p. 35938.
Turbo cornutus Solander.—Shikama, 1963, pl. 12.

3307 ‘“‘A fine young specimen of Strombus truncatus, D’ Avila, vol. I.

LOO ND a
Lambis truncata truncata (Lightfoot, 1786)
See lot 2967.
3356 ‘Four curious species of Buccina, viz... . one of Iris, S.

Martyn, vol. I. fig. 2.6... .”
Latirus iris (Lightfoot, 1786)
See lot 30.

3384 ‘Patella Auricularia, an extremely rare species, from Amboyna
se a unp. 40. IN.”
Dolabella auricularia (Lightfoot, 1786)
Scapula Martyn, 1784, fig. 99 (nonbinomial).
Patella auricularia (Lightfoot, 1786], p. 154.
Dolabella callosa Lamarck, 1801, p. 62.
Dolabella rumphii Blainville, 1819, p. 395.
Dolabella auricularia (Solander).—Kira, 1959, p. 102.

3388 ‘‘A fine pair of Helix ovipara, List. 1055. 1. from Surinam... .”
Strophocheilus ovatus (Miller, 1774)
Helix ovipara [Lightfoot, 1786], p. 155.

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

3389 ‘Two fine species of Cardium, viz. . .. impressum, S. Born.
Mus. tab. 2. fig. 15, 16. both from China and rare.”
Corculum impressum (Lightfoot, 1786)

Cardium impressum [Lightfoot, 1786], p. 155.

Cardium roseum Gmelin, 1791, p. 3245.

Corculum humanum Réding, 1798, p. 189.

Cardium junoniae Lamarck, 1816, p. 17.

Corculum impressum (Solander).—Shikama, 1964, p. 74.

Not Cardium impressum Deshayes, 1842 (= Protocardia sp., Creta-
ceous, France).

3494 “. . . Trochus punctulatus of Martyn, vol. I. fig. 36. r... .”
[error Calliostoma (Mauriella) punctulatum Martyn, 1784
for Trochus punctulatus [Lightfoot, 1786], p. 155.
3394] Trochus diaphanus Gmelin, 1791, p. 3580.
Not Trochus punctulatus Gmelin, 1791 (=?Cantharidus turriculum
Phil., 1855).
3494“) 2 Trochus]) qranosus, 7bid. 37. Fs 2

Modelia granosa (Martyn, 1784)
See lot 1942.

Both of the above two Martyn names were validated by the Inter-
national Commission of Zoological Nomenclature in Opinion 479
(1957).

3411 “. .. and two fine varieties of [Conus] nocturnus, S. Martyn,
IT. 687. and 688. all from China, and rare.”

Conus nocturnus [Lightfoot, 1786]

Conus nocturnus (Lightfoot, 1786], p. 156.

Conus nocturnus Hwass, 1792, p. 611.

Conus nocturnus Solander.—Tomlin, 1937, p. 281.
Conus nocturnus [Lightfoot]—Kohn, 1963, p. 162.

3458 “A most beautiful variety of Mytilus pictus, S. or painted
Muscle, bright green, waved with brown, from the Medi-
terranean .. . rare. Knorr. IV. tab. 15. fig. 5.”

Perna picta (Born, 1778)
Mytilus pictus [Lightfoot, 1786], p. 158.

Knorr’s figure is the only illustration cited by Born (1778, pp.
111-112) for his Mytilus pictus. It is also listed by Gmelin, but with a
question mark, under his Mytilus afer (1791, p. 3358), which is a new
name for Mytilus pictus Born.

Dall believes this to be the same as Mytilus ungulatus Linnaeus,
which name, however, seems to have been based on an abnormal speci-
men of Mytilus edulis as Dodge (1952, pp. 214-215) has stated. Lamy
(1936, p. 130) places Mytilus pictus Born in the synonymy of M. perna
Linnaeus, but for the present I maintain it as a distinct species although
it may turn out to be a subspecies or even ecological form of the latter.

NO. 3579 PORTLAND CATALOGUE—REHDER Di

3487 “A large and fine violet Solen, from China . . . extremely
rare. Solen rostratus, S. Valentyn Bivalves, no. 5.”
Hiatula diphos (Linnaeus, 1771)

Solen rostratus [Lightfoot], 1786, p. 160.
Solen rostratus Lamarck, 1818, p. 456.

It has been quite generally overlooked that Hiatula Modeer, 1793,
is an earlier name for Soletellina Blainville, 1824. H. and A. Adams
(1856, p. 392) used it correctly in place of Soletellina, as did Stoliczka
(1870, p. 114) and Tryon (1884, p. 167). Stoliczka (1870) designated
Solen diphos Linnaeus as the type. Thiele (1934, p. 909) thought
it was preoccupied by Hiatula Martini, 1774, which is, however, an
invalid name. Winckworth (1935, p. 322) in discussing Modeer’s
genera was unaware of Stoliczka’s action and designated Mya truncata
Linnaeus as type, hoping thereby to bury Hiatula in the synonymy of
Mya.

Although Hiatula (as Soletellina) has commonly been considered a
subgenus of Sanguinolaria Lamarck, 1799, I feel that the differences
between these two groups in the nature of the hinge, pallial sinus,
and general shape are of sufficient magnitude to justify their being
considered distinct genera.

3491 “A large and fine specimen of Conus fuscatus, S. or bastard
Imperial Crown, from China, very rare . . . Martyn, vol.
II. fig. 693.
Conus imperialis Linnaeus, 1758

Conus fuscatus Born, 1778, p. 126.
Conus fuscatus [Lightfoot, 1786], p. 160.

3553 ‘“Two specimens of a very uncommon species of terrestrial
Snail, nearly allied to Helix Ovipara, List. 1055. 1. the
country unknown.”

Strophocheilus ovatus (Miiller, 1774)
See lot 3388.

3561 “A very fine specimen of Bulla Zonata, S..... Born.
Mus. Caes. tab. 9. fig. 1.”
Hydatina zonata (Lightfoot, 1786)

Bulla amplustre Born, 1780, p. 204 (not Linnaeus, 1758).

Bulla zonata (Lightfoot, 1786], p. 164.

Bulla velum Gmelin, 1791, p. 3433.

Bulla fasciata Bruguiere, 1792, p. 380.

Hydatina (Hydatoria) zonata Solander.—Habe, 1950, p. 18.

Not Bulla zonata Turton, 1834 (=Scaphander lignarius L., 1758,
young).

28 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

The shell figured by Kira (1959, p. 104, pl. 39, fig. 15) as H. zonata,

lacks the white band, and may be a distinct species (?H. inflata
Dunker, 1877).

3624 “Seven curious specimens of Solens, viz. a small one of
Rostratus, Valentyn. Bwalves, no.5... .”
Hiatula dipkos (Linnaeus, 1771)
See lot 3487.

3655 “Two very fine specimens of Strombus truncatus, showing

the different stages of growth, from China, very rare—D’ Avila, .

vol. I. pl. 12 and 14.”
Lambis truncata truncata (Lightfoot, 1786)
See lot 2967.

3696 “A very fine pair of Voluta Incrassata, S. extremely scarce .. .
Martyn, vol. IT. 499. 500.”
Oliva incrassata (Lightfoot, 1786)
See lot 2315.

3711 “A large and complete specimen of Voluta Nobilis, S. from

China, extremely scarce ... Martyn, vol. III. fig. 775.
Lie.)
Aulica nobilis (Lightfoot, 1786)

See lot 89.

3734 “A large and very perfect specimen of Argonauta nodosa,
S. or tuberculated paper Nautilus, from the Cape of Good
Hope, very rare, Rum. 18. 1.”

Argonauta nodosa (Lightfoot, 1786]
See lot 2120.

3741 “Two curious varieties of Ostrea purpurea, S. from New
Holland, and an odd valve of the same species, from China,
very scarce. . . Born. mus. caes. pl. 6. fig. 11. 12.”
Ostrea cucullata Boon, 1778
See lot 3091.

3744 ‘A very fine and large specimen of Placuna Ephippium,
S. (Anomia, LZ.) or the Saddle Shell, very rare, from China.”
Placuna (Lightfoot, 1786]
See lot 3028.

3745 ‘A very fine variety of Helix Ovipara, Lister, tab. 23. having
a white margin to the mouth, with three of its eggs, and a
young shell as taken out of the Egg from St. Vincents. .
extremely curious and very rare.”
Strophocheilus oblongus (Miiller, 1774)
See lot 1930.

A I RL A A LOS -—-—--

NO. 3579 PORTLAND CATALOGUE—REHDER 29

3748 “A very perfect specimen of Argonauta hians, S. a scarce
variety of the brown paper Nautilus, from China, very
roroews\)., dumy18, Be?’

| Argonauta hians (Lightfoot, 1786]

See lot 1055.

3751 “A fine specimen of Voluta virescens, S. or brindled Music,

from Guinea, very rare. . . Martin, vol. III. fig. 932. 933.”
Voluta virescens [Lightfoot, 1786]
. See lot 610.
3758 “A very fine Bulla Zonata, from China, extremely rare, en-
closed in a small glass case. . . Born, mus. caes. tab. 9.
sfegeilen:

Hydatina zonata (Lightfoot, 1786)
See lot 3561.

3761 “A large and fine specimen of Venus nimbosa, S. from Florida,
very rare. . . Favanne, pl. 49, fig. I.”
Macrocallista nimbosa (Lightfoot, 1786)
Venus nimbosa [Lightfoot, 1786], p. 175.

Venus gigantea Gmelin, 1791, p. 3282.
Macrocallista nimbosa Solander.—Clench, 1942, p. 5.

3794 ‘Nine curious and rare species of Helix, among which are.
insignita. Lister, 67,68. . . the last undescribed.”
Carocolus marginella (Gmelin) subsp.

Helix insignita (Lightfoot, 1786], p. 176 [momen oblitum].

3798 “A very fine specimen of Ostraea purpurea, S. rich in colour,
from New Holland, exceedingly scarce. Born. mus. caes. tab.

Ge figalie 122:
Ostrea cucullata Born, 1778

See lot 3091.
3802 ©‘‘A very fine specimen of Helix undata, a carinated umbilicated
Land Shell, the Country unknown. Lister, 76. Favanne,

tab. 63. fig. Gro
Solaropsis undata (Lightfoot, 1786)
Helix undata [Lightfoot], 1786, p. 177.
Helix pellis-serpentis Gmelin, 1791, p. 3620.
Helix undata.—Dall, 1921, p. 99 (in part).
Not Helix undata Gmelin, 1791 (a nomen dubium, possibly young

Cochlitoma sp.).
I am restricting Lightfoot’s name to Lister’s figure, since the figures
in Favanne look to me more like S. gibboni Pfeiffer. The replacement
of Gmelin’s name by wndata is more readily acceptable since Pilsbry

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

in his “Manual of Conchology” (1889, p. 178) gave the name pellis-
serpentis erroneously to the species with two deep pits in the last
whorl, which must bear the name S. constrictor Hupe, 1853. Dall
has identified Lightfoot’s undata with S. brasiliana Deshayes, 1832.

3817.‘ ‘‘A large and fine specimen of Trochus Onustus, or the Carrier,
from Guadalupe, very rare. Favanne 12. C.1. 2.”
Xenophora conchyliophora (Born, 1780)

See lot 31.

3825 “A very fine specimen of Cardium protrusum, S. a curious
variety of the Venus’s Heart Cockle, from China, very rare.
Lister, 319. 156.”
Corculum impressum (Lightfoot, 1786)

Cardium protrusum [Lightfoot, 1786], p. 178.
See lot 3389.

3828 “A large and very fine Turbo undulatus, or waved Emerald
Turbo, extremely scarce, from Van Dieman’s Land, New
Holland. Martyn, Vol.1. fig. 29, P.”

Subninella undulata (Lightfoot, 1786)
See lot 408.

3830 ‘A very fine specimen of Patella umbraculum or Umbrella
Limpet, from China, extremely scarce. Humphrey’s Conch.
pl. 5, fig. 5. which was taken from this shell.”

Umbraculum umbraculum (Lightfoot, 1786)
Patella umbraculum [Lightfoot, 1786], p. 178.
Patella sinica Gmelin, 1791, p. 3705.

Patella umbellata Gmelin, 1791, p. 3720.

Patella wmbracula Réding, 1798, p. 6.
Umbraculum umbraculum (Humphrey).—Kira, 1959, p. 103.

3844 “A very perfect specimen of Conus pulcher, an undescribed
species from the Coast of Guinea . . . extremely rare, List.
rip ae
Conus pulcher [Lightfoot, 1786]

Conus pulcher [Lightfoot, 1786], p. 179.

Conus leoninus Gmelin, 1791, p. 3386 (not Lightfoot, 1786).
Conus papilionaceus Hwass, 1792, p. 665.

Conus pulcher [?Humphrey].—Tomlin, 1937, p. 298.

Not Conus pulcher A. Ad., 1854.

3447 “A large and fine Anomia Ephippiun, L. (placuna, 8.) from
[Error China .. . extremely rare.”

for Placuna [Lightfoot, 1786]

3847] See lot 1315.

NO. 3579 PORTLAND CATALOGUE—REHDER ol

3853 ‘A very fine specimen of Patella pulchra, Humphrey’s Conch.
pl. 2. fig. 8... the first is a native of the Cape of Good
EVO pe. 323)
Patella miniata Born, 1778
See lot 2302.

3866 ‘Two fine specimens of a curious and beautiful variety of
Conus undulatus, S. extremely scarce, Gualt. 25. I.”
Conus textile Linnaeus, 1758

Conus undulatus (Lightfoot, 1786], p. 180.
Not Conus undulatus Sby., 1858 (=?C. mucronatus Reeve, 1843).

3874 “A large and beautiful specimen of Voluta Amphora, S. or
clouded Aethiopian Crown. Martyn, vol. III. fig. 780.”
Cymbium amphora (Lightfoot, 1786)

See lot 708.

3902 “A very fine specimen of Voluta Cithara, S. having its Epi-
dermis on, from Japan, extremely scarce.—Seba ILI. 65. 1, 2.”
Cymbium (Melo) amphora (Lightfoot, 1786)

See lot 2122.

3906 “Nautilus scrobiculatus, S. or great umbilicated Nautilus, from
New Guinea, very rare... Lister, 552. 4... Knorr IV.
D225 ie’?
Nautilus scrobiculatus [Lightfoot, 1786]
Nautilus scrobiculatus [Lightfoot, 1786], p. 182.
Nautilus scrobiculatus Dillwyn, 1817, p. 339.

Nautilus umbilicatus Lamarck, 1822, p. 633 (not Linnaeus, 1758).
Nautilus scrobiculatus Solander.—Shikama, 1964, p. 101.

3911 ‘A cluster of Balanus Tulipa, Ellis. Philosoph. Trans. fig. 10,
partly over-run by a Gorgonia pretiosa, very curious and rare,
from Sicily.”

Balanus tulipiformis Darwin, 1854
Balanus tulipa (Lightfoot, 1786], p. 183.

Balanus tulipa Poli, 1791, p. 24.
Not Lepas tulipa Miiller, 1776 (=B. tintinnabulum L., 1758).

3913 “A capital specimen of Voluta Imperialis, S. or Great horned
Wild Music, from Luconia, extremely scarce . . . Martin,
vol. IV. 934. 935.”
Aulica imperialis (Lightfoot, 1786)
Voluta imperialis [Lightfoot, 1786], p. 183.

Voluta imperialis Lamarck, 1811, p. 62.
Aulica imperialis Solander.—Abbott, 1962, p. 98.

ane PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

3924 ‘A large and finely coloured specimen of Helix Undata, an
undescribed species of Land Snail, from whence un-
known .. . Favanne., pl. 63, fig. G. 3.”
Solaropsis undata (Lightfoot, 1786)
See lot 3802.

3926 ©6©“‘A large and fine specimen of Voluta Nobilis, S. an extremely
rare, and very beautiful shell of the Wild Music kind, from
China. . . . Lister, 799.6 . . . Martin III. 774.”
Aulica nobilis (Lightfoot, 1786)
See lot 89.

3939 ‘A fine Serpula Tortuosa, a new and undescribed Species, the
country unknown, Humph. Conch. pl. 11, fig. 4. unique.”
Vermicularia tortuosa (Lightfoot, 1786)

Serpula tortuosa (Lightfoot, 1786], p. 184.
Vermetus costalis ‘Rousseau’ Chenu, 1844, Vermetus, pl. 3.
Vermetus (Vermicularia) tortuosus Solander.—Tryon, 1885, p. 186.

3947 ‘A fine specimen of Arca Labiata, S. or valved Ark, called by
the French, Coquelochon de Moine, D’ Avila, vol. I. pl. 18.
its country unknown.”’

Cucullaea labiata (Lightfoot, 1786)

Arca labiata [Lightfoot, 1786], p. 185.

Arca concamera Bruguiere, 1789, p. 102.

Arca cucullus Gmelin, 1791, p. 3311.

Cucullaea auriculifera Lamarck, 1801, p. 116.

Arca concamerata Dillwyn, 1817, p. 232.

Cucullaea labiata (Solander).—Shikama, 1964, pl. 20, fig. 2.

Not Arca labiata Sowerby, 1833 (—Scapharca perlabiata Grant and
Gale, 1931).

3955 “A very large and fine specimen of Serpula Gigantea, an
undescribed Species, from Luconia, 21 inches long, eatremely
rare... SebalIlIl. 94. The largest figure.”

Kuphus polythalamia Linnaeus, 1766

See lot 97.

3965 ‘‘A fine specimen of Voluta Arausiaca, S. or Prince of Orange’s
Flag Musick from Amboyna, very rare. . . Rum 37. 2.”
Harpulina arausiaca (Lightfoot, 1786)

See lot 611.

3370 “A fine pair of Helix Alba a terrestrial Shell from the £.
[Error Jndies, one of them an exceeding great variety, being left
for handed, or turning the contrary way.—lJuster tab. 33. 32.
3970] and 46.—Favanne, 63. E.”
Nomen dubium
Not Helix alba Bouillet, 1835.

NO. 3579 PORTLAND CATALOGUE—REHDER 33

Each of the three figures cited represents a different species, and the
crudeness of their delineation forbids any exact identification.

3973 ‘A very fine Helix erubescens, a curious non-descript Land
Shell, the country unknown. . . Lister 24. 22.”
Strophocheilus pudicus (Miiller, 1774)

Helix erubescens (Lightfoot, 1786], p. 187.

Bequaert, in his study of the family Strophocheilidae (1948, p. 30)
is probably correct in assigning this name to the synonymy of Miiller’s
species.

3982 ‘A very fine Trochus tectus, the only perfect specimen known,
from the Jsland of All Saints, in the West Indies. A. bad
specimen is figured in Lister, 628.14 . . . ‘tis undescribed.”
Astraea tecta (Lightfoot, 1786)

Trochus tectus [Lightfoot, 1786], p. 187.

Trochus imbricatus Gmelin, 1791, p. 3581.

Astraea tecta tecta Solander, 1786.—Abbott, 1958, p. 30.

Not Trochus tectum Gmelin, 1791 (=Modulus tectum (Gmelin),
1791).

As Abbott has pointed out (1958), Lightfoot’s name does not pre-
occupy Gmelin’s Trochus tectum, since the latter trivial name is a noun
(Latin tectum=roof) in apposition to the genus, while Lightfoot
used the adjective tectus(=covered or tiled).

3983 ‘“‘A very fine specimen of Patella Auricularia, figured Rumph.
tab. 40. fig. N. and called by that author an operculum, it
comes from China, and is extremely scarce.”

Dolabella auricularia (Lightfoot, 1786)
See lot 3384.

4000 ‘A very large and fine specimen of Madrepora Agnus, or
Lamb Madrepora from New Holland . . . extremely scarce.
Seba vol. III. tab. CXI. fig. 6.”
Polyphyllia talpina (Lamarck, 1801)
Madrepora agnus (Lightfoot, 1786], p. 188 [nomen oblitum].
Fungia talpina Lamarck, 1801, p. 370.
Fungia talpa Lamarck, 1816b, p. 237.
Polyphyllia talpina.—Thiel, 1932, p. 89.

4001 “A large and perfect specimen of Murex Tribulus, L. var.
Pecten, or the Venus’s Comb, from China, exceedingly
scarce. Rum. 26. 3.”
Murex pecten [Lightfoot, 1786]

Murez tribulus var. pecten [Lightfoot, 1786], p. 188.
Murex histrix Réding, 1798, p. 145 (not Murex hystrix Linnaeus,
1758).

34 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Aranea gracilis Perry, 1810, pl. 47 (not Murex gracilis Montagn,
1803).

Aranea triremis Perry, 1811, pl. 45, no. 3.

Murex tenuispina Lamarck, 1822, p. 158.

Murex triremis Perry.—Abbott, 1962, p. 71

Not Murex pecten Montfort, 1811 (=M. ternispina Lam., 1822).
Any regret over the loss of the long-well-known name Murer |
tenuispina Lamarck is tempered by the fact that Perry’s name |
triremis had priority and has been used by recent workers for this |
species. Murex ternispina Lamarck is once more available since |
Montfort’s earlier name for this species is preoccupied by Lightfoot’s

name; Kira (1962, p. 63) has so used ternispina.

4002 ‘A very large and fine specimen of Echinus maximus, ez- |
tremely scarce, the country unknown. Seba III, pl. 14. fig. 5. 6.
and Favanne, pl. 58. fig. A. 2.”
Plagiobrissus grandis (Gmelin, 1788)
Echinus maximus (Lightfoot, 1786], p. 188 [momen oblitum].
Echinus grandis Gmelin, 1788, p. 3200.
Spatangus pectoralis Lamarck, 1816b, p. 29.
Plagiobrissus grandis (Gmelin).—Mortensen, 1951, p. 496.

H. L. Clark (1917, p. 207) had pointed out that Lamarck’s pectoralis,
long in use, must yield to Gmelin’s earlier name.

4017 “A fine specimen of Conus Architalassus, S. a curious granu-
lated variety of the High Admiral, extremely scarce, from |
Amboyna. Argenv. Suppl. tab. 1. fig. M. N. Martyn II. tab
min. 26. fig. 1. 2. page 214.”

Conus ammiralis Linnaeus, 1758
Conus architalassus [Lightfoot, 1786], p. 189.
Conus ammiralis coronatus Gmelin, 1791, p. 3389.
Conus archithalassus Hwass, 1792, p. 659.

I follow Kohn (1964, p. 161) in placing this name in the synonymy of
ammiralis L.

4022 ‘A very perfect specimen of Strombus sinuatus, or scollop-
winged Strombus, eatremely rare. Seba. III. pl. 62. fig. 3.
Favanne 22. A. 2.”

Strombus sinuatus [Lightfoot, 1786]

Strombus sinuatus [Lightfoot, 1786], p. 189.

Lambis lobata Réding, 1798, p. 65.

Strombus laciniatus Dillwyn, 1817, p. 663.

Strombus sinuatus Humphrey, 1786.—Abbott, 1960, p. 60.

Not Strombus sinuatus Perry, 1811 (=Lambis scorpius Linnaeus,
1758).

NO. 3579 PORTLAND CATALOGUE—REHDER 35

4023 “A very fine reversed Voluta ponderosa, S. or heavy Volute,
extremely scarce, from the East Indies. A direct one is
Sigured in Favanne, pl. 35. fig. I.”
Turbinella ponderosa (Lightfoot, 1786)

See lot 566.
4030 ‘A very perfect specimen of that beautiful species, the
Voluta Cithara S. from Japan. . . extremely scarce Seba

Wilsiab. 65.1. 2.”
Cymbium (Melo) amphora (Lightfoot, 1786)
See lot 2122.

Systematic List

The following list is arranged systematically by phyla, classes, and families.
All validly proposed Lightfoot names are included, followed by the modern name
of the taxon as I have determined it. In some instances the latter is accompanied
by one or two synonyms in current use.

MOLLUSCA-GASTROPODA
Fissurellidae
macroschisma, Patella=Macroschisma sp.
Patellidae

fungoides, Patella= Patella barbara Linnaeus
gorgonica, P.= Patella barbara Linnaeus
mytiliformis, P.=Nacella mytilina (Helbling)
oculus hirci, P.= Patella oculus Born
pulchra, P.= Patella miniata Born

Trochidae

annulatus, Trochus—Calliostoma annulatum (Lightfoot)
canaliculatus, T.=C. canaliculatum (Lightfoot)
punctulatus, T.=C. (Mauriella) punctulatum (Lightfoot)

Turbinidae

cornutus, Turbo=Turbo (Batillus) cornutus [Lightfoot]
granosus, Trochus= Modelia granosa (Lightfoot)
solaris, Trochus—Astraea imperialis (Gmelin)

sulcatus, Trochus=Cookia sulcata (Lightfoot)

tectus, Turbo=Astraea (Astralium) tecta (Lightfoot)

Pilidae
plicata, Helix=Pila urceus (Miiller)

36 PROCEEDINGS OF THE NATIONAL MUSEUM VOL, 121

Vermetidae
tortuosa, Serpula=Vermicularia tortuosa (Lightfoot)
Capulidaee
calyptra, Patella=Capulus sp.
Xenophoridae

onustus, Trochus=Xenophora conchyliophora (Born)

Strombidae

persicus, Strombus fusus= Tibia insulaechorab (Réding)
tricornis, Strombus=Strombus tricornis [Lightfoot]
truncatus, Strombus=Lambis truncata (Lightfoot)

Cypraeidae

nebulosa, Cypraea=Cypraea mauritiana Linnaeus
pantherina, Cypraea=Cypraea pantherina [Lightfoot]
pustulata, Cypraea=Jenneria pustulata (Lightfoot)
seriata, Cypraea=Cypraea tigris Linnaeus

Cymatiidae

muricatum, Buccinum=Colubraria muricata (Lightfoot)
maculosa (Gmelin)
pustulosum, Buccinum=Argobuccinum pustulosum (Lightfoot)
argus (Gmelin)

Muricidae-Muricinae

elongatus, Murex=Pterynotus (Marchia) elongatus (Lightfoot)
clavus (Kiener)
pecten, Murex tribulus= Murex pecten [Lightfoot]
triremis Perry
tenwspina Lamarck
Muricidae-Thaisinae

monodon, Buccinum=Acanthina monodon (Pallas)

Muricidae-Trophoninae
fimbriatus, Murex=Trophon geversianus (Pallas)
plicatus, Murex=Trophon plicatus (Lightfoot)

laciniatus (Martyn)
Melongenidae

calcaratum, Buccinum= Melongena (Volema) hippocastanum (Linnaeus)

NO. 3579 PORTLAND CATALOGUE—REHDER 37

Fasciolariidae
iris, Buccinum= Latirus iris (Lightfoot)
Olividaee
incrassata, Voluta= Oliva incrassata (Lightfoot)
Mitridae

filosa, Voluta= Mitra (Cancilla) filaris (Linnaeus)
incompta, Voluta= Mitra terebralis Lamarck

Turbinellidae

angulata, Voluta=Turbinella angulata (Lightfoot)

gravis, Voluta=Turbinella pyrum (Linnaeus)

ponderosa, Voluta=Turbinella ponderosa (Lightfoot)
rapa Lamarck

Vasidae
muricata, Voluta=Vasum muricatum (Born)
Harpidae

testudo, Buccinum=Harpa sp.

Volutidae-Volutinae

virescens, Voluta= Voluta virescens [Lightfoot]

Volutidae-Cymbiinae

amphora, Voluta= Melo amphora (Lightfoot)

cithara, Voluta= Melo amphora (Lightfoot)

haustrum, Voluta= Melo amphora (Lightfoot)

imperialis, Voluta=Aulica imperialis (Lightfoot)

melo, Voluta=Melo melo (Lightfoot)

nobilis, Voluta=Volutocorona nobilis (Lightfoot)
scapha (Gmelin)

pepo, Voluta=Cymbium pepo (Lightfoot)

scafa, Voluta=Cymbium cymbium (Linnaeus)

Volutidae-Zidoninae

arausiaca, Voluta=Harpulina arausiaca (Lightfoot)
ancilla, Voluta—Adelomelon ancilla (Lightfoot)

226—-756—67—_—_4

38 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Conidae

araneosus, Conus=Conus araneosus [Lightfoot]
architalassus, Conus=Conus ammiralis Linnaeus
augur, Conus=Conus augur [Lightfoot]
fuscatus, Conus=Conus imperialis Linnaeus
leoninus, Conus=Conus striatus Linnaeus
mappa, Conus=Conus ammiralis Linnaeus
nocturnus, Conus=Conus nocturnus [Lightfoot]
pulcher, Conus=Conus pulcher [Lightfoot]
papilionaceus Hwass
quercinus, Conus=Conus quercinus [Lightfoot]
undulatus, Conus=Conus textile Linnaeus

Terebridae
taurinum, Buccinum=Terebra taurinum (Lightfoot)
Aplysiidae

auricularia, Patella=Dolabella auricularia (Lightfoot)
callosa Lamarck

Umbraculidae
umbraculum, Patella=Umbraculum umbraculum (Lightfoot)
sinicum (Gmelin)
Hydatinidae

vesicaria, Bulla=Hydatina vesicaria (Lightfoot)
zonata, Bulla=Hydatina zonata (Lightfoot)
velum (Gmelin)

Bulimulidae
elongata, Voluta= Placostylus elongatus (Lightfoot)
fibratus (Martyn)
Strophocheilidae

erubescens, Helix=Strophocheilus pudicus (Miller)
ovipara, Helix=Strophocheilus oblongus (Miiller)

Sagdidae
alveolatus, Trochus=Sagda epistylium (Miiller)

Camaenidae

insignita, Helix=Caracolus sp.
otis, Helix=Lampadion otis (Lightfoot)
undata, Helix—Solaropsis undata (Lightfoot)

ee i ii eee —

NO. 3579 PORTLAND CATALOGUE—REHDER 39

Helminthoglyptidae-Cepolinae
vitellus, Helix=Polymita picta (Born)
Helicidae
alba, Helix=nomen dubium

MOLLUSCA-BIVALVIA
Arcidae
fusca, Arca=Arca sp.
nodulosa, Arca=Anadara granosa (Linnaeus)
Cucullaeidae
labiata, Arca=Cucullaea labiata (Lightfoot)
Mytilidae
castaneus, Mytilus= Modiolus vagina Lamarck
pictus, Mytilus= Perna picta (Born)
Pinnidae
rigida, Pinna=Atrina rigida (Lightfoot)
Isognomonidae
Isognoma=Isognomon [Lightfoot]

Tsognomon [Lightfoot]
rigida, Isognoma=Isognomon sp.

Anomiidae

Placuna [Lightfoot]
Ostreidae

purpurata, Ostrea=Ostrea cucullata (Born)

Unionidae
complanata, Mya=Elliptio complanatus (Lightfoot)
ovalis, Mya= Unio ovalis (Lightfoot)

crassa cytherea Kiister
Chamidae

pannosus, Chama lazarus=Chama lazarus Linnaeus

Cardiidae

hystrix, Cardium=Trachycardium aculeatum (Linnaeus)
impressum, Cardium=Corculum impressum (Lightfoot)
humanum Réding
protrusum, Cardium—=Corculum impressum (Lightfoot)
robustum, Cardium—=Dinocardium robustum (Lightfoot)
spinosum, Cardium=Acanthocardia echinatum (Linnaeus)

40 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 122

Solenidae
pallidus, Solen=Solen sp.
Tellinidae
marginalis, Tellna=Tellina laevigata Linnaeus
Garidae

cruenta, Tellina=Sanguinolaria cruenta (Lightfoot)
operculata (Gmelin)
rostratus, Solen=Hiatula diphos (Linnaeus)
Soletellina diphos (L.)

Solecurtidae

antiquatus, Solen=Solecurtus (Azorinus) chamasolen (Da Costa)
plebeius, Solen=Tagelus plebeius (Lightfoot)

Veneridae

nimbosa, Venus= Macrocallista nimbosa (Lightfoot)
punctata, Venus= Meretrix meretrix (Linnaeus)

Myidae
gigas, Mya=Panope glycymeris (Born)
Hiatellidae
arctica, Venus= Hiatella arctica (Linnaeus)
Teredinidae
gigantea, Serpula=Kuphus polythalamia (Linnaeus)
Clavagellidae

attrahens, Serpula=Penicillus (Warnea) attrahens (Lightfoot)
vaginiferum Lamarck

MOLLUSCA-CEPHALOPODA

Argonautidae

hians, Argonauta=Argonauta hians [Lightfoot]
navicula, Argonauta=Argonauta sp.
nodosa, Argonauta= Argonauta nodosa [Lightfoot]

Nautilidae

scrobiculatus, Nautilus= Nautilus scrobiculatus [Lightfoot]

COELENTERATA-ANTHOZOA
Fungiidae
agnus, Madrepora= Polyphyllia talpina (Lamarck)

|

NO. 3579 PORTLAND CATALOGUE—REHDER 41

BRACHIOPODA-INARTICULATA
Lingulidae
lingua, Mytilus=Lingula lingua (Lightfoot)
anatina Lamarck

ECHINODERMATA-ECHINOIDEA
Brissidae
maximus, Echinus= Plagiobrissus grandis (Gmelin)
Echinometridae

anemonoides, Echinus= Podophora atrata (Linnaeus)

ARTHROPODA-CRUSTACEA-CIRRIPEDIA
Lepadidae

cornucopiae, Lepas= Mitella cornucopiae (Lightfoot)
pollicupes (Gmelin)
vittata, Lepas=Conchoderma virgata (Spengler)

Balanidae

ponderosus, Balanus=Tetraclita squamosa stalactifera (Lamarck)
pyramidalis, Balanus=Balanus tintinnabulum (Linnaeus)
tulipa, Balanus=Balanus tulipiformis (Darwin)

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eee ee

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Acanthina monodon, 8, 24
Acanthocardia echinatum, 20
Adelomelon ancilla, 15, 24
Anadara granosa, 18
Aranea gracilis, 34

triremis, 34
Arca concamera, 32

concamerata, 32

cucullus, 32

fusca, 10

imbricata, 11

labiata, 32

nodulosa, 18

tetragona, 11
Argobuccinum pustulosum, 16
Argonauta gondola, 11

hians, 12, 29

navicula, 11

nodosa, 17, 28

tuberculata, 17

tuberculosa, 17
Aspergillum vaginiferum, 21
Astraea imperialis, 15

tecta, 33
Atrina rigida, 24
Aulica imperialis, 31

nobilis, 6, 28, 32

Balanus ponderosus, 16
pyramidalis, 12
stalactiferus, 16
tintinnabulum, 12
tulipa, 31
tulipiformis, 31

Buccinum calcaratum, 22
iris, 7, 18, 25
laciniatum, 20
monoceros, 8
monodon, 8, 24
muricatum, 20
prismaticum, 7
pustulosum, 16
taurinum, 25
testudo, 17

Bulimus bovinus, 9

48

Index

Bulla amplustre, 27
fasciata, 27
physis, 23
velum, 27
vesicaria, 23
zonata, 27, 29

Calcar, 8
Calliostoma annulatum, 19
canaliculatum, 19
punctulatum, 26
Caracolus marginella, 29
Cardium erinaceum, 20
hystrix, 22
impressum, 26
junoniae, 26
magnum, 12
Cardium protrusum, 30
robustum, 12
roseum, 26
spinosum, 20
ventricosum, 12
Cassis inflata, 18
Chama lazarus, 17
lazarus pannosus, 17
Cineras vittata, 16
Colubraria muricata, 20
obscura, 20
Conchoderma virgata, 16
Conus ammiralis, 22, 34
ammiralis coronatus, 34
arachnoideus, 15
araneosus, 15, 21
architalassus, 34
archithalassus, 34
augur, 11
fuscatus, 27
imperialis, 27
leoninus, 14, 30
mappa, 22
nocturnus, 26
papilionaceus, 30
pulcher, 30
quercinus, 13
striatus, 14
textile, 31
undulatus, 31

NO. 3579

Cookia sulcata, 21
Corculum humanum, 26
impressum, 26, 30
Cucullaea auriculifera, 32
labiata, 32
Cymbium amphora, 9, 17, 24, 31, 35
cymbium, 10, 23
flammeum, 9
maculatum, 10
melo, 10, 13
pepo, 16
Cypraea mauritiana, 22
nebulosa, 22
pantherina, 12
pustulata, 21
seriata, 22
tigris, 22
vinosa, 12

Dinocardium robustum, 12
Dolabella auricularia, 25, 33
callosa, 25
rumphii, 25
Dolichoschisma munita, 14

Echinus anemonoides, 7
grandis, 34
maximus, 34
Elliptio complanatus, 13, 18
Ellobium australe, 9
Epitonium feldmanni, 25

Fungia talpa, 33
talpina, 33

Harpa articularis, 18
ligata, 18
major, 18
testudo, 18
ventricosa, 18
Harpulina arausiaca, 8, 32
Helix alba, 32
erubescens, 33
insignita, 29
labyrinthus, 9
otis, 9
ovipara, 15, 25, 27, 28
pellisserpentis, 29
plicata, 8
undata, 29, 32
vitellus, 8
Hiatella arctica, 24
Hiatula diphos, 27, 28

INDEX 49

Hydatina inflata, 28
vesicaria, 23
zonata, 27, 29

Tsognoma, 6
lignea, 6
perna, 24
rigida, 21
Isognomon, 6, 10, 12, 24
perna, 10, 12

Jenneria pustulata, 21
Kuphus polythalamia, 6, 32

Labyrinthus, 9
Lambis davilae, 22
lobata, 34
truncata, 22, 25, 28
Lampadion, 9
labyrinthus, 9
otis, 9
Latirus iris, 7, 13, 25
prismaticus, 7
Lepas cornucopiae, 18
pollicipes, 18
virgata, 16
vittata, 16
Limax fibratus, 9
undulatus, 8
Lingula anatina, 15
lingua, 15
unguis, 15

Macrocallista nimbosa, 29
Macroschisma, 14
Madrepora agnus, 33
Melo diadema, 9
melo, 10
Melongena hippocastanum, 22
Meretrix meretrix, 18
Mitella cornucopiae, 18
pollicipes, 18
Mitra filaris, 14
terebralis, 17
Modelia granosa, 16, 26
Modiola castanea, 13
vagina, 13
Modiolus vagina, 13
Murex argus, 13
calearatus, 22
clavus, 13
elongatus, 13
fimbriatus, 20, 21

50 PROCEEDINGS OF THE NATIONAL MUSEUM

Murex histrix, 33
lamellosus, 20
maculosus, 20
pecten, 33
plicatus, 19, 20
prismaticus, 7
tenuispina, 34
ternispina, 34
tribulus pecten, 33
triremis, 34

Mya batava, 23
complanata, 18
gigas, 9, 18
ovalis, 22, 23
truncata, 27

Mytilus afer, 26
castaneus, 13
edulis, 26
lingua, 15
perna, 26
pictus, 26
ungulatus, 26

Nacella mytilina, 10, 13
Nautilus scrobiculatus, 31
umbilicatus, 31

Oliva angulata, 7
incrassata, 7, 20, 28

Ostrea cucullata, 24, 28, 29
purpurea, 24, 28, 29

Panope glycymeris, 9, 18

Patella auricularia, 25, 33
barbara, 12, 20
calyptra, 14
fungoides, 12
gorgonica, 20
macroschisma, 14
miniata, 20, 23, 31
mytiliformis, 10 , 13
mytilina, 10
oculus, 20 , 24
oculushirci, 20, 24
pulchra, 20, 23, 31
sinica, 30
umbellata, 30
umbraculum, 30
unguis, 15

Penicillus attrahens, 21

Perna picta, 26

Pharetra monoculoides, 15

Phorus onustus, 6

Pila urceus, 8

Pinna rigida, 24

Placenta, 7

Placostylus elongatus, 9, 25

placenta, 7, 12, 14
Plagiobrissus grandis, 34
Plejona muta, 8
Pleurodonte otis, 9
Podophora atrata, 7
Pollicipes cornucopia, 18
Polymita picta, 8
Polyphyllia talpina, 33
Pterynotus elongatus, 13
Pyrula galeodes, 22

hippocastanum, 22

Rostellaria curvirostris, 5

Sagda epistylium, 12
Sanguinolaria, 27
bertini, 7
cruenta, 7, 13
operculata, 7
Scapula, 25
Septaria arenaria, 6
Serpula attrahens, 21
gigantea, 6, 32
tortuosa, 32
Solaropsis brasiliana, 30
constrictor, 30
gibboni, 29
undata, 29, 32
Solecurtus chamasolen, 19
Solen antiquatus, 19
cultellus, 19
diphos, 27
gibbus, 11
pallidus, 11
plebeius, 11
rostratus, 27, 28
truncatus, 11
Soletellina, 27
Spatangus pectoralis, 34
Strombus bryonia, 22
fusus persicus, 5
laciniatus, 34
sinuatus, 34
tricornis, 6
truncatus, 22, 25, 28

VOL. 121
Placuna, 7, 12, 23, 24, 28, 30
ephippium, 7, 12, 23, 28, 30

|

NO. 3579

Strophocheilus oblongus, 15, 28
ovatus, 25, 27
pudicus, 33

Subninella undulata, 8, 30

Tagelus plebeius, 11
Tellina cruenta, 7, 13
cruentae, 7
laevigata, 24
marginalis, 24
operculata, 7
sanguinolenta, 7
semiplanata, 7
Terebra flammea, 25
taurinum, 25
Tetraclita squamosa stalactifera, 16
Tibia insulachorab, 5
Trachycardium aculeatum, 22
Trochus alveolatus, 12
annulatus, 19
canaliculatus, 19
cookii, 21
diaphanus, 26
doliarius, 19
granosus, 16, 26
imbricatus, 33
onustus, 6, 30
punctulatus, 26
solaris, 15
suleatus, 21
tectum, 33
tectus, 33
virgineus, 18
Trophon geversianus, 21
laciniatus, 20
plicatus, 20
Turbinella, 15
angulata, 14
ponderosa, 8, 35
pyrum, 19
rapa, 8, 19
Turbo cornutus, 25
granosus, 16
rubicundus, 16
sulcatus, 21
undulatus, 8, 30

Umbraculum umbraculum, 30
Unio cytherea, 23

ovalis crassus, 22

ovalis ovalis, 22

INDEX 51

Unio—Continued
violaceus, 18

Vasum muricatum, 25
Venus arctica, 24
gigantea, 29
nimbosa, 29
punctata, 18
Vermetus costalis, 32
tortuosus, 32
Vermicularia tortuosa, 32
Volema nuxmoschata, 22
Voluta amphora, 9, 31
ancilla, 15, 24
angulata, 14
anguria, 10, 13
arausiaca, 8, 32
armata, 17
cithara, 17, 31, 35
cymbium, 10
elongata, 9, 25
filosa, 14
gravis, 19
haustrum, 24
imperialis, 31
incompta, 17
incrassa, 20
incrassata, 7, 28
indica, 10
melo, 10
muricata, 25
navicula, 16
neptuni, 16
nobilis, 6, 28, 3:
pepo, 16
polyzonalis, 8
ponderosa, 8, 35
seafa, 6, 10, 23
scapha, 6
scolymus, 14
spectabilis, 15
tesselata, 17
vexillum, 8
virescens, 8, 23, 29
Volutocorona nobilis, 6

bo

Xancus, 15
angulatus, 14
pyrum, 8
Xenophora conchyliophora, 6, 30

U.S. GOVERNMENT PRINTING OFFICE; 1967

Proceedings of

the United States

National Museum

SMITHSONIAN INSTITUTION + WASHINGTON, D.C.

Volume 121 1967 Number 3580

THE MYODOCOPID OSTRACOD FAMILIES PHILOMEDIDAE
AND PSEUDOPHILOMEDIDAE (NEW FAMILY)

By Louis S. KornickER
Associate Curator, Division of Crustacea

Introduction

The Pseudophilomedidae, new family, is represented in the oceans
of the world by only one genus, Pseudophilomedes Miiller, 1894, con-
taining 4 species: 2 in the Mediterranean Sea (Miiller, 1894), 1 in the
eastern Atlantic (Brady and Norman, 1896), and 1 in the western
Atlantic (Kornicker, 1959).

The writer recently received 17 specimens of Pseudophilomedes
ferulanus Kornicker, 1959, as part of a collection of ostracods obtained
by Dr. John H. Day from the Atlantic Shelf off Beaufort, N.C.
Four additional specimens from the same area were in a collection
made in 1964 by personnel on the U.S. Bureau of Commercial Fisheries
R/V Gosnold. This material has been used to supplement 3 specimens
collected from the Great Bahama Bank, upon which the original
description was based (Kornicker, 1959).

I have taken this opportunity to review the genus. Specimens of
Pseudophilomedes foveolatus Miiller, 1894, the type species of Pseudo-
philomedes, were borrowed from the Zoological Museum, Berlin, and
the original description by Miiller (1894) is supplemented in this paper.

Specimens identified by Brady and Norman (1896) as males of

el

a PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Pseudophilomedes foveolatus Miller (called Philomedes foveolata by
Brady and Norman) were borrowed from the collection of the British
Museum. It is concluded that they are not congeneric with Pseudo-
philomedes but instead are members of the genus Huphilomedes Poul-
sen, 1962. They are identified as Huphilomedes asper (Miller, 1894)
and Huphilomedes sp. and are described and illustrated in this paper.
A supplemental description is given of a syntype of Huphilomedes
asper (Miller) borrowed from the Zoological Museum, Berlin.

As a result of this study, the subfamily Philemedinae Miller, 1912,
has been raised to familial rank.

I wish to thank Dr. John H. Day of the Beaufort Marine Laboratory,
Duke University, and personnel of the U.S. Bureau of Commercial
Fisheries for specimens of Pseudophilomedes ferulanus from the
Atlantic Shelf, and Dr. J. P. Harding for permission to study speci-
mens in the collection of the British Museum (Natural History),
identified as Philomedes foveolatus (Miiller) by Brady and Norman
(1896), and Dr. H. E. Gruner for permission to study syntypes of
Pseudophilomedes foveolatus Miller and Philomedes asper Miiller in the
collection of the Zoological Museum, Berlin. I wish also to thank
Miss Caroline Bartlett Gast for preparing plates for publication from
my penciled camera lucida drawings and especially for the carefully
stippled drawing of the ventral view of P. ferulanus (fig. 1). Criti-
cisms of the manuscript by Mr. I. Gregory Sohn and Doctors Raymond
B. Manning, Thomas E. Bowman, and Meredith L. Jones are greatly
appreciated.

Discussion of Classification

Classification of myodocopid Ostracoda was reviewed by Skogsberg
(1920) and more recently by Poulsen (1962). Of particular concern
in the present paper are the taxonomic relationships of Pseudophilo-
medes. Therefore, discussion of classification will be restricted to the
superfamily, Cypridinacea Baird, 1850. When Miiller (1894) de-
scribed Pseudophilomedes, he referred it to the family Cypridinidae
Baird, 1850. Two years later, Brady and Norman (1896) described
the genus Paramekodon (=Pseudophilomedes) referring it also to the
Cypridinidae. Later, Miller (1912) established the subfamily
Philomedinae in the Cypridinidae. He referred to the Philomedinae
the genera Philomedes Liljeborg, 1853, Pseudophilomedes, and Ruti-
derma Brady and Norman 1896. Skogsberg (1920) recognized the Philo-
medinae for Pseudophilomedes and Philomedes but followed Brady and
Norman (1896) in placing Rutiderma in the Rutidermatidae Brady
and Norman, 1896. The Philomedinae sensu Skogsberg has been
recognized by Sylvester-Bradley (1961), Poulsen (1962, 1965), and

NO. 3580 OSTRACOD FAMILIES—KORNICKER 3

Hartmann (1964). However, Poulsen (1962, p. 339) perceived that
Pseudophilomedes might require a higher ranking in the classification:

. according to the description the two genera [Paramekodon Brady and
Norman, 1896, and Pseudophilomedes Miiller, 1894] hold a rather unique position
in the family Cypridinidae (and in the Rutidermatidae and Sarsiellidae) by
having only two endites on the maxilla. Also in the reduced number of bristles
on the basale and endopodite of the mandible and by the reduced Ist and 2nd
endites and the end-joint of the 6th limb the two genera differ widely from all
other Cypridinidae. If further investigations should confirm these differences
these two genera may well have to be included in a family of their own.

After reviewing the genus Pseudophilomedes I find that I concur
with Poulsen; therefore, I have established the new family Pseudo-
philomedidae. The genus Pseudophilomedes Miiller, 1984, with its
synonym Paramekodon Brady and Norman, 1896, is referred to the
Pseudophilomedidae.

Miiller (1912, p. 33) considered Paramekodon Brady and Norman,
1896, to be a synonym of Pseudophilomedes Miller, 1894. Skogsberg
(1920, p. 348) agreed with Miller and stated: ‘In the identification
of Paramekodon with Pseudophilomedes Miiller certainly is correct.”
Sylvester-Bradley (1961, p. 399) also considered Paramekodon to be
a synonym of Pseudophilomedes. Poulsen (1962, p. 339) was of the
opposite opinion and stated: “If the description of Paramecodon [sic]
is reliable, it is based on a single specimen conserved dry, the species
differs in so many respects from Pseudophilomedes that the uniting
of the two genera into one—as done by G. W. Miiller is hardly
admissible.”

Differences between Pseudophilomedes and Paramekodon seem quite
minor; therefore, I have followed Miiller (1912), Skogsberg (1920),
and Sylvester-Bradley (1961) in considering Paramekodon to be a
synonym of Pseudophilomedes. Of possible significance in this regard
is the following statement by Brady and Norman (1896, p. 623) in
their publication in which the genus Paramekodon was established:

It was not until our Monograph was nearly completed, and on the point of
going to press, that there appeared the splendid work of Herr. G. W. Miiller on
the Ostracoda of the Gulf of Naples. This work has, in the case of some few
species, anticipated our descriptions, and we have, as far as possible, rectified
our nomenclature in conformity with it. For the rest we have thought it best
to let our MS. go to press as it was originally written.

It was in the work mentioned in the above statement that Miller
established the genus Pseudophilomedes. It is unfortunate that
Brady and Norman did not have more time to study Miiller’s paper
before publishing their monograph. The fact that they wrongly
considered a species of Philomedes to be males of Pseudophilomedes
foveolatus Miiller, 1894, shows that they held a misconception of the

4 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

morphology of Pseudophilomedes. The fact that they state (1896,
p. 660) that in their opinion Pseudophilomedes is not sufficiently
different from Philomedes to warrant the establishment of a new
genus and, then, on page 670 establish the new genus Paramekodon
for precisely the same reason Miiller established Pseudophilomedes
must be attributed to confusion resulting from the unexpected
publication of Miiller’s work.

Brady and Norman (1896, p. 665) described a new species, Strepto-
leberis favosa Brady and Norman, 1896, collected at depths of 836 to
2333 meters off the west coast of Morocco. From 2 dried specimens
they were able to examine only fragments of appendages. Concerning
the 2nd antenna, Brady and Norman (loc. cit.) commented: ‘‘The
antennae resemble most closely in character the same organs in
Paramekodon, having no appendicular branch but two setae in its
place, the basal joint somewhat smaller than usual, the length ex-
ceeding the breadth, first joint of the swimming branch rather slender.”’
Fragments of the furca were illustrated by Brady and Norman (ibid,
pl. 62, figs. 20, 21). The distribution of strong and slender claws on
the lamella is similar to that of the genus Pseudophilomedes and also
to some species in the genus Euphilomedes Poulsen, 1962. Claw no. 1
of the furea has a large tooth near the middle of the convex margin
and bears short spines between the tooth and the tip of the claw.
The claw is quite similar to claw no. 1 on species of Pseudophilomedes.
Streptoleberis favosa is probably closely related to Pseudophilomedes
and may belong in that genus. Streptoleberis crenulata (Brady, 1890),
the type species of the genus, is known only from its shell and belongs
in the category ‘‘genera dubia et species dubia’? (Miiller, 1912, p. 15;
Poulsen, 1965, p. 44). Streptoleberis rectirostris Brady and Norman,
1896, the third species that has been referred to Streptoleberis, is also
known only from the shell and Miiller (1912, pl. 51) correctly consid-
ered it as a species dubia.

At the present state of our knowledge there seems little reason for
considering the present subfamily Philomedinae to be more closely
related to the Cypridininae than to other subfamilies comprising the
Cypridinacea. Therefore, the subfamily Philomedinae is raised to
familial rank. Some distinguishing characters between females of
the Philomedidae and Pseudophilomedidae are as follows:

Philomedidae Pseudophilomedidae

Shell, no. of medial bristles behind rostrum 6+ 4-5
1st ant. 2nd joint, no. of bristles 2-3 1
2nd ant. endop., no. of joints 1-2 1
2nd ant. endop., no. of bristles 6+ 2-4
2nd ant. exop. 9th joint, no. of bristles 4-7 2-3
Mandible, basale, no. of medial plus ventral bristles 10+ 4
Mandible, endop. 2nd joint, no. of dorsal bristles 8+ 4

NO. 3580 OSTRACOD FAMILIES—KORNICKER 5

(Continued) Philomedidae Pseudophilomedidae
Maxilla, no. of endites 3 2
Maxilla, end joint with process (p) or only spines (s) s p
5th limb, no. of endites 3 1?
5th limb, 2nd joint with extremely long fanglike tooth
(f) of with shorter tooth (s) 8 f
6th limb, no. of endites 4 3-4
6th limb, end joint, no. of bristles 9-45 70
7th limb, terminal with comb opposed by 2 pegs (p) or
2 opposing combs (c) p c

Skogsberg (1920), Poulsen (1962, 1965), and Hartmann (1964) are
correct in considering the families Rutidermatidae Brady and Nor-
man, 1896, Cylindroleberidae Miller, 1906 (= Asteropidae Brady,
1874), and Sarsiellidae Brady and Norman, 1896, to have coordinate
rank. In view of the anatomy of the male Rutiderma, recognized for
the first time by Poulsen (1965), referral of the genus Rutiderma to
the Sarsiellidae as done by Sylvester-Bradley (1961) seems no longer
tenable. Thus, the families contained in the superfamily Cypridi-
nacea are: Cypridinidae Baird, 1850; Philomedidae Miiller, 1912;
Rutidermatidae Brady and Norman, 1896; Cylindroleberidae Miller,
1906; Sarsiellidae Brady and Norman, 1896; Pseudophilomedidae,
new family.

Key to Families of Cypridinacea

1. Well-developed gills * along posterior of body, protopodite of maxilla elongate

with row of ventral bristles . . . . . . . . Cylindroleberidae
Without both well-developed gills and Bloneate protaporite on maxilla. . . 2

2. Mandible of female with large pincers distally. .... . Rutidermatidae
Mandible without large pincers . ......-+-++s+e+-2- oe ee eee a

3. Fifth limb of female with large teeth . .......+++-.+++-. er 4
Fifth limb of female without large teeth . ......+-+-.--. Sarsiellidae

4. Maxilla with only 2stout endites ...... .. - Pseudophilomedidae
Maxilla with 8endites .... sea pees <\Scypayaeeee ne

5. Lamellar prolongation of selvage iy fuinge of hat . . . . Philomedidae
Lamellar prolongation of selvage without fringe of hair. . . . Cy pridinidae

* Gills are common to all Cylindroleberidae but are rare in the Cypridinidae.

Pseudophilomedidae, new family

Philomedinae Miller, 1912, p. 24 (part).

Typr GENuS.—Pseudophilomedes Miller, 1894.

Diagnosis (female; male unknown).—Shell: Oval in lateral view
with rostrum and posterior process. Antennal sinus varies from
shallow curvature to deep incisure. Punctate surface smooth or
with longitudinal ridges or medial sulcus. Inner lamella with 4 or

6 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

5 bristles in row behind rostrum and 1-6 bristles dorsal to posterior
process. Lamellar prolongation of selvage with fringe of hairs
along anterior and neutral margins of shell.

First antenna: First joint without bristles; 2nd joint with 1 dorsal
bristle; 3rd jomt with 1 ventral and 1 dorsal bristle; 4th joint with
1 distodorsal and 2 distoventral bristles; 5th joint with 1 distoventral
bristle having short terminal filaments; 6th joint reduced, with a
short distomedial bristle; 7th joint reduced, with 1 short a-bristle
with spines, 1 medium b-bristle, and 1 long c-bristle; 8th joint reduced,
with 2 long bare d- and e-bristles and 2 long f- and g-bristles. Sensory
bristle on 5th joint, b- and c-bristles of 5th joint, and f- and g-bristles
of 8th joint with forked tips and few ringed filaments.

Second antenna: Endopodite single jointed with 1 long hirsute
bristle and 1-3 short bare bristles. Exopodite with 9 joints: 2nd to
Sth joints each with 1 long bristle; 9th joint with 2-3 bristles.

Mandible: Coxale endite bifurcate at tip. Basale with 4 ventral
and 3 dorsal bristles. Exopodite elongate with 2 terminal bristles.
Endopodite: 1st joint with 14 bristles; 2nd joint with 3+ dorsal
bristles, and 2-3 ventral bristles; end joint with 1 long claw, 2 shorter
claws, and about 3 bristles.

Maxilla: Precoxale endite with 3 stout pectinate spines and 2-3
bristles; coxale endite with 2 stout pectinate spines and 2 bristles.
Endopodite with slender to stout terminal process.

Fifth limb: Second joint prolonged distally forming long fanglike
tooth followed along the inner margin by 3 teeth. Inner margin of
protopodite with single endite with 1-2 bristles. Epipodial appendage
with numerous bristles.

Sixth limb: End joint with 7-9 bristles; endites 3-4 in number, not
more than 1 bristle in place of epipodial appendage.

Seventh limb: Terminus with opposing combs. Number of bristles
in distal group 4-6, in lateral group 2-8; total number of bristles 6-14.

Frontal organ: Elongate with or without(?) rings proximally.

Eyes: Large medial eye and small lateral eyes.

Furea: Third claw more slender than 4th. Total number of claws
6-10; all well-developed claws separated from lamella by suture.

DistrisutTion.—Atlantic Ocean off Morocco, on Great Bahama
Bank, B.W.I., on Continental Shelf off North Carolina and Georgia,
U.S.A., Gulf of Naples (Mediterranean Sea).

Ecotoay.— The family is benthonic. It has been reported from
water with temperatures ranging from about 3° to 29° C., water depths
of 6 to 1435 meters, and salinities from about 35 to 38 parts per thou-
sand. Specimens have been collected from shelly sand, sandy mud,
and calcareous algae. Some environmental factors concerning each
species are presented in table 1.

OSTRACOD FAMILIES—KORNICKER

NO. 3580

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8 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Pseudophilomedes

Pseudophilomedes G. W. Miiller, 1894, p. 211.

Paramekedon Brady and Norman, 1896, p. 670.
Type spEciEs.—Pseudophilomedes foveolatus Muller, 1894.
GENDER.— Masculine.
Diagnosis.—Same as for family.

Key to Species of Pseudophilomedes

(species in brackets not described in present paper)

1. Shell with longitudinal ridges ........ [P. angulatus Miiller, 1894]
Shell: without: longitudinal ridges ..., < ss ses. be. hee ees ae ee 2

2. Endopodite of 2nd antenna with 2 short bristles
[P. inflatus (Brady and Norman, 1896)]

Endopodite of 2nd antenna with more than 2 short bristles. ...... 3
3. Endopodite of 2nd antenna with 3 short bristles ...... P. foveolatus
Endopodite of 2nd antenna with 4 short bristles. . ..... P. ferulanus

Pseudophilomedes ferulanus

Fiaures 1-6; Puate 1

Pseudophilomedes ferulana Kornicker, 1959, p. 235, figs. 46, 1A-B, 2A-B; 56,
A-D.—Darby, 1965, pp. 64-70, fig. 10, pls. 11, 12.

Hototypr.—USNM 113287; male; complete dry specimen. Spec-
imens from Atlantic Shelf, 8 specimens: USNM 113020-113022.

Marteriau.—The number of specimens, locations of sampling
stations, and some environmental factors concerning specimens
collected from the Atlantic Shelf are presented in table 2. All
specimens sufficiently developed for determining sex were females.
Five specimens were dissected for this study.

SUPPLEMENTAL DESCRIPTION OF FEMALE (male unknown).—Shell
(fig. 3): Oval, highest anterior to center, widest posterior to center;
posterior process slightly below middle of valve; rostrum subtrian-
gular with nearly horizontal lower margin; antennal sinus shallow;
surface punctate; each valve with arcuate sulcus extending from
dorsal margin to near middle of valve; left valve overlaps right valve
along anterodorsal and posterodorsal margins; hinge simple, extending
from highest point on valve to posterodorsal corner; about 15 muscle
scars in front of and below middle of each valve (fig. 3c), additional
muscles are attached near anterodorsal corner of valve (fig. 3h).
Inner lamella broadest behind rostrum and in front of posterior
projection; radial pore canals numerous (fig. 3f); selvage with lamellar
prolongation and fringing hairs (fig. 3g); inner lamella behind rostrum
with row of 4 bristles with short marginal hairs (fig. 3d) ; inner lamella
in front of posterior process with row of 6 hirsute setae (fig. 3e) ; inner

]

—— SS

NO. 3580 OSTRACOD FAMILIES—KORNICKER 9

lamella below antennal sinus with 1 short bristle; outer surface of
each valve with scattered tapered hairs.

Darby (1965 p. 64) states concerning the shell, “About six nodes
along the contact margin of both valves slightly dorsal to caudal
process.” These nodes each contain a hirsute setae, and the nodes
are well-defined pores rather than raised structures (pl. 17).

Darby (1965, p. 64) reports 4 or 5 bristles on the inner lamella
behind rostrum, indicating variability in the number of bristles in
that area; only 4 bristles were observed in the examined specimens,

Shell dimensions (in mm) are as follows (see table 2 for station
data) :

Specimen greatest greatest greatest
station no. length height width remarks
BST 30 1 1. 66 1. 16 1.00 eggs in brood pouch
ee 2 1. 30 0. 81 late instar
cs 3 1. 54 1. 00 eggs in body
ss 4 152, 1. 00 eggs in body
a 5 1. 54 0. 96 no eggs
GOS 1857 1 eae ip 1S eggs in brood pouch

First antenna (figs. 4a—d): First joint with spines along ventral mar-
gin; 2nd joint with dorsal bristle and marginal spines; 3rd joint with
1 ventral and 1 dorsal terminal bristle and marginal spines; 4th
joint with 1 subterminal dorsal and 2 terminal ventral bristles; 5th
joint with 1 long ventral, terminal sensory bristle with forked tip
and 2 proximal, ringed filaments; 6th joint minute and fused to 5th,
with 1 short spinous medial bristle; 7th joint reduced and fused to
8th joint, with 1 short spinous a-bristle, 1 medium b-bristle with
forked tip and 1 distal filament, and 1 long stout c-bristle with forked
tip and 2 proximal filaments; 8th joint reduced, with 2 long bare
d- and e-bristles, 1 long f-bristle with forked tip and 3 distal and 1-2
proximal filaments, and 1 long g-bristle with forked tip and 1 distal
and 2 proximal filaments; Ist to 4th joints with rows of slender spines
on medial surface.

Second antenna (figs. 4e, f): Endopodite (fig. 4/) single jointed, with
long hirsute bristle and 2 short bare bristles. Exopodite (fig. 4e) with
9 joints: 1st joint slightly longer than total length of following joints;
2nd to 9th joints trapezoidal, each joint smaller than preceding joints,
all without basal spines, 2nd to 8th joints with long bristles, bristle
on 2nd joint with spines and natatory hairs, bristles on 3rd to 8th
joints with sparse natatory hairs and stout marginal spines near
middle; 9th joint with medium spinous bristle and 2 long bristles
with natatory hairs; distal margins of 2nd to 8th joints with comb
of slender spines. Medial surface and anterior margin of protopo-
dite with clusters of slender spines.

226-965—67—2

10 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Mandible (figs. 49-7): Coxale endite (fig. 4h) bifurcate, 1 prong
annulate, other nonannulate, both with spines, more spines on annu-
late prong. Basale with 7 bristles, 4 ventral, 3 dorsal. Exopodite
with 2 bristles, outer bristle about half length of inner bristle. Endop-

mandible
frontal organ

end antenna

maxilla

5th limb
furca

- adductor muscles

6th limb
oS .

epipodial appendage

(on 5th limb)

-7thlimb

: SSR eRe

Ficure 1,—Pseudophilomedes ferulanus: ventral view, removed from shell; all bristles,
muscles, and appendages not shown; appendages and joint positions generalized.

(Spec. 1, Gosnold Sta. 1857.)

at i A i
sien a
Seen ll

No. 3580 OSTRACOD FAMILIES—KORNICKER 11

odite: 1st joint with 1 short and 2 long ventral bristles distally;
ond joint with 1 dorsal bristle proximal to middle, 3 dorsal bristles
near middle, and 3 short ventral bristles distally; end joint (fig. 42)
with 3 bristles and 3 claws, 1 of latter very long, 1 about a third to
half length of long claw, 1 on dorsal edge very short. Medial surface
of endopodite and basale with clusters of fine hairs.

Maxilla (figs. 1, 5a-d): Precoxale endite with 3 stout pectinate

ant,
——_$_—_$_—_—_———_-

S
>
=>
=<
re,
sd
©

J
JZ
Lom A
YJ
Coe mr
*
{]

Me i
maxilla
,
= K ae
aE a cy is
y= TILL sy o
Vowi»ny Rs gee

5th limb

Ficure 2.—Pseudophilomedes ferulanus: lateral view, left maxilla, 5th limb in place;
all bristles not illustrated. (Spec. 3, Beaufort Shelf Transect, Sta. 30.)

spines and 2 spinous bristles. Coxale with stout plumose bristle,
hirsute epipodial fringe and endite with 2 stout pectinate spines and
2 spinous bristles. Distal edge of basale with 1 spinous medial
bristle and 2 on inner margin adjacent to low node with 2 bristles
at base of coxale endite, shorter of latter bristles bare, other with
several marginal spines. Exopodite consisting of 3 lateral spinous
bristles distally on basale (figs. 1, 5a). Endopodite: 1st joint hirsute
with 2 spinous bristles, 1 near middle and 1 terminally; 2nd joint

12 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

with 1 short and 2 long terminal spinous bristles and a long stout
process with annulations and a rough surface of small pustules distally.

Fifth limb (figs. 5e-7): First joint with 2 large teeth: proximal
tooth trident with 3 bristles (fig. 5f); distal tooth bifid (fig. 5h).
Second joint prolonged distally forming long fanglike tooth followed
along the inner margin by a single tooth and 2 trident teeth (fig. 5e);
posterior surface with 2 stout spinous bristles and a minute bristle
(fig. 52) ; inner edge of joint with spinous bristle (fig. 51). (It was not
possible to discern with certainty that this bristle was not in fact on a
proximal margin of the distal tooth of the first joint.) Third joint
with 2 short hirsute bristles (fig. 57). Fourth joint hirsute with 1
short and 2 long terminal bristles and 1 short subterminal bristle
(figs. 5g,i). Epipodial appendage with 42-46 bristles. Single endite
with 2 short bristles.

Sixth limb (fig. 6a): First endite with 1 to 2 bristles; 2nd endite
with 2 bristles; 3d and 4th endites each with 5 to 6 bristles; end joint
with 9 to 10 bristles (end joint of late instar with only 7 bristles) ; single
bare bristle in place of epipodial appendage. Surface with clusters
of hairs; end joint fringed with hair.

Seventh limb (figs. 6)-d): Symmetrical terminus with opposing
combs of 5 or 6 teeth (all teeth not shown in figs. 6 b,c). Adult with
6 terminal and 6 to 8 lateral bristles (instars with fewer bristles), each
bristle with 1 to 5 bells (fig. 6d) and short marginal spines distally.
Distribution of bristles on seventh limbs of 4 specimens from station
BST 30 are as follows:

limb A limb B*
specimen no. 1

terminal bristles 6 6

lateral bristles 8 a
specimen no. 2

terminal bristles 4 4

lateral bristles 4 6
specimen no. 3

terminal bristles 6 lost

lateral bristles 6 lost
specimen no. 4

terminal bristles 6 6

lateral-bristles 8 7

* Limbs A and B are opposing 7th limbs; they have not been classified as either left or right.

Furea (figs. 6e, f-h): Each lamella with 8 to 10 claws;3rd claw thinner
than 4th but about the same length; each claw separated from lamella
by suture with exception of 9th and 10th claws which, when present,
appear as nodes. Claw 1 with stout spines in row proximally on
lateral margin, slender spines distally on lateral and medial margins,
1 large spine near middle of medial margin, and groups of slender

No. 3580 OSTRACOD FAMILIES—KORNICKER 13

spines distally on anterior surface; claws 2-8 with spines along medial
and lateral margins; lamella, in area of distal and proximal claws,
with hairs.

Eyes: Median eye (fig. 67) large, pigmented. Lateral eyes (fig. 69)
small, black in preserved specimens.

Frontal organ (figs. 67, &): Elongate, proximal part divided into
about 8 short segments, distal part (fig. 6k) tapered, with short spines
near terminus.

Eggs (fig. 67): As many as 8 eggs observed in thoracic region; 2 to
6 eggs in brood pouch.

Parasites: Two larval copepods of genus Sphaeronella in brood
pouch of 1 specimen having eggs inside body (specimen no. 4, Beaufort
Shelf Transect 30).

Pseudophilomedes foveolatus

Figures 7, 8
Pseudophilomedes foveolata Miller, 1894, pp. 211-212, pl. 3, figs. 34, 35, 45-49,
51, 53, 54; pl. 4, figs. 127. [Not Philomedes foveolata Brady and Norman,
1896, pp. 659-661, pl. 56, figs. 4, 5.]

SyntypEs.—Zoological Museum of Berlin, Division of Crustacea,
catalog no. 9154.

Marerrau.—A vial containing 1 whole specimen and a left and
right valve in alcohol was received from the Zoological Museum,
Berlin. The accompanying label contained the following information:
Kat. Nr, 9154; Species Pseudophilomedes foveolata G. W. Miller,
1894; Fudort Neapel. An asterisk appears in the upper left-hand
corner of the label.

The small size of the specimens leads me to believe that they are
late instars of females. I have designated the whole specimen as
specimen number 1 in this paper and on the prepared slide.

DESCRIPTION OF LATE INSTAR OF FEMALE (male unknown).—Shell
(figs. 7a-d): Lateral outline oval, highest anterior to center; posterior
process slightly below middle of valve; rostrum subtriangular with
nearly horizontal lower margin; antennal sinus shallow; surface punc-
tate; ovoid muscle scars in front of and below middle of each valve.
Inner lamella broadest behind rostrum and in front of posterior pro-
jection; selvage with lamellar prolongation and fringing hairs unbroken
around antennal sinus; inner lamella behind rostrum with row of 4
bristles with short marginal hairs (fig. 76); inner lamella in front of
posterior process with 1 (or 2 closely spaced?) hirsute seta (fig. 7c);
inner lamella below antennal sinus with 1 short bristle; outer surface
of each valve with scattered surface hairs.

Miiller (1894, pl. 3, fig. 7) illustrates 4 bristles on the inner lamella
behind the rostrum but does not show them to be hirsute. He evi-

14 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

dently also overlooked the faint seta on the inner lamella in front of
the posterior process and the small bristle below the antennal sinus.
Shell dimensions (in mm) are as follows:

specimen no. greatest length greatest height remarks
1 0. 73 0. 46 whole specimen
2 0. 64 0. 43 separated left valve
3 0. 73 0. 48 separated right valve

Miiller (1894, p. 212) gives a length of 0.93 mm for this species.
The smaller size of specimens available for the present study indicates
that they are instars.

First antenna (figs. 7e,f): Second joint with 1 spinous dorsal bristle
and sparse clusters of spines on medial surface; 3rd and 4th joints
each with 1 ventral and 1 dorsal spinous terminal bristle; 5th joint
with 1 long ventral, terminal sensory bristle with forked tip; 6th joint
minute, welded to 5th joint, and with 1 short medial bristle; 7th joint
reduced, welded to 8th joint, and with 1 short spinous a-bristle,
1 short b-bristle with forked tip and 1 distal filament, and 1 long
c-bristle with forked tip; 8th joint reduced, with 2 long bare d- and
e-bristles, 1 long f-bristle and 1 long g-bristle, both with forked tips.

The illustration of the Ist antenna illustrated by Miler (1894,
pl. 8, fig. 48) bears 2 ventral bristles terminally on the 4th joint (con-
sidered 5th joint by Miiller). The presence of only 1 bristle in that
position on the specimen I examined may be due to its not being
adult. A few filaments were observed on long bristles of joints 5-8,
but it was not possible to determine the exact number and precisely
on which bristles they belonged.

Second antenna (figs. 7g-i): Endopodite (fig. 7h) single jointed, with
1 long hirsute bristle and 1 short bare bristle. Exopodite with 9
joints: 1st joint slightly longer than total length of following joints;
2nd to 9th joints trapezoidal, each joint smaller than preceding joints,
all without basal spines; 2nd to 8th joints with long bristles with
stout spines along outer margin, becoming shorter, more slender, and
numerous at distal end; 9th joint with 1 short and 1 long spinous
bristle; all bristles without natatory hairs; distal margins of 2nd to 8th
joints with comb of slender spines; tip of bristle on 2nd joint not
reaching 9th joint.

The 2nd antenna shown by Miiller (1894, pl. 3, fig. 35) is similar to
the specimen described in this paper. Miller apparently over-
looked the row of short spines along the distal margin of joints 2-8
of the exopodite.

Mandible (figs. 77, k): Coxale endite (fig. 77) bifurcate with distal
spines. Basale with 7 bristles, 4 ventral, 3 dorsal. Exopodite with
2 bristles, outer bristle shorter than inner bristle and with wreaths
of long hairs. Endopodite: 1st joint with 1 short and 2 long ventral

NO. 3580 OSTRACOD FAMILIES—KORNICKER 15

bristles terminal; 2nd joint with 1 short dorsal bristle proximal to
middle, 3 dorsal bristles near middle, 1 short ventral bristle distally,
and 2 short terminal bristles near ventral margin; end joint (fig. 7/)
with 3 bristles and 2 claws, 1 of latter very long, other about 3rd
length of long claw. Terminal end of list endopodite joint with
spines in row.

The mandible of the species illustrated by Miiller (1894, pl. 3,
fig. 51) has only 2 ventral bristles distally on the 1st endopodite joint,
only 2 dorsal bristles in the distal group on the 2nd endopodite joint,
and does not show 2 small bristles terminally on the 2nd endopodite
joint or a small centrally located terminal bristle on the end joint.
These were probably overlooked by Miiller.

Maxilla (fig. 71 m): Precoxale endite with 3 stout pectinate spines
and 2 bristles. Coxale endite with 2 pectinate spines and 2 bristles.
Distally of coxale endite a short lobe with 3 bristles. Basale with 1
medial bristle and 2 bristles on inner margin. Exopodite consisting
of 3 lateral bristles located distally on basale. Endopodite: Ist joint
with 1 subterminal and 1 terminal bristle; 2nd joint with 1 short and
2 long bristles and a slender process with annulations distally.

Miiller (1894, pl. 3, fig. 49) illustrates the slender process on the end
joint of the endopodite as a bristle. Although the process resembles
a stout bristle, it is ringed only distally and is less tapered than a
bristle would be. Miiller’s illustration does not have the 3 bristles I
interpret as the exopodite. The specimen I examined does not have
a bristle on the distal margin of the coxale, possibly it broke off during
the dissection. Unfortunately the left maxilla was lost so that I had
the opportunity to examine only the right maxilla.

Fifth limb (figs. 8a-e): First joint with 2 large teeth; proximal tooth
trident with 2 or 3 bristles, forward tooth spinous on posterior margin ;
distal tooth bifid with small secondary tooth above base of lead tooth,
forward tooth with spines on leading and posterior margins, spines
larger on leading margin. Second joint prolonged distally forming
long fanglike tooth followed along the inner margin by a single tooth
and 2 trident teeth; posterior surface with 1 long and 1 short bristle.
Third joint with 2 spinous bristles near base of 4th joint. Fourth
joint hirsute with 4 or 5 bristles. Inner margin of protopodite with
single endite with 1 or 2 minute bristles. “at

The taxonomically important inner margin of the 2nd joint with its
3 teeth is not shown in the view of this appendage illustrated by
Miiller (1894, pl. 3, figs. 43, 46). The fanglike tooth distally on the
2nd joint is much longer on the specimen I examined than would be
suspected from examination of Miiller’s illustrations. Possibly short-
ness of that tooth in Miiller’s illustration is the result of foreshortening,
which results when an appendage is mounted obliquely on the slide.

16 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

The 2 teeth of the Ist joint of the specimen illustrated by Miiller and
the one I examined are quite similar. Miller apparently interprets
the 2 bristles, which I consider to be on the posterior surface of the
2nd joint, to be on the inner lobe of the 3rd joint. This is quite
possible, but difficult to verify, because the suture between 2nd and
3rd joints is not visible on specimens I examined. The 4th joint
illustrated by Miiller (figs. 43, 46) does not have the short terminal
bristle present on the specimen I examined. This may have been
overlooked by Miiller, or possibly the number of bristles on the 4th
joint varies slightly within the species.

Sixth limb (fig. 8f): First endite with 2 bristles; 2nd endite with 5
bristles; 3rd endite with 4 bristles; end joint with 5 (?) bristles; single
bristle in place of epipodial appendage.

The appendage illustrated by Miiller (1894, pl. 3, fig. 54) is quite
similar to the one I examined with the exception of bearing 8 bristles
on the end joint. Since the end joint I examined belongs to an im-
mature specimen, it is possible that it has fewer bristles; because of the
position of the 6th limb on the slide, I was not able to see the end joint
clearly, so it is likely that more than 5 bristles are present.

Furca (figs. 8g-i): Each lamella with 6 claws; claws 1, 2, 4 being
stout claws, and claws 3, 5, 6 slender claws; claw 3 shorter than claw 4 :
each claw separated from lamella by suture; claw 1 with stout spines
in row proximally on lateral margin, slender spines distally on lateral
and medial margins, and 1 large spine near middle of medial margin.
Remaining claws with spines along medial and lateral margins; lamella,
in area of distal and proximal claws, with hairs.

Miiller (1894, pl. 3, figs. 34, 53) figures a furca with the 3rd claw
almost equal in length to the 4th claw. This suggests that the length
of the 3rd claw may be somewhat variable within the species. Miil-
ler’s figures do not show peripheral spines on claws 3, 5, 6. It seems
probable that these were overlooked by Miiller. In this regard, it is
interesting that Brady and Norman (1896, p. 672) state that the larger
furcal claws of Pseudophilomedes inflatus are ciliated on the edge and
in the figure of this appendage (1896, pl. 59, fig. 10) show claws 3 and
8-10 without marginal spines. In view of Miiller’s omission of mar-
ginal spines on slender claws, it seems likely that they also were over-
looked by Brady and Norman. It would be most unusual and seems
quite unlikely for spines to be absent from the slender claws, especially
claw 3.

Family Philomedidae Miiller

Philomedinae Miller, 1912, p. 24 (part).

TypE Genus.—Philomedes Liljeborg, 1853. Gender: Masculine.
D1aGnosis.—Shell: Shell variable in outline, but generally elongate

NO. 3580 OSTRACOD FAMILIES—KORNICKER hz

with convex dorsal and ventral margins; surface smooth or with
complex ornamentation; anterior with rostrum and shallow-to-deep
antennal sinus; posterior rounded or with process; lamellar prolonga-
tion of selvage with fringe of hairs.

First antenna: First joint without bristles; 2nd joint with 1 disto-
lateral bristle, 1 distodorsal bristle, and usually 1 distoventral bristle;
3rd joint with 1 ventral and 1-4 dorsal bristles; 4th joint with 1-4
distoventral and 1-2 distodorsal bristles; 5th joint of male consider-
ably reduced but always represented ventrally by a sensory bristle
broadening proximally and with numerous long filaments; 5th joint
of female not reduced, and with a stout distoventral bristle with short
filaments; 6th joint of male not reduced, and with a slender disto-
medial bristle; 6th joint of female considerably reduced but always
with a slender distomedial bristle; 7th joint reduced, and with 1 short
distodorsal bristle, 1 long distomedial bristle with short filaments,
and 1 long distoventral bristle with filaments (a-, b-, and c-bristles
respectively); Sth joint reduced and with 2 long bare distolateral
bristles (d- and e-bristles), and 2 long distomedial bristles with
filaments (f- and g-bristles); c- and f-bristles on male extremely long,
about length of shell.

Second antenna: Endopodite of male with 3 joints, each joint
with 2 or more bristles; endopodite of female with 1-3 joints having
total of 6 or more bristles. Exopodite: 3rd joint of male at least
twice length of 2nd joint; in female joints decreasing in length distally;
joints usually with basal spines; 2nd to Sth joints each with 1 bristle;
9th joint usually with 6-7 bristles.

Mandible: Coxale in female large, bifurcate at tip, in male reduced
or absent. Basale with 5-9 ventral bristles, 1-4 dorsal bristles, and
5-6 medial bristles. Exopodite elongate with 2 terminal bristles.
Endopodite: 1st joint with 4~7 ventral bristles; 2nd joint with 5 or
more dorsal bristles and ventrally at least 1, but usually about 6
bristles. End joint with 2-3 claws and 3-4 bristles.

Maxilla: Endites 3 in number; exopodite elongate with 3 bristles;
maxilla on female usually larger than on male.

Fifth limb: Exopodite: 1st joint of female with 2 or more strong
teeth; 2nd joint of female forming large squarish sclerotized process
having large inward projecting tooth distally and an inner margin
either smooth or with small nodes or 2 large teeth; Ist and 2nd joints
of male without stout teeth; 3rd joint with 3 bristles on inner lobe
and 2 on outer lobe; 4th joint elongate with 5-6 bristles. Endites 3
in number with a varying number of bristles. Epipodial appendage
with numerous bristles (more than 30).

Sixth limb: Endites 4 in number; end joint with 9-45 bristles; 2-4
bristles in place of epipodial appendage.

226-965—67——3

18 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Seventh limb: Terminus with comb opposed by 2 pegs; cleaning
bristles 6-35 in number.

Frontal organ: Elongate with or without rings.

Eyes: Large medial eye. Lateral eyes in female large with numer-
ous ommatidia, in male absent or small without numerous ommatidia.

Furca: Considerable variation in distribution of primary and sec-
ondary claws. Total number of claws 6-17; all or some claws sepa-
rated from lamella by sutures.

The following genera are referred to the Philomedidae: Philomedes
Lilljeborg, 1853: ?Pleoschisma Brady, 1880: ? Tetragonodon Brady and
Norman, 1896: Scleroconcha Skogsberg, 1920; Ewphilomedes Poulsen,
1962; Paraphilomedes Poulsen, 1962.

Euphilomedes Poulsen

TYPE spEcIES.—Euphilomedes nodosus Poulsen, 1962, by subsequent
designation, Kornicker (in press). Gender: masculine.

Euphilomedes asper, new combination
Fiaures 9, 10, lla-d, 12

Philomedes aspera Miller, 1894, pp. 210-211, pl. 3, figs. 3, 17, 21; pl. 8, fig. 1.
Philomedes foveolata.—Brady and Norman, 1896, pp. 659-661, pl. 56, figs. 4, 5.
[Not Pseudophilomedes foveolata Miiller, 1894.]

SyntypPes.—Zoological Museum of Berlin, Division of Crustacea,
catalog no. 9152.

Mareriau.—1. A glass slide in the collection of the British Museum
(Natural History) labeled ‘“Philomedes foveolata Bra & Noro” at
one end of the slide and at the other end ‘‘Naples 1887”; a 3rd label
bears the number 1900-3-6-452. The slide contains a dissected
ostracod with parts under 6 cover slips. This slide probably contains
the specimen upon which Brady and Norman (1896) based their
description. Because of the poor condition of the slide, another
specimen (see paragraph 2, below) was used for the description and
most illustrations used in this paper. However, the appendages and
carapace on Brady and Norman’s slide were carefully compared
wherever possible with those from the specimen I dissected. Both
specimens are almost identical, but where minor differences were
observed they are discussed when the appendage is described.

2. A vial in the collection of the British Museum (Natural History)
containing a smaller vial with 2 whole undissected specimens in
alcohol. The larger vial contains the label ‘Pseudophilomedes
foveolata, Naples 1887, Norman Coll., B. M. regd. no 1911, 11.8.
36962964.” The specimens in this vial are designated in this paper

NO. 3580 OSTRACOD FAMILIES—KORNICKER 19

as specimens A and B. Specimen A wis dissected and used in de-
scriptions and illustrations.

3. A jar in the collection of the Zoological Museum of Berlin
containing a small vial with 1 whole ostracod and 1 left and 1 right
valve. The jar contains the label “Type, Philomedes aspera G. W.
Miller, Kat. Nr 9152, Fundort Napoli.” The small vial contains the
label, “9152.” These specimens are syntypes; Miiller (1894) did not
designate a holotype for the species. The whole specimen is a
mature male. ‘The left and right valves are from different specimens;
their sex is unknown. I have designated the whole specimen as
number 1, the right valve as number 2, the left valve as number
3, and have illustrated each valve. All specimens were returned to
the Zoological Museum of Berlin in the same vial in which they were
received. ‘The whole specimen was not dissected, but as the furca
was projecting from between the valves, it was possible for it to be
described and illustrated in this paper.

SUPPLEMENTARY DESCRIPTION.—Shell (figs. 9a; 12a—f): Lateral
outline oval, elongate with greatest height near middle, prominent
rostrum and broad rostral incisure (figs. 9a, 12a-d); surface of valves
with numerous large oval pits (fig. 9b). Posterodorsal margin with
straight hinge; posterior hinge element of each valve consisting of
angular sclerotized process; medial hinge element straight; anterior
hinge element not prominent. Adductor muscle scars consisting of
about 16 or 17 individual scars below middle of valve (figs. 9e, 7;
12e). Inner lamella broadening at posteroventral process and behind
rostrum (figs. 96, c; 12e). Selvage with wide, corrugated, lamellar
prolongation having fringe of slender spines along outer margin.
Twelve to 14 bristles forming row on inner lamella behind rostrum
(figs. 9b, f; 12d); inner lamella below rostrum with small hair followed
by a wide space and then 10 hairs on left valve and 14 on right valve
(figs. 9d, h). Posteroventral process with small hairs on inner lamella
(figs. 9c, g; 12e); small hairs, singly or in pairs, forming row on list
on posteroventral and posterior part of inner lamella (figs. 9e, 9).
Radial pore canals numerous but faint. Hairs with taper distributed
on lateral surface of rostrum, some forming row near ventral and
posterior margins of valves; additional hairs sparsely distributed on
valve surface.

Muscle scars were somewhat obscured on all specimens. Four
muscle scars located in the anterior part of the central muscle field
of Miiller’s syntypes were not observed on the specimens of Brady and
Norman.

20 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Shell dimensions (in mm) are as follows:

collector Specimen greatest length greatest height remarks

Brady and Norman A 1. 34 0. 85 mature male
B 1. 35 0. 81 Ch ance

Miller 1 1. 30 0. 80 . i:
2 1. 28 0. 78 zs .
3 0. 94 0. 60 instar

Brady and Norman (1896, p. 659) give measurements of length as
1.6 mm and height as 1 mm for specimen BM 1900-3-6-452; because
of the poor condition of the mounted specimen, I was unable to
measure it.

Miiller (1894, p. 211) gives the length of the male as 1.3 mm.

First antenna (fig. 97): First joint with clusters of spines on medial
surface; 2nd joint with clusters of hairs on medial surface, short spines
projecting from ventral and dorsal margins, and distally with a dorsal,
ventral, and lateral bristle; 3rd joint with clusters of hairs on medial
surface and 1 ventral and 2 dorsal bristles; 4th joint with clusters of
hairs on medial surface and 2 dorsal and 4 ventral bristles distally;
5th joint inferred to be inserted between 4th and 6th joints bearing sen-
sory bristle with broad base and numerous filaments; 6th joint with
medial bristle distally; end joints with 2 long stout c- and f-bristles,
1 short spinous dorsal a-bristle, and 2 long b- and g-bristles with
filaments and 2 bare d- and e-bristles.

The above description of the 1st antenna differs from Brady and
Norman’s (1896, p. 660) in having 2 rather than 1 bristle on the dorsal
margin of the 3rd joint and in having a total of 7 rather than 6 bristles
on the end joint. The 1st antenna on slide no. 1900-3-6-452 con-
taining a specimen dissected by Brady and Norman confirms the
presence of 2 dorsal bristles on the 3rd joint and a total of 7 bristles on
the end joint.

Second antenna (figs. 9k, 1; 10a): Endopodite (fig. 9k) with 3 joints:
Ist joint with 5 proximal bristles and 1 distal bristle; 2nd joint elongate
with 2 bristles near middle; 3rd joint prehensile with a long slender
bristle about one-third distance from proximal end of joint and 2
short bristles and about 5 crescentic ridges at distal end. Exopodite
with 9 joints; 1st joint elongate with small medial spine distally; 2nd
joint about one-third length of 1st with spined bristle reaching 6th
joint; 3rd joint about twice length of 2nd; joints 4—9 trapezoidal, each
joint smaller than preceding joint; joints 3-8 with slender basal spines
and row of short hairs distally on lateral margin; bristles on joints 3-8
with natatory hairs; end joint (fig. 10a) with 4 long distal bristles with
natatory hairs and 2 smaller mediodorsal bristles with sparse marginal
hairs.

Brady and Norman (1896, p. 660) report only 3 bristles on the 1st
joint of the endopodite of the 2nd antenna ; however, 6 bristles are

No. 3580 OSTRACOD FAMILIES—KORNICKER ot

present on this joint on slide no. 1900-3-6-452 containing a specimen
dissected by Brady and N orman. The 1st endopodite joint of
Philomedidae usually contains more than 3 bristles, so it is probable
that Brady and Norman were unable to make an accurate count of
bristles on that joint.

Mandible (figs. 106, c): No coxale endite. Basale ventral margin
with 7 subequal bristles with wreaths of long hairs, some also with
short marginal spines distally ; dorsal margin with 1 bristle near middle
and 2 distally; subventral medial surface with 4 small proximal
bristles and 1 bristle proximal to middle. Exopodite reaching about
middle of 1st endopodite joint, with 2 subequal terminal bristles.
Endopodite 1st joint with 2 short and 3 long distoventral bristles;
2nd joint with 3 dorsal bristles proximally, 6 dorsal bristles near
middle, 2 ventral bristles subdistally and 2 long and 2 short ventral
bristles distally. End joint (fig. 106) with 2 large subequal claws,
1 short dorsal claw, and 4 bristles. Medial surface of basale and 1st
and 2nd endopodite joints with clusters of hairs.

Maxilla (fig. 10d): Three endites: 1st endite with about 8 bristles;
2nd endite with about 7 bristles; 3rd endite with 1 proximal bristle
and numerous distal bristles. Coxale with long plumose bristle.
Exopodite with 1 short and 2 long bristles.

Fifth limb (figs. 10e, g): Epipodial appendage with about 33 bristles.
Three endites with numerous bristles. Exopodite 1st and 2nd joints
each with a flat, transparent, bladelike tooth and several bristles;
3rd joint with 2 stout plumose bristles on outer lobe and about 3 bare
bristles on inner lobe; 4th and 5th joints with 6 bristles.

Sixth limb (fig. 10f): First endite with 3 bristles; 2nd endite with
1 proximal and 3 distal bristles; 3rd and 4th endites each with 1 proxi-
mal and 7 distal bristles; end joint with 14 bristles; 3 plumose bristles
in place of epipodial appendage. Lateral and medial surface of end
joint with clusters of hairs. Some bristles of end joint broken on
specimen examined.

The description of the 6th limb given by Brady and Norman (1896,
p. 660) differs from the above. The 1st endite seems to have been
pulled off on Brady and Norman’s dissected specimen (slide no. 1900-
3-6-452), but part of 1 bristle remains to show that the endite was
present. The missing Ist endite on the specimen evidently led Brady
and Norman to mistakenly consider the 2nd endite to be the Ist
endite, the 3rd endite to be the 4th, and the 4th to be part of the end
joint. Brady and Norman described the 1st lobe [their terminology]
to have 2 setae, and the 2nd lobe to have 1 lateral and 6 distal setae.
The dissected specimen (slide 1900-3-6-452) actually has 1 lateral
and 3 distal setae on the 1st lobe [their terminology] and 1 lateral and
7 distal setae on the 2nd lobe. The remainder of the description of

29 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Brady and Norman conforms to the above description and describes
accurately the appendage on slide 1900-3-6-452. However, Brady
and Norman do not mention 2 slender bristles on the end joint; this
brings the total number of bristles on this jomt to 14, nor do they
mention 3 bristles in place of the epipodial appendage; those bristles
are present on the specimen on slide 1900-3-6-452.

Some variability was observed in the location of 2 relatively short
and slender medial bristles on the margin of the end joint. These
bristles occupy positions nos. 5 and 8 (counting from the posterior end
of the joint) on the appendage on slide 1900-3-6—452; whereas, on
specimen A dissected by the writer, these bristles occupy positions 6
and 9 on the left appendage, and 7 and 9 on the right appendage.

Seventh limb (figs. 107, &): Four bristles of subequal length in
distal group, each with 5 bells; 4 shorter bristles in proximal group,
each with 3 bells (fig. 107); marginal hairs not observed on bristles.
Terminal comb with 9 or 10 teeth, some with basal spines and flanges;
2 subequal pegs opposing comb (fig. 10%).

Furea (figs. 107, 1, m): Each lamella with 10 or 11 claws: primary
claws 1, 2, and 4, remaining claws secondary (fig. 10/). Secondary
claw no. 3 slightly larger than claws 5 to 10 (figs. 107, m), claws 6 to
10 about equal in length. All claws separated from lamellae. Pri-
mary claws with lateral and medial row of pointed teeth; tooth length
varying only slightly in length on each claw. Secondary claws with
row of spines along each margin. Clusters of long hairs medially on
lamella between claws 1 to 8.

Specimen number 1 of Miiller’s syntypes contains 11 claws on the
left lamella and 10 on the right, all other specimens examined contain
10 on each lamella. Both Miiller (1894) and Brady and Norman
(1896) describe their specimens as having 10 claws on each lamella.

Copulatory limbs (fig. 11a): Elongate terminating in 3 lobes, each
lobe with 2 or more short bristles, 1 lobe with sclerotized tip.

Frontal organ (figs. 116, c): Elongate, 2-jointed with 2nd joint
tapering distally and having 2 small spines at tip.

Eyes: Medial eye large (fig. 116). Lateral eyes large with about
13 ommatophores, each weakly divided by a suture into 2 parts
(fig. 11d).

Discussion.—Differences between Brady and Norman’s Philo-
medes foveolatus (Miiller) and Pseudophilomedes foveolatus Miiller are
too large to be the result of sexual dimorphism. For example, each
caudal lamella of Brady and Norman’s specimens has 10 or 11 claws,
whereas P. foveolatus of Miiller has only 6; also, upper claws of Brady
and Norman’s specimens are subequal in length, whereas those on
Miiller’s P. foveolatus decrease uniformly in size proximally; in addi-
tion, the 3rd claw of the furca on Brady and Norman’s specimen is

<i el

NO. 3580 OSTRACOD FAMILIES—KORNICKER 23

about half the length of the 4th claw, but it is almost the same length
of the 4th claw on P. foveolatus. Also, Brady and Norman’s speci-
mens have 13 to 14 medial bristles on the rostrum of the carapace
compared to only 4 on Pseudophilomedes foveolatus. Specimens
collected and described by Brady and Norman conform in every way
to males of Euphilomedes; there is no basis for considering them to be
males of Pseudophilomedes, which unfortunately remain unknown.

Euphilomedes asper (Miiller, 1894) was described by Miiller from
males collected from the Bay of Naples. His description of the species
mentions only the 2nd antenna, the furca, and the external morphology
of the carapace; the secondary appendage of the 2nd antenna, the
furca, and the outside of the carapace are illustrated. It is difficult to
identify a species with certainty having so few characters for compar-
ison. The specimens of Brady and Norman considered by me to be
E. asper (Miiller) have a carapace and secondary branch on the 2nd
antenna identical to that illustrated by Miiller. The furca on Brady
and Norman’s specimens has the same number of furcal claws as in
Miiller’s illustration and a similar distribution of primary and second-
ary claws, but it differs in one respect: the 3rd claw of E. asper
(Miiller) is slightly smaller than the 5th claw, whereas the opposite is
true on Brady and Norman’s specimens. The difference in size of the
3rd and 5th claws is so small in either case, I am inclined to believe
that Miiller’s drawing of the furca is slightly inaccurate. This belief
is supported by study of syntypic material of Philomedes asper which
shows the 3rd and 5th claws to be about the same size (fig. 129).

Euphilomedes sordidus (Miiller, 1890) was described from females
collected along the coast of northern Japan. Miiller (1912, p. 26) in
a key to species of Philomedes distinguished P. asper from P. sordidus
by the size relationship of the 3rd and 5th furcal claws: on P. sordidus
the 3rd and 5th claws are about the same size, whereas, on P. asper,
according to Miiller, the 3rd claw is smaller than the 5th. This dif-
ference between the species disappears if the 3rd claw of EL. asper is
not actually smaller than the 5th, as is shown to be the case in the pres-
ent paper. Euphilomedes sordidus was inadequately described and
most appendages are unknown, so that it is difficult to distinguish it
from E. asper on the basis of what is presently known about the species.
The wide geographic separation of the two species suggests that they
are probably not conspecific. The Japanese form (female) has 4
lateral and 6 distal bristles on the 7th appendage, whereas, the Gulf of
Naples form (male) has 4 lateral and 4 distal bristles, but to what
degree this is due to sexual dimorphism is not known. The antennal
sinus of the female P. sordidus (Miiller, 1890, pl. 25, fig. 17) may be
shallower than of P. aspera (Miiller, 1894, pl. 8, fig. 1).

24 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Euphilomedes species

Figures 1le-j

Marterrau.—Vial from British Museum collection containing
separated left and right valves and several pieces of the soft parts
of an ostracod, and also a label with the number 50002. The vial
with the ostracod was in a larger vial with the label ‘Pseudophilo-
medes foveolata Naples—Shallow water—April 1887, formerly a slide
dissected. Norman coll. 1900. 3. 6. 451.” All appendages were
present in the vial except the caudal furca.

Discusston.—Because of the missing furca and poor condition of
the valves, a complete description of the species was not attempted.
A few appendages have been illustrated to show that the specimen
could not belong to the genus Pseudophilomedes and is a female
Euphilomedes.

No explanation is available for Brady and Norman’s statement
(1896, p. 660) that only males of Philomedes foveolatus (Miiller)
were collected. In order to disturb the specimen as little as possible,
most illustrations were made from the original partly dissected speci-
men without benefit of cover glass.

Description (incomplete female).—Shell (fig. 11e): Oval, elongate,
prominent rostrum, and broad rostral incisure; surface with oval
pits. Endopodite of second antenna with 2 joints (fig. 11f): Ist
joint with 1 long and 5 short bristles; 2nd joint elongate with long
hirsute bristle near middle and distal bristle (distal bristle represented
by stump on specimen examined). Exopodite of 2nd antenna (figs. 11f,
g) with 9 joints: 1st joint elongate, joints 2-9 trapezoidal, each joint
smaller than preceding joint; joints 2-8 with row of short hairs along
media-distal margin; bristles on joints 2-5 with marginal spines and
without natatory hairs; bristles on joints 6-8 with natatory hairs
and without marginal spines; joint 9 with 4 stout bristles and 1 slender
bristle, all with natatory hairs, and 2 subequal short spinous bristles
(fig. 11g). Maxilla with 3 endites typical of genus. Fifth limb with
large quadrate 2nd joint typical of genus. Sixth limb (fig. 11h)
with 4 endites and end joint projecting posteriorly. Seventh limb
(fig. 117) with 6 lateral and 6 distal bristles; terminal row of teeth
with 2 opposing pegs. Frontal organ 2-jointed (fig. 117): 1st joint
elongate; 2nd joint tapering from broad base to slender tip.

Shell dimensions: Maximum length 1.42 mm, maximum height
0.88 mm.

The structure of the endopodite of the 2nd antenna, the maxilla,
and the 5th and 6th limbs clearly shows that this species does not
belong in Pseudophilomedes. Without having the furca available,
it is not possible to be absolutely certain that the species does not

NO. 3580 OSTRACOD FAMILIES—KORNICKER 25

belong to Philomedes rather than Ewphilomedes, but the low number
of cleaning bristles on the 7th limb make the latter genus the more
probable.

Literature Cited

Brapy, G. §., and Norman, A. M.

1896. A monograph of the marine and freshwater Ostracoda of the North
Atlantic and of northwestern Europe, 2-4: Myodocopa, Cladocopa,
and Platycopa. Trans. Roy. Dublin Soe. Sci., ser. 2, vol. 5, pp.
621-746, pls. 50-68.

Darsy D. G.

1965. Ecology and taxonomy of Ostracoda in the vicinity of Sapelo Island,
Georgia. Rept. 2 in Four Reports of Ostracod Investigations,
pp. iii-vi, 1-76, figs. 1-11, pls. 1-33. [Offset report issued by the
University of Michigan.]

HarTMANN, G.

1964. Neontological and paleontological classification of Ostracoda. In
Puri, Symposium Ostracods as ecological and palaeocological indi-
cators. Pubbl. Staz. Zool. Napoli, suppl. 33, pp. 550-587.

KornicxkeEr, L. 8.

1959. Ecology and taxonomy of Recent marine ostracodes in the Bimini
Area, Great Bahama Bank. Publ. Inst. Marine Sci., vol. 5 (1958),
pp. 194-399, figs. 1-89.

1967. HEuphilomedes arostrata, a new mydocopid ostracod from the Maldive
Islands, Indian Ocean. Proc. U.S. Nat. Mus.

Mi.uter, G. W.

1890. Neue Cypridiniden. Zool. Jahrb., Abt. f. syst., vol. 5, pp. 211-252,
pls. 25-27.

1894. Die Ostracoden des Golfes von Neapel und der angrenzenden Meeres-
Abschnitte. Monogr. 21 in Fauna und Flora des Golfes von
Neapel, viii + 404 pp., 40 pls.

1912. Ostracoda. In Das Tierreich, pt. 31, 434 pp., 92 figs.

PoutsEN, E. M.

1962. Ostracoda-Myodocopa, 1: Cypridiniformes-Cypridinidae. Copen-
hagen, Carlsberg Foundation, Dana Report No. 57, 414 pp.
181 figs.

1965. Ostracoda-Myodocopa, 1: Cypridiniformes-Rutidermatidae, Sar-
siellidae and Asteropidae. Copenhagen, Carlsberg Foundation,
Dana Report No. 65, 483 pp., 156 figs.

Port, H. 8.

1963. Preliminary notes on the Ostracoda of the Gulf of Naples. Experi-

mentia, 19, 368, pp. 1-6.
Puri, H. 8.; Bonapucez, G.; and MatLoy, J.

1964. Ecology of the Gulf of Naples. In Puri, Symposium Ostracods as
Ecological and Palaeoecological indicators. Pubbl. Staz. Zool.
Napoli, suppl. 33, pp. 87-199, figs. 1-67.

SxoacsBeEr«G, T.

1920. Studies on marine ostracods, 1: Cypridinids, halocyprids and poly-

copids. Zool. Bidr. Uppsala, suppl. vol. 1, 784 pp., 153 figs.
SYLVESTER-BRADLEY, P. C. ,

1961. Ostracoda (part). Pt.3 in Arthropoda of Part Q in Moore, Treatise

on invertebrate paleontology, xxiii + 442 pp., 334 figs.

26 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficure 3.—Pseudophilomedes ferulanus, complete specimen: a, outline, all punctae not
shown; b, dorsal view, anterior to right. Medial view, right valve: c, muscle scars and
some punctae; d, anterior process; ¢, posterior process; f, anteroventral part; g, anterior
margin rostrum with lamellar prolongation. Medial view left valve; h, muscle attach-
ments below anterodorsal corner. (Scale same, in microns: a, b; c; d, e;f, h; g. Spec. 1,

BST Sta. 30.)

aE Mite tei Maia a ot

| NO. 3580 OSTRACOD FAMILIES—KORNICKER 27

SS ey 7,

SI
a

Ficure 4.—Pseudophilomedes ferulanus, 1st antenna: a, right; }, distal end, 5th and end
joints, left; c, ciliated bristles on end joints; d, tips of closely ringed e- and d-bristles on
end joint. 2nd antenna: ¢, medial view; f, endopodite. Mandible: g, lateral view; h,

| coxale; i, distal part. (Same scale, in microns: 4, é, g} b-d, f;h,7. Figs. a, c-i from spec.

| 1, BST Sta. 30; fig. b from spec. 2, same station.)

28 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Ficure 5.—Pseudophilomedes ferulanus, maxilla: a, left; b, distal process of right, anterior
to left; c, lst endite, anterior to left; d, 2nd endite, anterior to left. 5th limb: e, anterior
view, part of 2nd joint; f, trident tooth on Ist joint; g, anterior view, distal part, only 1
bristle of epipodial appendage shown with hairs; h, bifid tooth on Ist joint; 7, posterior
view, distal part, only 1 bristle of epipodial appendage shown with hairs. (Same scale,
in microns: a, g, 1; b-f, h. Figs. af, h from spec. 1, BST Sta. 30; figs. g, i from spec. 4,
same station. Roman numerals on a, c, d refer to endites; arabic numbers on e-g, 7 refer
to joints.)

NO. 3580 OSTRACOD FAMILIES—KORNICKER 29

Ficure 6.—Pseudophilomedes ferulanus: a, 6th limb; b, 7th limb; c, distal part of 7th
limb; d, bell on distal part of bristle of 7th limb; e, left lamella of furca; f, 1st claw of left
and right lamellae; g, lateral eye; h, subventral view of furca showing muscles; 1, egg
from brood pouch; j, medial eye and frontal organ; &, tip of frontal organ. (Same scale,
in microns: a, b, e, h, i; c, d, k; f, g,j- Figs. a-f, h, i from spec. 1; fig. 7 from spec. 4;
figs. g, k from spec. 2. All from BST Sta. 30.)

30 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

Ficure 7.—Pseudophilomedes foveolatus: a, left view of shell, broken circle locates adductor
muscles, only representative punctae shown. Medial view of valves: b, rostrum of
right; c, posterior process, right; d, lamellar prolongation of selvage on anteroventral
margin of left. Antenna: ge, left 1st; f, medial view, joints 5-8 of left Ist; g, right 2nd;
h, endopodite of same; i, lateral view, joints 7-9 of right 2nd. Left mandible: 7, medial
view, 2 outer bristles on basale broken; k, terminal end. Left maxilla, lateral view:
l, bristles on endite II not shown; m, endite II, anterior margin to left. (Same scale,
in microns: a; b-d, f, h, 1, k-m; e, g,7. Spec. 1, BST Sta. 30.)

NO. 3580 OSTRACOD FAMILIES—KORNICKER 31

Ficure 8.—Pseudophilomedes foveolatus, 5th limb: a, posterior view, right; b, oblique
view from posterior of left, processes on Ist joint not shown; ¢, posterior view, right,
showing only 2nd and distal part of 4th joint; d, posterior view, forward bifid tooth of
[st joint; ¢, posterior view, trident tooth of Ist joint. 6th limb: f, oblique view, right,
not all bristles shown on end joint. Right caudal lamella: g, complete; h, part; 7, lateral
view, claw 1. (Same scale, in microns: a-f, h, 1; g. Spec. 1, BST Sta. 30. Roman
numerals on f refer to endites; Arabic numbers on a, 5, ¢ refer to joints, on h to claws.)

ee

32 PROCEEDINGS OF THE NATIONAL MUSEUM VOL. 121

oly
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OO
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26 a
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9
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OQ
Oe?

Ficure 9.—Euphilomedes asper, shell: a, lateral outline showing muscle scars, punctae only
on part of rostrum and along posteroventral margin. Medial view of valves: b, rostrum,
right; ¢, posterior process, right; d, anteroventral part of inner lamella, right; lamellar
prolongation of selvage not shown; ¢, muscle scars, right; f, rostrum, left, lamellar prolon-
gation of selvage not shown; g, posterior process, left, lamellar prolongation of selvage not
shown; A, anteroventral part of inner lamella, left; 7, muscle scars, left. Antenna: 7, |
right Ist, ends of many bristles not shown; &, endopodite of 2nd; /, lateral view, pro-
topodite, exopodite, and joint 2 of endopodite of 2nd, bristles on joints 3-9 of exopodite |
not shown. (Same scale, in microns: a; b, ¢, e-g, i, k; d, h, j, 1. All from spec. A.)

NO. 3580 OSTRACOD FAMILIES—KORNICKER 33

Ficure 10.—Euphilomedes asper: a, medial view, joints 7-9, left 2nd antenna; J, lateral
view, 2nd and end joints of endopodite, right mandible; ¢c, right mandible; d, left maxilla;
e, 5th limb; f, 6th limb; g, joints 1-4, 5th limb, all bristles not shown; h, 4th joint and 2
bristles on outer lobe of 3rd joint of 5th limb; 7, claws 3-6 and part of 2nd claw of furca;
j, distal part of 7th limb; , terminal of 7th limb; J, left lamella of caudal furca; m, claws
3-6 and part of 2nd claw. (Same scale, in microns: a, b, d, h; c, ¢, 1; f,; g, i,k, m. Fig. m
from spec. 1900-3-6-452; remaining figs. from spec. A. Roman numerals on d-f refer to
endites; Arabic numerals on a, g, h refer to joints, on 7, m to claws.)

34 PROCEEDINGS OF THE NATIONAL MUSEUM VoL. 121

Figure 11.—Euphilomedes asper: a, copulatory appendage; b, frontal organ and medial
eye (M. E.) with joints 1-2 of Ist antenna, all bristles on 2nd joint not shown; c¢, tip of
frontal organ; d, lateral eye. Euphilomedes species: ¢, outline of left valve from inside,
all punctae not shown; f, endopodite and joints 1-3 of exopodite of 2nd antenna, terminal
bristle on 2nd joint of endopodite broken; g, medial view, joints 7-9, left 2nd antenna;
h, 6th limb; 7, distal part of 7th limb; 7, frontal organ and medial eye. (Same scale, in
microns: a, ¢, g; d,f, h-j;b; e. Figs. a-d from spec. A; figs. e-j from spec. 50002. Roman
numerals on h refer to endites; Arabic numbers on J, f, g refer to joints.)

NO. 3580 OSTRACOD FAMILIES—KORNICKER 35

Ficure 12.—Euphilomedes asper, shell: a, outline showing central muscle scars and lateral
eye; broken lines represent c- and f-bristles. Medial view of valves: b, right, broken
circle outlines muscle scars, small circles represent punctae; c, outline of left, showing
muscle scars; d, rostrum and incisure of right; ¢, posterior process of right; f, muscle scars
of left. Caudal furca: g, left lamella. (Same scale, in microns: a-c; d-g. Figs. a, g
from spec. 1; figs. b, d, and ¢ from spec. 2; figs. c, f from spec. 3. All are syntypes.)

PROG UES NAW MUS) VOL. 121 KORNICKER

PLATE 1

Pseudophilomedes ferulanus, sections: a, ligament and left valve overlapping right on
anterodorsal margin of carapace; b, ligament and muscle to median eye on dorsal margin
at highest part of carapace; c, hinge and ligament near posterior of hinge; d, free margin
at anteroventral part of left valve; ¢, free margin at anterodorsal part of left valve;
free margin at ventral part of right valve; g, side of left valve showing spiral pore canal
leading to hair and reticulate pattern of shell, inner lamella at left; 4, attachment of
adductor muscles on left valve; 7, sockets of two medial bristles of caudal process.
(Photographs taken with oil immersion objective, 100x.)

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