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SMITHSONIAN INSTITUTION 
UNITED STATES NATIONAL MUSEUM 



PROCEEDINGS 



OF THE 



UNITED STATES NATIONAL MUSEUM 



VOLUME 121 

NUiMBERS 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 tlae 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 tal)le of contents of the volume. 

In the Bulletin series, the first of wliich 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 Bidletin series under the heading Contributions from the United 
States National Herbarium. 

Frank A. Taylor 
Director, United States National Museum 
II 



CONTENTS 

Number 

Barnard, J. Laurens. New species and records of Pacific 
Ampeliscidae (Crustacea: Amphipoda). 20 pages, 4 figures. 
Published March 8, 1967 3576 

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

Cresset, Roger. Revision of the family Pandaridae (Cope- 
poda: Caligoida). 133 pages, 356 figui'es. Published 
February 16, 1967 357O 

New genus: Pannosus. 

New species: Pandarus floridanus, P. katoi, Dinemoura discrepnns, 
Nesippus iigris. 

Cresset, Roger. Caligoid copepods parasitic on sharks of 
the Indian Ocean. 21 pages, 54 figures, 2 plates. Published 

March 15, 1967 3572 

New species: Nemesis aggregatus, Eudnctylina pusilla, E. pollex, 
Kroyeria gemnrsa. 

Humes, Arthur G., and Ho, Ju-Shet. New cyclopoid 

copepods associated with the alcyonarian coral Tubipora 

musica (Linnaeus) in Madagascar. 24 pages, 69 figures. 

Pubhshed March 8, 1967 3573 

New species: Lichomolgus organicus, L. conjunclus, Hippomolgus 
Jalipes, H. cognatus. 

KiER, Porter. Bredin-Archbold-Smithsonian Biological Sur- 
vey of Dominica: 1, The echinoids of Dominica. 10 pages, 
3 figures, 2 plates. Pubhshed December 30, 1966 3577 

KoRNiCKER, Louis S. Supplementary descriptions of two 
myodocopid ostracods from the Red Sea. 18 pages, 6 figures. 
Published February 16, 1967 357 1 

KoRNiCKER, Louis S. The myodocopid ostracod families 
Philomedidae and Pseudophilomedidae (new family). 35 
pages, 12 figures, 1 plate. Published February 16, 1967 . . 3580 

New family: Pseudophilomedidae. 

New combination: Euphilomedes asper. 

Lewis, Allan G. Copepod crustaceans parasitic on teleost 
fishes of the Hawaiian Islands. 204 pages, 70 figures. Pub- 
lished June 22, 1967 3574 

New species: Pseudotaeniacanthus puhi, Anchistrotos moa, Hat- 
schekia breviramus, Lepeophtheirusf fallolumdus, Caligus kapuhili, 
Brachiella regia. 

Ill 



IV CONTENTS 

McFadden, Max W. Soldier fly larvae in America north of 
Mexico. 72 pages, 156 figures. Published February 1, 
1967 3569 

Pettibone, Marian H. Some bathyal polynoids from Central 
and Northeastern Pacific (Polychaeta: Polynoidae). 15 
pages, 5 figures. Published February 16, 1967 3575 

Now genera: Balhyadmetella, Balhymoorea. 

Now species: Bathijadmelclla commando. 

New combination: Bathymoorea rcnotubulata. 

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 3568 

New species: Melanotus indislinctus , M. dietrichi, M. miscellus, 
HI. lanei, M. beameri, M. lanceatus. 

Rehder, Harald a. Valid zoological names of the Portland 
Catalogue. 51 pages, 2 figures. Published March 21, 1967 . 3579 

Stone, Alan. 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 3578 

New species: Mycetobia limanda, Olbiogaster danista, 0. evansi, 
Plecia porca. 

U.S. GOVERNMENT PRINTINS OmCE 262-119-h 




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 Sarah E. Thompson ^ 



Click beetles and wire worms 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 (1853, 
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 Hmited area. Descriptions of a few neAv 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 Avide knowledge to print. 

' Quatc: 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 Kocky 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, beameri, 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 cribricoUis 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 mil 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 aU the species, are based on the 




Figure 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 wiU be indi^ddual variants that 
do not seem to quite fit either part of a couplet. Many revisions have 
failed to eliminate aU deficiencies, but for the majority of specimens 
the key should provide a satisfactory shortcut to their proper 
identification. 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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 (Natiu-al History) (BMNH); Zoologisches Museum 
der Humboldt-Universitat zu Berlin (ZM). 

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




Figure 2. — Distribution of the three types of female bursae of Melanotus similis. 
O 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 



MELANOTUS — QUATE AND THOMPSON 



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 pan- 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 "biu-sa," 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 muTor 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) ai-e 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 Usting 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 Museiun der Humboldt-Universitat zu Berlin and the 
British Museum (Natm'al 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 Natm'al History, Stockholm. 
WhUe 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 Aiizona; 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 Smwey; 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. 



MELANOTUS — QUATE AND THOMPSON 



List of North American Melanotus 

SiMiLis Group 
(Mandible without pit) 



1. similis (Kirby) 

2. spadix (Erichson) 

3. decumanus (Erichson) 

4. castanipes (PaykuU) 

5. communis (Gyllenhal) 

6. indistinctus, new species 

7. dietrichi, new species 

8. miscellus, new species 



9. verberans (LeConte) 

10. emissus (LeConte) 

11. lanei, new species 

12. pilosus Blatchley 

13. opacicollis (LeConte) 

14. clandestinus (Erichson) 

15. ignohilis Melsheimer 



Americanus Group 
(Mandible with pit) 



16. depressus (Melsheimer) 

17. morosus Candfeze 

18. cribulosus (LeConte) 

19. corticinus (Say) 

20. Sagittarius (LeConte) 

21. hyslopi van Zwaluwenburg 

22. prasinus Blatchley 

23. piceatus Blatchley 

24. difficilis Blatchley 

25. macer (LeConte) 

26. testaceus (Melsheimer) 

27. trapezoideus (LeConte) 

28. tenax (Say) 

29. periinax (Say) 

30. infaustus (LeConte) 



31. parallelus Blatchley 

32. americanus (Herbst) 

33. cribriventris Blatchley 

34. obsciiratus Blatchley 

35. beameri, new species 

36. concisus Knull 

37. lanceatus, new species 

38. hamatus Knull 

39. longulus longulus (LeConte) 

40. I. oregonensis (LeConte) 

41. gradatus (LeConte) 

42. insipiens (Say) 

43. leonardi (LeConte) 

44. taenicollis (LeConte) 



45. cribricollis Candfeze 



Cribricollis Group 

46. chiricahuae Knull 



List of Synonymical Names in North American Melanotus 

abdominalis (Erichson) = 2. spadix (Erichson) 

angustatus (Erichson) = 32. americanus (Herbst) 

blatchleyi Leng=33. cribriventris Blatchley 

canadensis Candfeze = 3. decumanus (Erichson) 

carinus Blatchley=20. Sagittarius (LeConte) 

cuneatus (LeConte) = 3. decumanus (Erichson) 

debilis Blatchley=17. morosus Candfeze 

divarcarinus Blatchley=16. depressus (Melsheimer) 

dubius (LeConte) = 26. testaceus (Melsheimer'l 

exuberans (LeConte) = 1. similis (Kirby) 

fissilis (Say) = l. 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. crihriventris Blatchley 

longulus (LeConte) = 17. morosus Caiidfeze 

ochraceipennis Melsheimer=l. similis (Kirby) 

paganus Candeze=5. communis (Gyllenhal) 

paradoxus (Melsheimer) = 4. castanipes (PaykuU) 

parumpundatus (Melsheimer) = 16. depressus (Melsheimer) 

peninsularis Candfeze=14. clandestinus (Erichson) 

perplexus Blatchley=14. clandestinus (Erichson) 

prolixus (Erichson) = 1. similis (Kirby) 

scrobricollis (LeConte)=4. castanipes (Paykull) 

secretus (LeConte) = 15. ignobilis Melsheimer 

simulans Blatchley=17. morosus Candfeze 

sphenoidalis (Melsheimer) = 1. similis (Kirby) 

tenellus (Erichson) =42. insipiens (Say) 

texanus Candfeze=4. castanipes (Paykull) 

variolatus LeConte = 40. longulus oregonensis (LeConte) 

vetulus (Erichson) = 19. coriicinus (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.] 
Ctenonychu^ 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: Elater fulvijyes 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, punctiu-es 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 '\\dth 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 Mith 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 spui's; 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, pronotum partly or completely 

reddish orange and elytron black 2 

Pronotum and elytron of same color 3 

2(1). Pronotum almost entirely orange and abdomen entirely black; antenna 
of cf usually exceeding tip of hind angle by 1 segment or less, ? 

antenna ending about at tip 43. leonardi 

Pronotum black in center and orange elsewhere, and abdominal venter 
partly or entirely orange; antenna of cf ending before tip of hind 

angle by about 1 segment and less in ? 44. taenicollis 

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

Antennal segments 4 and most of following wider than long, and lobes 
lighter in color than rest of antenna; antenna ends 2 or more segments 
before tip of hind angle; frontal margin 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 on lateral margin 21 

SiMiLis Group 
(Mandible without pit on lateral margin) 

5(4). Antennal segment 3 small, subequal to 2 or at least much nearer size of 

2 than 4 6 

Antennal segment 3 intermediate in size between 2 and 4, clearly larger 
than 2 8 

6(5) . Antennal segments beyond 3 triangular, same color as body, triangular, 

erect male hairs well developed 7 

Antennal segments beyond 3 testaceous, lighter in color 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; cf 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 smaU; cf 

paramere with apical blade 14. clandestinus 

Large species, elytron more than 9.0 mm; frontal margin not strongly 
jutting over nasale; nasale and parantennal fovea poorly developed, 
but usually present; sides of pronotum often straight and divergent 
posteriorly with large hind angles; cf paramere with apical 
blade 4. castanipes 

8(5). Frontal and pronotal punctures not greatly enlarged; punctures on 
venter of abdomen oblong and lateral ones but little larger than 
central ones; cf paramere little shorter than aedeagus and with 
apical blades 9 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Frontal and pronotal punctures very large (fig. 100; punctures on 
venter of abdomen teardrop shaped and lateral ones several times 
larger than central ones; cf paramere about two-thirds length of 
aedeagus and abruptly tapering to sharp, recurved apex; New Mexico 

to Baja California 46. chiricahuae 

9(8). Pronotum wider than long 10 

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

31. parallelus (part) 
10(9). Pronotal punctures not so thick as to appear granulose; vestiture of 

thorax and elytra of about same length and density 11 

Pronotal punctures very dense and appearing granulose; vestiture of 
pronotum short and fine, nearly velvety, considerably denser than 

on elytra 13. opacicoUis 

11(10). Pronotal punctures smaller than those on frons, dense and nearly con- 
tiguous on all sides; small species, elytron length less than 7.5 mm . 12 
Pronotal punctures sparser and larger; most species with elytron length 

7.5 mm or more 13 

12(11). Vestiture of thorax rather sparse, that on venter sparser than on dorsum; 
antennal segment 3 intermediate in size between 2 and 4; frontal 

margin not upturned 11. lanei 

Vestiture of thorax long and thick, that on venter as long and thick 

as on dorsum; antennal segment 3 shorter, closer to length of 2 than 

4; frontal margin thin, shelflike and a little upturned ... 12. pilosus 

13(11). Frontal margin not strongly protuberant, usually without a depression 

before margin 14 

Frontal margin strongly protuberant, when viewed from side extends 
out in shelflike extension, marked depression behind margin; 9 bursa 
without spines or with only patch of 5-8 small spines on each side 
near base; cf paramere with margins before apical blade straight 
and without indentation; large species, elytron 10-12mm. 

3. decumanus 

14(13). Frontal margin well defined, dark brown or black, not weakened in 

center; elytron rarely as long as 13 mm and most species considerably 

less 15 

Frontal margm usually weakened in center, 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 spmes 
in large basal packet and small, separated distal patch (fig. 5/). 

4. castanipes, females 
15(14) . Antennal segment 4 about lYz times as long as wide and considerably wder 

than 3; elytron often less than 10 mm long 16 

Antennal segment 4 usually slender, about two times as long as wide and 
only a little wider than 3; average elytron length 10.8 mm; cf gono- 
style with thick patch of 30-40 long, soft hairs on apical blade (fig. le) ; 
9 bursa with dense packet of long, close-set spines arranged nearly in 

trapezoid (fig. 5e) 2. spadix 

16(15). Pronotal punctures smaller than those on front, those in center generally 
separated by at least own diameter; elytron usually less than 8 mm 
long; cf aedeagus parallel sided until near apex and sharply tapering 
to acute tip 17 



i 



MELANOTUS — QUATE AND THOMPSON 



11 



Pronotal punctures subequal in size to those on front; elytron usually 
more than 8 mm; cf aedeagus often with convergent sides and tapering 

to tip 18 

17(16). Antenna same color as head and thorax; cf paramere evenly tapering to 
apical blade; outer margin of apical blade nearly straight . 9. verberans 

Antenna usually lighter in color than head and thorax; cf paramere 
with marked constriction at base of blade; outer margin of blade 

rounded 10. emissus 

18(16). Male aedeagus broad until near apex and then 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 

9 bursa in center , 20 

19(18). Pronotal punctures large and separated by less than own diameter; 
dark reddish brown to black species; spines of 9 bursa without large 
base (fig. 8b) ; d^ genitalia as in figs. 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) ; cf genitalia as in figs. 4j-l 8. miscellus 

20(18). Male genitalia as in fig. 4j 5. communis 

Male genitalia as in fig. 4k 6. indistinctus 

Male genitalia as in fig. 41 7. dietrichi 

Americanus Group 

(Mandible with pit on lateral margin) 

21(4). Antennal segment 3 small, subequal to 2 (in doubtful cases of some 

99, 3 less than VA length of 2) 22 

Antennal segment 3 intermediate in size between 2 and 4, clearly larger 
than 2 (in doubtful cases of some 99, more than 1}^ length 

of 2) 27 

22(21). Small species, elytron less than 6 mm 23 

Larger species, elytron more than 6 mm 24 

23(22). Pronotum wider than long; pronotal punctures smaller than those on 
front and sparse, widely separated, equal size throughout; East 

Coast to Nebraska 32. americanus 

Pronotal punctures coarse, larger than on front on anterior two-thirds 

and fine, sparse on basal one-third in cf ; Southwest . 36. concisus 

24(22). Antenna exceeds tip of hind angle by 1-4 segments; parantennal fovea 

large; flagellar segments elongate 25 

Antenna of cf barely exceeds tip of hind angle, 9 antenna shorter; 
parantennal fovea, clearly smaller than antennal segment 2; flagellar 
segments triangular, not elongate; pronotal punctures large and dense; 
cf paramere with pecuhar, winglike flaps near center; Texas. 

38. hamatus 

25(24). Elytron length 6-8 mm 26 

Large species, elytron 8.5-11.5 mm; nasale quadrate, about as wide as 
high; antenna exceeds tip of hind angle by about 2 segments; cf para- 
mere with definite apical blade and antennal hairs long and dense. 

19. corticinus 
26(25). 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 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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

34. obscuratus 

27(21). Elytron more than 5 mm 28 

Very small species, elytron less than 4.5 mm; nasale flat; frontal margin 
rounded, moderately thin and jutting; pronotal punctures nearly as 
large as those on front; parantennal and mandib\ilar foveae relatively 
large; southeast U.S. to Texas 42. insipiens 

28(27). Pronotal punctures and vestiture uniform except usual small increase 

in density and size along sides 29 

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

29(28). Parantennal fovea very large, as large or larger than antennal segment 3; 
nasale large and protuberant, especially in cf ; 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 
above combination 31 

30(29). Frontal margin scarcely overhanging protruding nasale, not extended 
in angular apex; pronotal punctures as large as those on front; cf 
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; cf 
paramere without apical blade; elytron 8.2-9.3 mm . . . 21. hyslopi 

31(29). 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 34 

32(31). Parantennal fovea and nasale large and well developed, nasale about as 
high as wide; antenna not greatly elongate; smaller species, elytron 

7.5 mm or less 33 

Parantennal fovea and nasale weak, nasale wider than 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 

33(32). Pronotal punctures smaller than those on front; frontal margin often 
rounded in center; cf paramere expanded at base, abruptly rounded 

at tip 26. testaceus 

Pronotal punctures about same size as those on front (but widely 
separated); frontal margin often coming to blunt point in center; 
cf paramere slender, gently tapering to tip . . . . 27. trapezoideus 

34(31). 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 35 



MELANOTUS — QUATE AND THOMPSON 



13 



Frontal margin not strongly protruding 36 

35(34). Pronotal punctures separated by no more than own diameter; flagellar 
segments elongate, d* antenna exceeds tip of hind angle by 3-4 seg- 
ments; elytron about 7 mm 25. macer 

Pronotal punctures sparse, separated by 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 

36(34). Antenna extending to or beyond hind margin of pronotum .... 37 

Antenna short, ending before hind margin of pronotum in botii sexes; 

nasale flat, shallow; <f paramere slender, aedeagus broad until near 

apex 23. piceatus 

37(36). Nasale clearly defined, not obsolescent; color reddish brown or darker. 38 
Nasale obsolescent; color brownish yellow; cT paramere without apical 
blade; pronotal punctures large and compact; vestiture rather thick; 

small species, elytron 5.0-6.5 mm; Texas 35. beameri 

38(37). Antenna, legs, and body of same color 39 

Antenna and legs reddish brow^n and body black (teneral specimens 
with body dark browm but darker than appendages); pronotum 
convex and rounded dorsally, rounded and concave at base of hind 
angles; central pronotal punctures small and sparse; c? paramere 

without apical blade 29. pertinax 

39(38). Western U.S., not occurring east of New Mexico 40 

Eastern U.S. and as far west as western Texas, Kansas, etc .... 41 
40(39). 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 

41(39). Larger species, elytron more than 7.5 mm 42 

Smaller species, elytron less than 7.5 mm 44 

42(41). Parantennal fovea well developed; pronotal punctures well sepa- 
rated 43 

Parantennal fovea shallow; nasale flat; pronotal punctures very close 
together; cf paramere tapering and lateral margin straight; color 

usually reddish brown 18. cribulosus 

43(42). Sides of pronotum parallel or concave over distal 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 usually dark reddish brown, darker than 

morosus 16. depressus 

44(41), Nasale well developed, about as wide as high 45 

Nasale flat, not protuberant, wider than high 46 

45(44). Pronotal punctures about as large as those on front; pronotum flattened 

and not rounded 27. trapezoideus (part) 

Pronotal punctures small, smaller than those on front; pronotum 

rounded 32. americanus (part) 

46(44). Side of pronotum straight, without 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 U.S 48 

Male paramere without apical blade; thorax wider than long; largely 

distributed along east coast 28. tenax 

48(47). Male paramere not unusually slender, base of apical blade angled at 

about 90° 30. infaustus 

Male paramere very slender, base of apical blade gently rounded and 
not strongly angulate 37. lanceatus 

The similis Group 

(Mandible with pit on lateral margin) 
1. Melanotus similis (Kirby) 

Figures Aa-d, 8b, c; Plate la 

Perimecus similis Kirby, 1837, p. 149. 

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

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

Cratonychus fissilis. — LeConte, 1853, p. 477. 

Melanotus fissilis. — Candeze, 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 synonj^my.] 
Melanotus exuberans. — Candfeze, 1860, p. 354. — Thomas, 1941, p. 260. 

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

Head: Front coarsely punctate, most punctiu-es hexagonal shaped, 
distance between punctures less than half puncture diameter, center 
slightly protuberant; clypeus lightly punctate, parantennal fovea 
shallow, width of nasale IK 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 V/2 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); punctiu-es moderately 
coarse at center, about size of those of front, separated by distance 
equal to less than puncture diameter, punctiu-es 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. 35G8 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 pimcture 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. 

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

Types ofjissilis: Specimens lost. 

Lectotype of laticollis: Male, "Amer. Sept., Norwish" (ZM). 
Lectotype selected by Quate, 1960, from co types of 1 male and 5 
females. 

Lectotype of ochraceipennis: Female, "Pa./Ziegler" (MCZ). Se- 
lected by Quate, 1962. Fu'st 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 exuberans: 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. 

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

Alabama: 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. California: Riverside (Blythe); May, July. Colorado: El Paso, 
Montezuma (Berkeley); July, August. Connecticut: Fairfield, Hartford, 
Litchfield (January, reared?), Middlesex, New Haven, Tolland; May-November. 
Florida: Brevard, Hendry, Manatee, Orange, Osceola, Palm Beach, Seminole, 
Volusia; February, August. Georgia: Chatham, Floyd, Fulton, Peach, Thomas, 
Warren, Okefenoke Swamp; March, September. Illinois: Boone, Champaign, 
Cook, DeKalb, Edgar, Iroquois, Jackson, Jo Daviess, Johnson, Kanakakee, Lake 
McHenry, McLean, Marion, Marshall, Mason, Peoria, Putnam, Union, Vermihon; 
March, September. Indiana: Clark, DeKalb, Greene, Knox, Lake, Marion, 
Orange, Porter, Pulaski, Stark, Steuben, Tippecanoe, White; April, October. 



16 PROCEEDINrGS 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: Atchtnson, 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. Louisiana: Lafourche, 
Orleans, Plaquemines, St. Landry, St. Tammany; January, September. Maine: 
Penobscot, Washington; June, Jvily. Maryland: Anne Arundel, Calvert, Har- 
ford, Montgomery, Plummers Isl., Prince Georges, Queen Annes, Washington; 
April, December. Massachusetts: Berkshire, Essex, Franklin, Hampden, 
Hampshire, Middlesex, Norfolk, Plymouth, Suffolk, Worcester; April, September. 
Michigan: 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. Missouri: 
Boone, Buchanan, Callaway, Cape Girardeau, Lawrence, Pike, St. Louis, Wayne; 
April, October. Montana: Custer; August. Nebraska: Dakota, Dawes, 
Dodge, Holt, Lancaster, Lincoln, Otoe, Red Willow, Scotts Bluff, Sioux, Washing- 
ton; May, August. New Hampshire: Coos, Strafford; May, June. New 
Jersey: 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. North Carolina: Buncombe, Chowan, Iredell, Pasquotank, 
Polk, Richmond, Swain, Wake, Watauga, Wilson; April, July. North Dakota: 
Cass, Grand Forks; Jvme. Ohio: Ashtabula, Athens, Delaware, Fairfield, Frank- 
lin, Fulton, Greene, Hamilton, Hocking, Holmes, Lucas, Pike, Preble, Summit, 
Warren; May, October. Oklahoma: Atoka, Cleveland, Latimer, Oklahoma, 
Tulsa; March, July. Pennsylvania: 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?). Rhode Island: Washing- 
ton; June, July. South Carolina: Oconee; April, August. South Dakota: 
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. Utah: Cache, San 
Juan, Utah, Washington, Zion Canyon; June, July. Vermont: Orange, Wind- 
ham; May. Virginia: Fairfax, Montgomery, Nelson; June. Washington, 
D.C. : March, December. West Virginia: 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 Scotia: Annapohs Royal; 
June. Ontario: Belleville, Muskoka, Ottawa, Prince Edward, Ridgeway, 
Sudbury, Toronto; May, July. Quebec: Aylmer, Montreal, Mt. St. Hilaire, 
Rigaud, St. Johns; June, August. 

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



NO. 35G8 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 a,sjissilis. 

Melanotus similis exhibits a good deal of variation throughout its 
range. Variations in the ratio of the thoracic wddth to length, in the 
size of the pronotal punctiu-es, in the shape of the distal enlargement 
of the male paramere, and the spines of the female biu-sa 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 wdder 
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/length 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 
great 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 ^vith 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 



[8 PROCEEDINGS OF THE NATIONAL 


MUSEUM 


VOL. 121 


Iowa Lincoln, 


North Platte, 




New York Illinois Missouri Kansas Nebraska 


Nebraska 


Arizona 


1. 20 1. 19 1. 17 1. 16 1. 17 


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 exuberans. Many specimens of this form 
appear quite different from tjrpical 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 "exu- 
berans" types intermediated with clear intergrades to typical similis. 
Specimens have been seen with the coarse pronotal punctation of 
"exuberans," 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 "exuberans" is a synonym of similis 
and represents a geographical variant not defined well enough to 
justify segi'egating 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.Jissilis Say. 

2. Melanotus spadix (Erichson) 

Figures 4e, 8e 

Cralonychus spadix Erichson, 1842, p. 103. 
Melanotus spadix. — Candfeze, 1860, p. 354. 
Melanotus abdominalis of authors, not Erichson, 1842. 
Melanotus decumanus. — Dietrich (not Erichson), 1945, p. 55. 

Male. — -Large species, body color reddish browTi to dark reddish 
brown, covered with yellowish hau's evenly distributed over body. 

Head: Front coarsely punctate, most punctm'es 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 
hau's 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 punctm'e 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-thu'ds 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. — SimUar 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. 

Type.— Holotype of spadix: Male, "Amer. Sept." (ZM 17035). 
Only specimen in type series. 

Distribution. — 'Eastern U.S. west to Nebraska and Arkansas. 

Arkansas: Washington; July. Florida: Manatee; March. Georgia: 
Clarke, Fulton, Habersham; April, July. Illinois: Piatt, Whiteside; June. 
Indiana: Clark, Lake; June, July, December. Maryland: Anne Arundel, 
Baltimore, Montgomery, Plummer's Is.; May, June. Michigan: Marquette, 
Oceana; June, July. Montana: Barry; June. Nebraska: Washington; June. 
New Jersey: Atlantic, Burlingtion, Middlesex, Ocean; May-August. New 
York: Erie, Queens, Suffolk; May- July. North Carolina: Moore, Polk; 
May. Pennsylvania: Dauphin; August. South Carolina: Oconee; June. 
Tennessee: Sevier; June. Virginia: 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 spadix 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 spadix 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 spadix, 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) 

Figures 4/-/i, M; Plate 16 

Cratonychus decumanus Erichson, 1842, p. 104. — LeConte, 1853, p. 474. 
Melanotus decumanus. — Candfeze, 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. 
Cratonychxis incertus LeConte, 1853, p. 474. 
Melanotus canadensis Candfeze, 1860, p. 342. — Blatchley, 1910, p. 751. — Thomas, 

1941, p. 258.— Dietrich, 1945, p. 55. 

Male. — 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, sm-face 
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 IK 
segments, erect male hairs dense and easUy observed, average ratio 
of segments 2:3:4 varies from 5:10:20 to 5:8:19, 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 punctiu-e 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 figiu-ed, 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. 

Female. — Similar to male. Larger, antenna little shorter. Gen- 
italia as figm-ed; 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. 

Types. — Holotype of decumanus: Female, "Amer. Sept." (ZM). 
Only specimen in type series. 



NO. 8568 MELANOTUS — QUATE AND THOMPSON 21 

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

hectotype of incertus: Male, labelled mth green circle "Middle 
States." (MCZ). First specimen to type series selected as lectotype 
by authors in 1962. 

Holotype oi canadensis: Male, "Canada" (BMNH). 

Distribution. — ^Eastern U.S. and Canada west to South Dakota, 
Colorado, and New Mexico. 

Alabama: Mobile; May. Connecticut: Fairfield, Hartford, New Haven; 
March, August. Georgia: Bibb, Clarke, Crawford, DeKalb, Fayette, Fulton, 
Richmond, Walton; April, June. Indiana: Greene, Vigo; June, October. Illi- 
nois: Champaign, Marshall; May, June. Iowa: Dickinson, Woodbury; July. 
Kansas: Douglas, Greenwood, Leavenworth, McPherson, Reno; May, June. 
Kentucky: no data. Maryland: Montgomery; June. Massachusetts: 
Hampshire, Middlesex; June. Michigan: Cheboygan; July. Nebraska: 
Cuming, Dakota, Lancaster, Lincoln, Otoe, Thomas; May, June. New Hamp- 
shire: Straflford; June. New Jersey: 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. North Carolina: Beaufort, Brunswick, 
Buncombe, Columbus, Duplin, Hyde, Jackson, Johnston, Montgomery, Wake; 
May, June. Ohio: Clark, Erie, Hamilton; May, July. Pennsylvania: Dela- 
ware, Northampton, Philadelphia; June, September. Rhode Island: Wash- 
ington; June, July. South Carolina: Aiken, Dorchester, Edgefield, Florence, 
Oconee; May, June. South Dakota: Brookings, Union; June. Tennessee: 
Smith. Virginia: Alexandria, Dinwiddie, Essex, Faquier, Nelson, Nottaway, 
Prince Edward; May, June. Nova Scotia: 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 (Paykiill) 

Figures 4i, 8/; Plate Ic 

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. 



22 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. 

Melanosis inaequalis. — Lang, 1920, p. 174. — Thomas, 1941, p. 258. 

Cratonychus scrobicoUis LeConte, 1853, p. 476. 

Melanotus scrobicoUis. — Leng, 1920, p. 174. — Thomas, 1941, p. 258. — Fattig, 

1951, p. 19. 
Melanotus texanus Candfeze, 1860, p. 351. 

Male. — 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, punctiu-es 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. 

Female. — 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. GenitaHa 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. 

Types. — 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 vaUdity of 
these types may be questioned. 

Lectotype of paradoxus: 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. 3oG8 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 Candeze 1903, 
130," (BMNH). 

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

Arizona: Coconino, Pima; June, July. California: "S. Cal." Colorado: 
Larimer, "Waldo Canon"; June, July. Illinois: McHenry; May. Maine: 
Penobscot; June, July. Massachusetts: Suffolk; May. Michigan: Alpena, 
Crawford, Marquette, Oakland; June, July. Minnesota: St. Louis. New 
Hampshire: Carroll, Cheshire, Coos; April, August. New Mexico: San Miguel. 
New York: Cattaraugus, Cortland, Erie, Essex, Niagara, Oswego, Tompkins, 
Warren; March, September. North Carolina: Wake; June. Ohio: Summit; 
May. Pennsylvania: Carbon; June. Rhode Island: Washington; August. 
Utah: Juab, Utah; January, July. Wisconsin: Dodge, Milwaukee; May, July. 
Nova Scotia: Bridgetown; July. Ontario: Prince Edward; May. Quebec: 
Mt. St. Hilaire, Regaud, Deparquest, Hull, Aylmer, Hudson; May, July. Mex- 
ico: 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 spadix or similis. The weakened 
frontal margin, when viewed from directly m 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 mto 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 margm 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. 

Specunens 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 synonymizmg of the two species as proposed long 
ago by Schwartz (1892) woidd seem warranted; however, w^e accept 
the decision of recent students of the European fauna who regard the 
two species as distinct (Binaghi, 1939; Jeuniaux, 1955). 

5. Melanotus communis (Gyllenhal) 

Figures 4j, 9a,b; Plate Id 

Slater communis Gyllenhal, 1817, p. 138. — Say, 1839, p. 184, 

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

Cratonychus communis. — Erichson, 1842, p. 102. — LeConte, 1853, p. 477. 

Melanotus communis. — Emmons, 1854, p. 88. — Candfeze, 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 Candfeze, 1860, p. 359. 

Male. — ^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, sm'face with 
transverse depression anteriorly, sometimes very weak or reduced to 
pan- of shallow, anterolateral depressions, maigin black or dark 
reddish bro^^^l, not strongly protuberant above clypeus; clypeus 
lightly punctate, parantennal fovea shallow, comma shaped, width 
of interfoveal area 1^-2 times height. Mandible without pit; palpi 
yellowish brown. Antenna 1 or 2 segments longer than pronotum, 
reddish bro\Mi, 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 25 

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 mdentation 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 figm'ed; 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 nun). 40 spec. 

Types. — Lectotype of communis: Male, "Amer. Bor. ?? — 401/62" 
(Schoenherr Collection, Riksmuseum, Stockholm). Fu'st 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 w^e believe they are not the type specimens of 
Gyllenhal. 

Distribution. — Eastern U.S. west to South Dakota, Oklahoma, 
and Texas. 

Alabama: Mobile; March. Arizona: Graham; August. Arkansas: Hemp- 
stead, Washington; April, June. Connecticut: Hartford, Litchfield, Middlesex, 
New Haven, Tolland; (Jan.), March-November. Colorado: Pingree Pk. 
(County?); August. Florida: Dade, Palm Beach; May, July. Georgia: 
Fulton, Harris; June. Illinois: Champaign, Cook, Gallatin, Lake, Piatt, Pike, 
Pope, Union, Vermilion, Wabash; March-September. Indiana: 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. Louisiana: East Baton Rouge, Saint 
Tammany; June. Maryland: Calvert, Harford, Prince Georges, Washington; 
February-November. Massachusetts: Essex, Franklin, Hampden, Hamp- 
shire; April-September. Michigan: Kent, Monroe, Ottawa, Washtenaw; 
April-June. Minnesota: St. Louis, no date. Mississippi: George; April-May. 
Missouri: Boone, Callaway, Mississippi, Phelps, St. Louis, Stoddard; March- 
November. Nebraska: Cherry, Lancaster, Lincoln, Sarpy; April- July. New 
Jersey: Bergen, Burleigh, Burlington, Camden, Essex, Middlesex, Morris, 
Ocean, Union; February-September. New York: Dutchess, Erie, Nassau, 



26 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Niagara, Orange, Rockland. Schenectady, Tompkins, Wayne, Westchester; 
March-December. North Carolina: Wake (Mills River); June, September. 
Ohio: Athens, Franklin, Hocking, Richland; March, November. Oklahoma: 
Woodward; June. Pennsylvania: Allegheny, Bucks, Philadelphia, York; 
March-September. South Carolina: Oconee; September, December. South 
Dakota: Bon Homme, Brookings, Union ; June-July. Tennessee: Montgomery; 
April, Texas: Harrison, Gonzales; April, June. Virginia: Alexandria, Canal, 
Fairfax, Loudoun, Nottaway, Spotsylvania; March-December. Washington, 
D.C: March-September. West Virginia: Marion; no date. 

6. Melanotus indistinctus Qiiate, new species 

Figures 4A;, 9f,g 

Male. — 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 pimctures 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 (7.4-9.4). 50 
spec. 

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

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

Holotype, male (USNM): Pittsfield, Pike Co., 111., July 6, 1946, A. T. McClay. 

Paratypes, 122 males (USNM, CU, ILL, U. Mo., U. Nebr., TEX AM, CAS, 
Lane CoU'n): Illinois: 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. Virginia: Holland, Nansemond Co., April 28, 1947, W. D. Fronk. 
Kentucky: no other data. Mississippi: Lincoln Co., May 18, 1938, peach 
tree, W. F. Turner. Missouri: St. Louis, August 12, 1935, June 22, 1937, 
USD A 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, CHnton Co., July 3, 1948, E. C. Becker. Louisiana: 
Shreveport, Caddo Co., June 2, 1949, J. H. Robinson; Ohvier, 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. McCoUoch. Oklahoma: Muse, LeFlore Co., May 7, 
1939, Sanderson. Texas: LuUng, 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. S. 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, 111., 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 bm'sa 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 Quale, new species 

Figures 4:1, 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. 

Female. — Unknown (or inseparable from communis). 

Distribution. — ^Connecticut to Pennsylvania, west to Iowa. 

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

Paratypes, 67 males (USNM, CU, ILL, CIS, U. Mo., Ohio U., Lane CoU'n) : New 
York: Poughkeepsie, Dutchess Co., June 19, 1933, P. J. Chapman; Staatsburg, 
Dutchess Co., June 23, 1934, II. Dietrich; Rhinebeck, Dutchess Co., Nov. 5, 
1934, Dietrich; Yonkers, Westchester Co., Nov. 21, 1940; Kensico,? Co., April 
27, 1935, J. A. AngeU. 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. Washington, D.C: Rock 
Creek, Dec. 30, 1906; same, April 21, 1907, C. E. Burden. Maryland: Cumber- 
land, Allegany Co., Nov. 4, 1915, W. F. Pennington; Wohville,? Co., May 2, 
1913. West Virginia: same as holotype. Ohio: Columbus, H. II. 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. KnuU. Indiana: 
Beverly Shores, Porter Co., Aug. 6, 1932. Illinois: 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. Louisiana: Baton Rouge, 
June 20, 1916, T. H. Jones. 

Other specimens: Connecticut: Middlesex Co., April. New York: Colum- 
bia Co., June, Queens Co. Pennsylvania: Westmoreland Co., June, July. 
Maryland: Allegany Co., Nov. Ohio: Athens Co., June, July. West 



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 

Figures An, 9c 
Melanosis communis var. B. — Dietrich, 1945, p. 57. — Severin, 1949, p. 16. 

Male. — 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; paratopes 8.5±0.4 mm (7.6-9.7). 
40 spec. 

Female. — Similar to male except sexual differences. Genitaha 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. 

Distribution. — Ontario, eastern U.S. west to IlUnois. 

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. Maryland: Sparrows Pt., Baltimore, July 3, 1932, 
J. W. Green; Edgewood, Harford Co., Nov. 3, 1918, H. Dietrich. Virginia: 
Pennington Gap, Lee Co., Hubbard and Schwarz. Florida: Apalachicola, 
Franklin Co., Taxodium distishum, W. F. Fiske. Georgia: Savannah, Chatham 
Co., March 9, 1940, Van Dyke. Mississippi: Lucedale, George Co., Feb. 27, 
1931, March 22, 1932, H. Dietrich; New Augusta, Perry Co., Feb. 12, 1931, H. 
Dietrich. Indiana: LaPorte Co., June 9, 1953, wheat, K. Pruess. Illinois: 
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; WiUow 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, Prison 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 \vill be found in some collections identified by Lane as 
"species 2." 

9. Melanotus verberans (LeConte) 

Figures 4m, 9i; Plate le 

Cratonychus verberans LeConte, 1853, p. 478. 

Melanotus verberans. — Cand^ze, 1860, p. 356. — Blatchley, 1910, p. 753. — Thomas, 
1941, p. 260. 

Male. — 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, vnth smaller and sparser pronotal punctures, genitalia 
with aedeagus nearly parallel sided and abruptly tapering to apex, 
paramere rather broad and with Uttle 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. — SLmUar 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. 

Types. — Lectotype of verberans: Male, labelled with pink circle 
"Md." (Ad!CZ). Only specimen now in type series, selected as lecto- 
type by authors in 1962; location of second type unknown. 

Distribution. — 'Maryland to Florida and west to South Dakota and 
Texas. 

Alabama: Locality?; January. Arkansas: Washington; May. Florida: 
Pinellas; March, May. Georgia: Ribb, Peach; April, June. Illinois: Mason; 
July. Indiana: Tippecanoe; April. Kansas: No data. Maryland: Baltimore; 
July. Mississippi: Forrest, George; January, April. Nebraska: Lancaster; 
May. New Jersey: Middlesex; April. New York: Essex, Niagara; May, 
November. North Carolina: Duplin, Moore, Orange, Wake, Wayne; April, 
May, October. Ohio: Athens; November. South Carolina: Bamberg, 
Oconee; May, Julj', December. South Dakota: Bon Homme; June. Texas: 
Bowie, Victoria; March. Virginia: Fairfax; April. 

10. Melanotus emissus (LeConte) 

Figures 4o, 9k; Plate Ih 

Cratonychus emissus LeConte, 1853, p. 478. 

Melanotus emissus. — Cand^ze, 1860, p. 372. — Thomas, 1941, p. 260. 

Male.' — 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 verherans), apical blade broader, and bearing four bristles. 

Elytron: 6.6 ±0.6 mm (5.9-8.4). 32 spec. 

Female. — 'Not positively identified (see discussion below). 

Type.— 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. 
Maryland: Baltimore; July. Nebraska: Lancaster; July. North Carolina: 
Bladen, New Hanover, Pender; May. South Dakota: Union, Yankton; June, 
July. Virginia: 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 verherans and the other a 
distinct form (fig. ^k). 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 verherans, but that 
cannot be answered at this time. 

11. Melanotus land Quate, new species 

Figures 5a, 9h; Plate 1/ 

Male. — 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 VA 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. Pennsylvania: Darby, Delaware Co., July 4. 
Iowa Ames, Story Co., June 6, 1931, J. F. Glawe;same, June 10, 1925. South 
Dakota: Brookings, June 22, 1943, H. C. Severin; Kimball, Brule Co., June 17, 
1947, Severin; Turner Co., June 15, 1929, Severin. Nebraska: Lincoln, June 27, 
1909, F. H. Shoemaker; Fairmont, FUlmore Co., June 17, 1914, G. W. Denning; 



NO. 3568 MELANOTUS — QUATE AND THOIVIPSON 31 

Bradshaw, York Co., July 9, 1917, Amnranlhus, E. J. Yates. Kansas: Douglas 
Co., J. C. Bridwell; Topeka, Popenoe. Arkansas: Hope, Hempstead Co., 
May 19, 1931, June 11, 1954, J. W. Green; "Vinita, Ind. T., June 7-8, '99, Wick- 
ham." Florida: 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. Melanotiis pilosus Blatchley 

Figures 5b, 91; Plate Ig 

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

Male. — 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, bm"sa 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). 

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

Distribution. — Central U.S. 

Arkansas: Washington; May. Illinois: Wabash; June. Indiana: Bar- 
tholomew; Tippecanoe; June. Iowa: Linn; June. Kansas: Riley, Sedgwick; 
June. Montana: Big Springs St. Pk.; June. Nebraska: Antelope, Chase, 
Custer, Keith, Lancaster, Lincoln, Scotts Bluflf; June. Oklahoma: Hayes; July. 

13. Melanotus opacicollis LeConte 

Figures 5d,e, 9j; Plate li 
Melanotus opacicollis LeConte, 1866, p. 390. — Blatchley, 1910, p. 751. 

Male. — 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. 

Female. — Similar to male. Bursa with 6 to 11 spines, much le^s 
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. 

Types.— Lectotype: Female, "I11./477" (MCZ). First specimen 
in type series selected as lectotype by authors in 1962. 
Distribution. — Indiana and ]\Iississippi, west to Kansas. 

Arkansas: Hempstead; May. Illinois: Greene, Marion, Rock Is.; May, 
June. Indiana: Owen; July. Iowa: no other data. Kansas: Linn, Mont- 
gomery, Riley; June. Louisiana; ? Co.; May. Mississippi: George, Greene; 
May. Montana: Boone, Callaway, Clinton, Jackson; May, July. 

14. Melanotus clandestinus (Erichson) 

Figure 5/ 

Cratonychus clandestinus Erichson, 1842, p. 112. — LeConte, 1853, p. 474. 
Melanotus clandestinus. — Candfeze, 1860, p. 343. — Thomas, 1941, p. 258.— 

Dietrich, 1945, p. 55.— Fattig, 1951, p. 20. 
Melanotus peninsularis Candfeze, 1889, p. 46. — Fattig, 1951, p. 22. [New 

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

Male. — ^Body color reddish bro^\^l, head and pronotum usually 
darker than appendages and elytra, covered with white (sometimes 
yello\\dsh) 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-lK segments, ratio of segments 2:3:4=2:2:7, 
segment 4 about Iji 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 sii:igle 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. 

Female. — Unknown to us. 

Types. — 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. 
(PU). 

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



NO. 3568 MELANOTUS — QUATE AND THOMPSON 33 

Alabama: Mobile; July. Florida: Highlands, Hillsborough, Lee, Levy, 
Palm Beach, Pinellas, Suwannee, Volusia; May, July. Mississippi: George, 
Jackson, Pearl River; June. New Jersey: Atlantic, Burlington, Cape May, 
Ocean; June, August. North Carolina: Columbus, Johnston, Moore; June, 
July. South Carolina: 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 

Figures 5g, 10a 

Melanotus ignobilis Melsheimer, 1846, p. 152. — Candeze, 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. 

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

Melanotus secretus. — Candeze, 1860, p. 344. — Thomas, 1941, p. 258. — Dcen 
and Cuthbert, 1955, p. 193.— Fattig, 1951, p. 20. 

Male. — 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. 

Female. — Antenna short of pronotal base by about 1 segment; 
segment 3 about IK times 2, but about K 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 

Types. — 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" (MCZ 2510). First specimen of type series selected as 
lectotype by authors in 1962. 

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

Alabama: Colbert, Mobile; May, June. Arkansas: Washington; June, July. 
Florida: Highlands, Alachua, Putnam, Volusia; April, May. Georgia: 
Charlton, Fulton; July. Illinois: Marshall, Putnam; June, July. Indiana: 
Clark; June. Louisiana: Madison; June, July. Mississippi: Greene, Jack- 
son, Perry; May, July. Montana: Phelps, St. Louis; June. Nebraska: 
Lancaster; July. New Jersey: Ocean; June, July. New York: Niagara, 
Tompkins; July. North Carolina: Wake; June. Pennsylvania: Dau- 
phin; June. South Carolina: Charleston, Oconee; March, July. South 
Dakota: Brookings; July. Tennessee: Morgan; June, August. Texas: 
McCuUoch; 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 apphed 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, hghter 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) 

Figures 5i, 10b; Plate Ij 

Ctenonychus depressus Melsheimer, 1846, p. 151. 

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

Melanotus depressus.— Candhze, 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. — Candfeze, 1860, p. 355.— Blatchley, 1910, p. 754. — 

Thomas, 1941, p. 260.— Dietrich, 1945, p. 57.— Fattig, 1951, p. 21. 



NO. 3668 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.] 

Male. — 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 -vvdth shallow, slitlike pit. Antenna exceeds 
tip of hind angle by 1 to 1,^ segments; reddish bro\vn, 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 Uttle 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: 8.7±0.6 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, peghke spines nearly throughout entire length, accessory 
gland originates near base, clavate, spermathecal duct branches near 
base. 

Elytron: 9.3 ±0.7 mm (8.2-10.2). 44 spec. 

Types. — 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 divarcarinus: Male, Posey Co., Ind., June 2, 1909, 
W. S. B. (PU). 

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

Distribution. — Connecticut to North Carohna, west to Nebraska 
and Texas. 

Arkansas: Benton, Hempstead, Pike, Washington; May, July. Connecti- 
cut: Fairfield, New Hampshire; June. Georgia: Catoosa, Fulton, Gwinnett, 
Lumpkin, Moran, White, Worth; May, July. Illinois: Champaign, Clay, 
Jackson, Johnson, Kane, Marion, McLean, Peoria, Pope, Putnam, Rock Island, 

221-532 — 67 i 



36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Scott, Union, Vermilion, Washington; April, July. Indiana: Spencer, Tippe- 
canoe; June. Iowa: Henry, Story; June, August. Kansas: Chautaugua, 
Cowley, Douglas, Miami, Montgomery, Riley; May, June. Kentucky: Jeffer- 
son; June. Louisiana: Madison; June. Maryland: Anne Arundel, Washing- 
ton; June, July. Mississippi: 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. New York: Dutchess, Essex, Seneca, Tompkins; May, 
July. North Carolina: Guilford; June. Ohio: Clifton, Delaware, Erie, 
Franklin, Greene, Hocking, Pickaway, Ross; May, July. Oklahoma: Payne, 
Sequoyah; July. Pennsylvania: Adams, Allegheny, Northampton, Phila- 
delphia; May, July. Tennessee: Sevier; June. Texas: Bexar, Brazos, Brew- 
ster, Dallas, Kerr, Madison, Val Verde; April, July. Virginia: Fairfax, Loudoun; 
May, July. Washington, D.C: June. West Virginia: 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 chne 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 
Indiana 


Arkansas, 
niinois 


Missouri 


Kansas 


Nebraska 


Texas 


c? 9 


C? 


9 


cP 


9 


& 


9. 3 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 crihulosus. 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 specunens 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 specmiens 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. 3568 MELANOTUS — QUATE AND THOMPSON 37 

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

17. Melanotus morosus Candeze 

Figures 5h, lOe 
Crafonychus longulus LeConte, 1853, p. 480. [Not LeConte, 1853, p. 473.] 
Melanotus morosus Candfeze, 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. 

]Male. — 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. 

Female. — Antenna extends to or beyond base of pronotum by 1 
segment; genitalia as figm'ed, 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. 

Types. — 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 3rd specimens not 
true types, labelled "Tex."; 4th specimen apparently infaustus. 

Holotype of debilis: Female, Marshall Co., Ind., June 26, 1904, 
W.S.B. (PU). 

Lectotype oi 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 Candeze 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. Florida: Manatee, Osceola, Pinellas; 
March. Georgia: Catoosa, Clarke, Cobb, Fulton, Hall, Rabun, Seminole, 
Thomas; March, June. Illinois: Champaign, Cook, Knox, Putnam, Scott, 
Washington; May, July (Nov.). Maryland: Anne Arundel, Baltimore, Har- 
ford, Prince Georges; June, October. Massachusetts: Barnstable, Middlesex, 
Nantucket, Worcester; May, July. Mississippi: George, Greene, Harrison, 
Lamar; April, May. Montana: Jefferson, Lawrence, St. Louis; April, July. 
Nebraska: Douglas; June. New Hampshire: Carroll; July. New Jersey: 
Atlantic, Camden, Ocean; May, August. New York: Orange, Suffolk, Tompkins, 
Washington; June, August. North Carolina: Buncombe, Duplin, Hyde, 
Moore, Sampson, Swain, Wake, Wayne; April, July. Ohio: Adams, Delaware, 
Fairfield, Franklin, Greene, Hocking, Scioto; June, August. Pennsylvania: 
Allegheny, Indiana, Monroe, Northampton, Philadelphia; April, July. South 
Carolina: Oconee; April. Tennessee: Carter, Sevier; June, September. 
Texas: Bastrop; April. Virginia: 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) 

Figures 5;, lOd 

Cratonychus cribulosus LeConte, 1853, p. 478. 

Melanotus cribulosus. — Candfeze, 1860, p. 357. — Hyslop, 1915, p. 17. — Thomas, 
1941, p. 260.— Severin, 1949, p. 17.— Srivastava, 1958, p. 87. 

Male. — 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 mth pit variable from ovoid to sUt shaped. Antenna exceeds 
tip of hind angle by )$ to iK segments, reddish brown, ratio of seg- 
ments 2:3:4 = 2.5:4:5, segment 4 about Ijz 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, mth 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 sUghtly shorter in com- 
parison to pronotum; genitaha as figured, bursa with few naiUike 
spines with large bases, accessory gland short and clavate, sperma- 
thecal duct branches near base. 

Elytron: 7.2-7.3 mm. 

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

Distribution.^ — Central U.S. 

Illinois: Champaign, Kane, Mason, McHenry, Warren; June, July. Kansas: 
Cowley, Douglas, Gove, Greenwood, Gray, Kingman; June, July. Nebraska: 
Lancaster, Saline; June, July. Oklahoma: Cleveland; April, May. South 
Dakota: 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) 

Figures 5n, lOi 

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- 

dfeze, 1860, p. 371.— Blatchley, 1910, p. 745.— Thomas, 1941, p. 259. 
Cratonychus glandicolor. — LeConte, 1853, p. 477. 

Male, — 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 3 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. 

Female. — 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, biu-sa 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. 

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

Illinois: Kankakee, Vermilion; June, July. Indiana: Clark; June. Mart- 
land: Baltimore; June. Montana: Montgomery; May. New Jersey: 
Warren; June. North Carolina: Buncombe; June. Pennsylvania: Adams; 
May. Tennessee: Sevier; June. Virginia: Norfolk; May. Ontario: 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; Plate Ifc 
Cratonychus Sagittarius LeConte, 1853, p. 480. 



NO. 3568 MELANOTTJS — QUATE AND THOMPSON 41 

Melanotus Sagittarius. — Candfeze, 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.] 

Male. — 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 angidate 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 iKo 
times height. Mandible mth deep, teardrop-shaped pit. Antenna 
exceeds tip of hind angle by 2 to 2K 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 = l. 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 cm*vature, 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. 

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

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

Types. — •Lectotype of Sagittarius: Male, labelled with white circle 
"Eastern States & Canada" (MCZ). First specimen in type series 
selected as lectotype by authors in 1962. Second specimen labelled 
wiih 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. 

Florida: Pinellas; February, April. Illinois: Putnam; July. Indiana: 
Marion; May. Maryland: Baltimore; July. Massachusetts: Hampshire, 
Middlesex; June, July. Mississippi: George; April. New Jersey: Middlesex; 
July. New York: Dutchess, Rockland, Tompkins, Westchester; June, July. 
Ohio: Athens; June. Pennsylvania: Allegheny, Bradford, Monroe; May, 
July. Virginia: Page; July. West Virginia: Hardy; July. 



42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

21. Melanotus hyslopi Van Zwaluwenburg 

Figures 51, 10/ 

Melanotus hyslopi Van Zwaluwenburg, 1921, p. 210. — Thomas, 1941, p. 261. — 
Dietrich, 1945, p. 69.— Fattig, 1951, p. 22. 

Male. — 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. 

Female. — Antenna exceeds tip of hind angle by 1^2-2 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. 

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

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

Connecticut: Litchfield; June. Illinois: Champaign, McHenry; July, Sep- 
tember. Maryland: Anne Arundel, Baltimore, Frederick; June, July. Mas- 
sachusetts: Barnstable, Bristol, Essex, Hampshire, Middlesex; June, August. 
Michigan: Oceana; August. Mississippi: Forrest; April. New Hampshire: 
Carroll; August. New Jersey: Atlantic, Bergen, Camden, Ocean; May, July. 
New York: Erie, Essex, Niagara, Rockland, Suffolk, Sullivan, Tompkins; May, 
September. North Carolina: Buncombe, Macon; July. Ohio: Delaware, 
Fairfield, Hocking, Summit; May, July. Pennsylvania: Dauphin, Monroe, 
Northampton, Philadelphia; May, July. Tennessee: Sevier; June, July. 
Virginia: 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 punctiu-es, longer antennae, larger paranten- 
nal fovea, and a more angulate frontal margin. 

22. Melanotus prasinus Blatchley 

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

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

Head: Front vnih pair of shallow anterolateral transverse depres- 
sions, margin dark reddish bro^vn, sometimes strongly protuberant 
above clypeus and turned up in front, fairly thin, parantennal fovea 
moderately small, but well defined, arc 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 vAdth/length= 1.0-1 .07; 
punctures at center usually separated by distance equal to about 
tmce own diameter, shghtly 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. GenitaHa as figured; paramere without apical blade. 

Elytron: 6.8 mm (6.2-7.3). 14 spec. 

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

Elytron: 7.5 mm. 1 spec. 

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

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

Connecticitt: Windham; August. Illinois: Champaign; July. Maryland: 
Plummers Isl.; April, June. Massachusetts: Barnstable, Middlesex; July. 
Montana: Carter; June. New Hampshire: HiUsboro; no date. New Jersey: 
Bergen, Cape May; June, July. New York: Kings; July. North Carolina: 
Buncombe; July. Ohio: 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 

Figures 5k, lOj 
Melanotus piceatus Blatchley, 1927, p. 141; 1930, p. 48. — Fattig, 1951, p. 22. 

Male. — 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 
IK 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 IK 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 mthout apical blade. 

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

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

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

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

Distribution. — Southeastern U.S. 

Alabama: Colbert, Mobile; May, June. Florida: Brevard, Charlotte, 
Highlands, Hillsborough, Pinellas, Volusia; February, May. Georgia: Chatham; 
March. South Carolina: 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 

Figure 6fir 

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

Male. — Body color reddish broA\Ti, covered with whitish vestiture, 
evenly distributed over body and longer than usual. 

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

Pronotum flattened dorsally, \^'ider 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 mth last visible sternite usually inflated to give tip 
enlarged appearance. Genitalia as figured; paramere with apical 
blade. 

Elytron: 7.5-9.7 mm. 

Female. — Unknown to us. 

Type. — Holotype of difficilis: Male, Posey Co., Ind., July 8, 1903, 
W.S.B. (PU). 

Distribution. — Central U.S. 

Kansas: Douglas; June. Illinois: Alexander; June. Mississippi: George; 
May. Oklahoma: Delaware; June. Texas: Austin, Dallas, Navarro; May. 

25. Melanotus macer (LeConte) 

Figures 6e,f, 101 

Cralonychus macer LeConte, 1853, p. 473. 

Melanotus macer. — Candfeze, 1860, p. 339. — Blatchley, 1910, p. 752. — Dietrich, 
1945, p. 54.— Fattig, 1951, p. 20. 

Male. — 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 usuaUy 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, usuaUy extending cephalad of base of hind angles, 
Genitaha as figured; paramere with apical blade. 

Elytron: 7.2-7.6 mm. 

Female. — Unknown to us. 

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

Distribution. — North Carolina west to Kansas. 

Iowa: Story; June. Kansas: Cherokee, Riley; June, July. Mississippi: 
George; May. New York: Nassau; July. North Carolina: Moore, Wake; 
May. 

The two species difficilis and macer can be separated from other 
North American Melanotics 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) 

Fiqitres 66, llc,d 
Ctenonychus testaceus Melsheimer, 1846, p. 151. 
Melanotus testaceus. — Thomas, 1941, p. 258. 
Cratonychus dubius LeConte, 1853, p. 479. [New synonymy.] 
Melanotus dubius.— Candhze, 1860, p. 372.— Thomas, 1941, p. 260.— Fattig, 1951, 

p. 21. 
Melanotus angustatus of authors, not Erichson, 1842. 

Male. — 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. 3568 MELANOTUS — QUATE AND THOMPSON 47 

and well defined, semicircular; nasale protuberant, width 1 to IK times 
height. Mandible with deep, oval pit. Antenna exceeds tip of hind 
angle by 1 to 1 K 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, plaquehke 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. 

Types. — 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) 
Usting of N.Y., but nonetheless is regarded as the type. 

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

Arkansas: Hempstead, Washington; April, August. Florida: Pinellas; 
April. Georgia: Fulton; June. Illinois: Alexander, Lawrence; April, July. 
Montana: Callaway; May. New Jersey: Ocean; June. New York: 
Suffolk, Tompkins; June, August. North Carolina: Wayne; April. Ohio: 
Delaware, Greene, Hocking, Scioto; May, June. Texas: 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 apphed 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. — Candeze, 1860, p. 348. — Blatchley, 1910, p. 752. — 

Thomas, 1941, p. 259. — Dietrich, 1945, p. 55. — Jewett, 1946, p. 9.— Fattig, 

1951, p. 20. 

Male. — 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 1^ 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 figiu-ed; paramere without apical 
blade. 

Elytron: 6.1 ±0.5 mm (5.2-7.3). 35 spec. 

Female. — ^Antenna exceeds tip of hind angle by about one-half 
segment; internal genitaha as figured, bursa not elongate or enlarged 
at base of accessory gland and mth more than 50 peghke spines. 

Elytron: 5.1-6.9 mm. 6 spec. 

Types. — 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. 

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

Arkansas: Garland, Washington; June. Connecticut: Litchfield, New 
Haven; June, July. Illinois: Champaign, Knox, McHenry, Putnam, Rock 
Island; May, July. Massachusetts: Berkshire, Hampshire, Middlesex, Nor- 
folk; June, July. Minnesota: Hennepin; July. Mississippi: George; June. 
New Hampshire: Carroll; August. New Jersey: Bergen, Gloucester; June. 
New York: Erie, Essex, Oswego, Rockland, Suffolk, Tompkins, Westchester; 
April, September. North Carolina: 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) 
Figures 6d, llo; Plate Im 

Elater tenax Say, 1839, p. 185. 
Cratonychiis tenax. — LeConte, 1853, p. 479. 

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

Male. — 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 IK 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, subparaUel; 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. 

Female. — 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. 

Types. — Types of tenax: "Mass." Specimens lost. 

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

Georgia: Houston; June. Illinois: Greene; June. Maryland: Prince 
Georges; June. Massachusetts: Middlesex, Nantucket; June. New York: 
Albany; June. North Carolina: Craven; May. South Carolina: 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 tenax. 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 sperm athecal duct of tenax, 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) 

Figures 6/, lOk 

Elater pertinax Say, 1839, p. 185, 

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

Melanotus pertinax. — Candfeze, 1860, p. 359. — Blatchley, 1910, p. 755. — Thomas, 

1941, p. 260.— Dietrich, 1945, p. 58.— Severin, 1949, p. 17.— Fattig, 1951, 

p. 21. 

Male. — 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 cljrpeus; parantennal 
fovea small but moderately deep circular or semicircular, foveal 
tragus seldom developed; nasale flat, width about 2 tunes 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 iKo 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. 

Female. — 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. 

Types. — Types of pertinax: "Mass." Specimens lost. 

Distribution. — Maine to North CaroHna, west to Wisconsin and 
Illinois. 

Connecticut: Litchfield, New Haven; May, July. Georgia: Clarke; May. 
Illinois: Champaign, Knox, McHenry, McLean; May, July. Maine: York; 
July. Massachusetts: 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. North Carolina: 
Buncombe; June. Pennsylvania: Lycoming; July. South Carolina: 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. 



NO. 3568 MELANOTUS QUATE AND THOMPSON 51 

30. Melanotus infaustus (LeConte) 

Figures Gi, lli,j; Plate lo 

Cratonychus infaustus LeConte, 1853, p. 478. 

Melanotus infaustus. — Candfeze, 1860, p. 357. — Thomas, 1941, p. 260. — Severin, 
1949, p. 17.— Fattig, 1951, p. 21. 

Male. — Body color reddish brown, covered with whitish vestitm-e 
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 w^ell 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 brow^n, ratio of segments 
2:3:4 = 3:6:9, segment 4 about 1}^ times as long as maximum u^dth; 
erect male hairs short and only moderately dense. 

Pronotum about as long as wide, punctm-es at center usually 
separated by distance equal to or less than owm 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. 

Female. — 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. 

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

Indiana: Tippecanoe; June. Illinois: Whiteside; May. Kansas: Douglas, 
Reno; June. South Dakota: 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 tenax, 
trapezoideus , and prasinus in the Midwest and South or with lanceatus 
in Texas. The slender thorax is helpfid 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 

Figures 6h, llg; Plate In 

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

Male. — Body color reddish brown, covered with white or yellowish 
vestitiire, 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, arc 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 Jissilis group). Antenna exceeds tip of 
hind angle by 1 to IK 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. 

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

Elytron: 5.9 mm. 

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

Distribution. — Southern U.S. 

Arkansas: No data. Florida: Highlands, Marion, Pinellas; February, April. 
North Carolina: 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) 

Figures 6j, lie,/ 

Elater americanus Herbst, 1806, p. 74. 

Cr atony chus americanus. — Erichson, 1842, p. 114. — LeConte, 1853, p. 479. 



NO. 3568 MELANOTUS — QUATE AND THOMPSON 53 

Melanotus americanus. — Candfeze, 1860, p. 361. — Blatchlcy, 1!)10, p. 747. — 

Fattig, 1951, p. 21. 
Cralonychus anguslaius Erichson, 1842, p. 113. — LeConte, 1853, p. 475. [New 

synonymy.] 
Melanotus angustaliis. — Candeze, 1860, p. 345. — Blatcliley, 1910, p. 753. — Thomas, 

1941, p. 258. 

Male. — Body color reddish bro^^^l, covered ^\^th white vestiture 
evenly distributed over body. Small species. 

Head: Front \\dth pair of shallow, anterolateral, or single, curved 
anterior depressions, margm 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, w^ell developed. 
Mandible with deep, teardrop-shaped pit. Antenna exceeds tip of 
hind angle by 1^2-2 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 apicaUy, but without apical 
blade. 

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

Female. — 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, 
diverticidum branching from spermathecal duct near center of duct. 

Elytron: 5.7i0.3 mm (4.8-6.7). 93 spec. 

Types. — 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. 

Distribution. — Eastern U.S. west to Indiana and Illinois. 

Connecticut: Hartford, Litchfield; June. Delaware: Sussex; I\Iay. 
Georgia: DeKalb, Fulton, Rabun; INIay, June. Illinois: McHenry, Putnam, 
Rock Is.; June, July. Indiana: Marion; July. Maryland: Anne Arundel, 
Baltimore, Frederick, Plummers Is.; June, July. Massachusetts: Barn- 
stable, Hampshire, JNIiddlesex; June, September. Nebraska: Cass; May, 
July. New Jersey: Bei'gen, Middlesex, Morris, Union; June, July. New 
York: Putnam, Suffolk, Westchester; April, August. North Carolina: Bun- 
combe, Cherokee, Cumberland, Moore, Montgomery, Pender, Pickens, Swain, 
Wake; April, July. Ohio: Delaware, Fairfield, Franklin, Greene, Hocking; 
May, June. Pennsylvania: Allegheny, Dauphin, Northampton, Perry; June, 
221-532— 67-— 5 



54 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

July. Rhode Island: Washington; July. South Carolina: Oconee; March. 
West Virginia: Greenbrier; July. 

Melanotus americanus is one of the smaller species of North American 
Melanotus. This featm:"e, with the small thu"d antennal segment and 
high rounded pronotiim bearing small, scattered punctures, makes 
the species more readUy 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 6 A;, 11 /i 

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 Candfeze, 1860. New synonymy.] 

Male. — ^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 3K 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 cm-vature; 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. 

Female. — ^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. 

Types. — 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 55 

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

Distribution. — New Jersey west to Kansas. 

Illinois: Rock Is.; no date. Indiana: Allen; June. Kansas: Douglas; no 
date. Maryland: Baltimore, Plummers Is.; April. New Jersey: Warren; 
June. Ohio: 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. 

Blatcliley apparently believed the male and female belonged to two 
different species. He first named the male longicornis, but this name 
is preoccupied by longicornis Candeze. Leng observed the homonymy 
and renamed Blatchley's species blatchleyi. 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 
cribriventris. 

34. Melanotus obscuratus Blatchley 

Figures 7b, 111 
Melanotus obscuratus Blatchley, 1927, p. 141; 1930, p. 45. 

Male. — ^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 figm-ed; paramere with well-defined apical blade. 

Elytron: 6.0-6.4 mm. 2 spec. 

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

Elytron: 7.4 mm. 1 spec. 

Type.— Holotype: Male, Ormoiid, Fla., April 13, 1913, W. S. B. 
(PU). 



56 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Distribution. — Southeastern U.S. 

North Carolina: Brunswick; June. Florida: Volusia; April. 

Three females in the type series belong to morosus as sliown 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 ohscuratus and agreeing with that specimen in external characters. 

35. Melanotus beameri Quate, new species 

Figure 6 m 

Male. — Body color light reddish bro^vn, 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, slitlike pit. Antenna 
reddish brown, exceeds tip of hind angle b}^ 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 mth little or no anterior cur- 
vature and nearly parallel but a httle 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. 

Female . — 'Unkn o wn . 

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 Jet. 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 follo\ving 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 Kniill 

Figures 7a, 12a 
Melanotus concisus Knull, 1959, p. 281. 

Male (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- 
taha 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. 

Type.— Holotype : Male, Ai-tesia (Eddy Co.), N. Mex. (Knull 
Collection). 

Distribution. — 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 genitaha. Positive identification of the female cannot 
be made, however, until associated with male in the field. 

37. Melanotus lanceatus Quate, new species 

Figures 7c, 12b 

Male. — 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 b}^ 
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. 

Female. — 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. 

Distribution. — 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, 
KnuU; 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. S. 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 hght, B. J. 
Adelson. Hidalgo Co.: Edinbiu-g, 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.: CotuUa, 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 bm-sal 
spines, but the origin of the accessory gland is in a different position 
in each. Externally, the smaller pronotal punctm'es 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 shoidd 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 
greater 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 KnuU 

Figure 7d 
Melanotus hamatus KnuU, 1959, p. 280. 

Male. — 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 ^nthout apical blade. 

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

Female.- — ^Unknown to us. 

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

Distribution. — -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) 

Cratonychits longidits LeConte, 1853, p. 473. 
Melanotus longulus. — Candfeze, 1860, p. 339. 

Male.— 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 

angiUate 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)2 segments, ratio of segments 2:3:4=3:4:7, flagellum usually 
strongly serrate, segment 4 about IK 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. 

Female. — ^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 biu-sa. 

39. Melanotus longulus longulus (LeConte) 

Figures 7e, 12c,d 

Cratonychus longulus LeConte, 1853, p. 473. 

Melanotus longulus. — Candeze, 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 ciu-vatm-e, 
usually slightly convergent posteriorly. 

Male genitalia as figiu-ed; 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 figiu-ed, accessory gland longer than 
biu-sa 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. 

Types.— 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 ISIts., 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 Alts., July 2, 1954 
July 3, 1955. Pinal Co.: Magna, Aug. 18, 1921; Oracle, July 7, 1950; Tucsonj 
August 1935. Santa Cruz Co.: Mt. Washington, Nogales, 1829 m, July 13, 1919; 
Patagonia, July 1936. Yavapai Co.: Congress Jctn., July; Prescott, July 1, 1919. 
Yuma Co.: Yuma, March 7, 1879. California. 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; Clare mont, Apr. 19, 1926 
May 13, 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; Tanbarlc Flat, June 20-July 26, 1952. Orange Co.: Newport 
Bay, May 14, 1940. Riverside Co.: Andi'eas Cnyn., Apr. 3, 1927; Coachella, 
May 19, 1927; Idyllwild, June 7, 1940; Indio, June 2, 1918; Alurray 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. Nevada. 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. Texas. Brewster Co.: Alpine, 
July 11, 1938. Utah. 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. Baja California. 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 
/. longulus. Western populations, particularly in and around Los 
Angeles Co., Calif., are colored like I. oregonensis with a black or 
nearlj^ 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. Genitaha 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) 

Figures 7/, 12e,/ 

Cratonychns oregonensis LeConte, 1853, p. 480. 

Melanotus oregonensis. — Candfeze, 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. — Fall, 1934, p. 24. 



62 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Larger than I. longulus; frontal margin more broadly flattened than 
in longulus. Usually black with lighter colored antennae. Antenna 
flagellum often strongly serrate. Pronotal piinctm-es 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 figiu-ed; apical blade weakly developed, gently 
rounded at base. 

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

Male elytron: 9.0 ±0.7 mm (7.3-10.3). 50 spec. 

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

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

Lectotype of variolatus: Male, labelled with gold circle (Calif.) 
(MCZ 2522). Thu-d specimen of type series selected as lectotype 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. Alay 8, 
1910 (CAS 3132). 

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

California. Alameda Co.: Berkeley, May 9, 1940, June 9, 1949; Castle 
Rock Pk., May 25, 1932; ]\Ioraga, 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, 1931; 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; MUl 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.: Half moon 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. Idaho. 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. Nevada. No 
other data. Oregon. 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, 
INLay 14, 1954; Talent, May 4, 1954. Lake Co.: Lakeview, May 27, 1957. 
Yamhill Co.: June 1913, May 1934. Utah. Cache Co.: June 26, 1954; Lewis- 
ton, JNIay; Bear River, Logan, 1925. Juab Co.: Eureka, July 2 (with /. 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. Washington. 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 Columbia. 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 genitaUc 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 improven, however, and we have seen few specimens from the 
borders of the two forms' area. In oiu- 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 thu'd 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 SieiTa 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 chne, it is abrupt and not 
gradual as outlined by Van Dyke. On the average, specunens 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 hkely a phenotypic 
expression of a more vigorous clunate, shorter simimer 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 synonjnnized with longulus by Van 
Dyke (1932) on the grounds that it was based on individual variants 
and in this we concur. 

41. Melanotus gradatiis LeConte 

Figure 7g; Plate Ip 
Melanotus gradatus LeConte, 1866, p. 390.-Blatchley, 1910, p. 753.-Thomas, 
1941, p. 260. 
Male.— 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; margm 
not strongly protuberant above nasale, evenly rounded or flattened 
in front when viewed from above; parantennal fovea moderately large, 
deep, semicircidar, foveal tragus smaU; nasale raised slightly, width 
from' 1% to IK 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 lYi times as 
long as maxunum width; erect male hairs very short, fine, rather 
sparse, and easily overlooked. 

Pronotiun 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 
usuaUy 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 

Female.^ — 'Antenna extends just to tip of hind angle or short of tip 
by one-half segment; internal genitalia as figured, biu-sa 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. 

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

Distribution.— Maryland west to Kansas and Arkansas. 

Arkansas: Lawrence, Washington; April, June. Illinois: Gallatin, Pope, 
Pulaski, St. Clair; May, June. Indiana: Clark; June. Kansas: Douglas; 
June. Ohio: 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. 

Cr atony chus insipiens. — LeConte, 1853, p. 480. 

Melanotus insipiens. — Candeze, 18G0, p. 361. — Fattig, 1951, p. 21. 

Cratonychus tenellus Erichson, 1842, p. 114. — LeConte, 1853, p. 480. 

Melanotus tenellus. — Candeze, 1860, p. 362. — Fattig, 1951, p. 22. 

Male. — 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, semicircidar; 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 spmes, 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: 3.6±0.2 mm (3.0-4.3). 50 spec. 
Types. — 'Type of insipiens: Specimens lost. 

Lectotype of tenellus: Female, "Amer. Sept." (ZM). Lectotype 
selected by Quate in 1960. 

Distribution.^ — New Jersey to Florida, west to Texas. 

Alabama: Mobile; June. Arkansas: Hempstead; June. Florida: Columbia, 
Pinellas, Osceola, Volusia, Wakulla; April, May. Georgia: Charlton, Houston; 
May. New Jersey: Burlington, July. North Carolina: 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 punctm'es are much smaller and 
sparser. M. 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) 

Figures Ik, 12h, i 

Cratonychus Leonardi LeConte, 1853, p. 475. 

Melanotus leonardi. — Candfeze, 1860, p. 349. — Thomas, 1941, p. 259.— Dietrich, 
1945, p. 56.— Fattig, 1951, p. 20. 

Male. — 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 easUy. 

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 \N'ith 
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. 

Female. — 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. 

Types. — 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. 

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

Connecticut: Litchfield; June. Indiana: Marion; June. Maine: Hancock, 
Oxford, Washington; July. Massachusetts: Suffolk, Worcester; June. New 
Hampshire: Cheshire, Strafford; no dates. New York: Essex, Jefferson; April, 
June. North Carolina: Buncombe; June. Texas: Bexar; June. Vermont: 
No data. Ontario: Mustota Dist., OrrvUle; June. Quebec: Duparquet, Mt. 
St. Hilaire, Perkins Mills; June, July. 

44. Melanotus taenicollis (LeConte) 

Figures 71, 12j 

Cratonychiis taenicollis LeConte, 1853, p. 475. 

Melanotus taenicollis: Candfeze, 1860, p. 348. — Thomas 1941, p. 259. — Dietrich, 
1945, p. 56. 

Male. — Black and orange or reddish brown, body largely black 
Avith 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 mth anterior depression; margin scarcely protuberant 
in front of nasale, coining to point in center when view^ed 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. 

Female. — 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: 6.7 ±0.4 mm (6.1-7.5). 16 spec. 

Types. — Lectotype of taenicollis: Male, labelled with pink circle 
"Middle States; Pa." (MCZ 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. 
Distribution. — Eastern 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 leonardi. 

The crihricollis Group 

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

45. Melanotus crihricollis Candeze 

Figures 7j, \2l,m; Plate IZ 
Melanotus crihricollis Candeze, 1860, p. 358. 

Male. — 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 semicircidar. 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 rectangidar, 
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 subparaUel, each v/ith 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. 

Female. — Internal genitalia as figured, accessory gland large and 
spherical, bursa with large, dense patch of spil^elike spines. 

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

Type. — ^Holotype of crihricollis: Female, "Amer. Bor." (BMN). 
Only specimen in type series. 



NO. 3568 MELANOTUS — QUATE AND THOMPSON 69 

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

Arizona: Cochise, Maricopa, Pima, Pinal; April, September. California: 
San Diego; April. Baja California: 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. cribricoUis and chiricahuae are readily separable from other North 
American Melanotus by the large pronotal punctures, shallow mandib- 
idar 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. cribricoUis 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 7i, I2k 
Melanotus chiricahuae Knull, 1962, p. 34. 

Male. — Large, dark reddish-bro^vn 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 K 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, pimctures 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. 

Female. — 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. 

Distribution. — Southwestern U.S. 

Arizona: Cochise, Santa Cruz; June-August. New Mexico: Hidalgo; August. 
221-532—67 6 



70 PROCEEDESTGS OF THE NATIONAL MUSEUM vol. 121 

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

Species Incertae 
Melanotus abdominalis (Erichson) 

Cratonychus abdominalis Erichson, 1842, p. 104. — LeConte, 1853, p. 481. 
Melanotus abdominalis.' — Candfeze, 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 margm, fine dense pronotal pimctures, and a long, 
slender, heavily spmed female bursa. It is imlike any species which 
we have encountered in North America and is imfamiliar 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 (=Jissilis) or 
communis, but the types apparently are lost, and we are unable to 
apply the name to a known species. 

Melanotus despectus Candeze 

Melanotus despectus Candfeze, 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 sho^vn to 
be types, despectus would fall as a synonym of ignobilis. 

Melanotus effetus Candeze 

Melanotus effetus Candfeze, 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. 



Literature Cited 

Arnett, R. H., Jr. 

1955. Supplement and corrections to J. A. Hyalop's genotypes of the 

elaterid beetles of the world. Proc. U.S. Nat. Mus., vol. 103, no. 
3336, pp. 599-619. 
Becker, E. C. 

1956. Revision of the Nearctic species of Agriotes (Coleoptera: Elateridae). 

Canadian Ent., vol. 88, suppl. 1, pp. 1-101. 

BiNAGHI, G. 

1939. I Melanotini della fauna italiana: Spheniscocomus Schw. e Melanotus 
Eschs. (Coleoptera: Elateridae). Mem. Soc. Ent. Italiana, vol. 17, 
pp. 205-239 (1938). 
Blackwelder, R. E. 

1939. Fourth supplement to Leng Catalogue of Coleoptera of America, 
north of Mexico. [Elateridae: pp. 39-42.] 
Blackwelder, R. E. and R. M. 

1948. Fifth supplement to Leng Catalogue of Coleoptera of America, north 
of Mexico. [Elateridae: p. 19.] 
Blanchard, C. E. 

1845. Histoire des insectes, vol. 2. 
Blatchley, W. S. 

1910. An illustrated descriptive catalogue of the Coleoptera or beetles 
(exclusive of the Rhynchophora) known to occur in Indiana. 
[Elateridae: pp. 699-773.] 
1920. Canadian Ent. News, vol. 52, pp. 45, 46. 
1927. Some new species of Coleoptera from Indiana and Florida. Ent. 

News, vol. 38, pp. 139-144. 
1930. Blatchleyana, 
Brooks, A. R. 

1960. Canadian Ent., suppl., vol. 20, p. 40. 
CANDi:zE, E. 

1860. Monographic de Elateridae, 3. Mem. Soc. Sci. Liege, vol. 15, 512 pp. 
1889. Elaterides nouveaux. Ann. Soc. Ent. Belgique, vol. 33, pp. 67-123. 
CoMSTOCK, J. H., and Shingerland, ? 

1891. Cornell Agric. Exp. Sta. Bull., no. 33, p. 262. 
Deen, O. T., and Cuthbert, F. P., Jr. 

1955. The distribution and relative abundance of wireworms in potato- 
growing areas of the southeastern states. Journ. Econ. Ent., vol. 
48, pp. 191-193. 
Dejean, p. S. 

1833. Catalogue des Coleopteres de la coUecion . . ., ed. 2, 176 pp. 
Dietrich, Henry 

1945. The Elateridae of New York State. Cornell Univ. Agric. Exp. Sta. 
Mem. 269, 79 pp. 
Emmons, Ebenezer 

1854. Insects of New York. Vol. 5 of pt. 5 (Agriculture) in Natural history 
of New York. 

71 



72 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Erichson, W. F. 

1842. Ueber die Elateren mit Kammformig gezahnten Krallen. Germar 
Zeitschr., vol. 3, pp. 88-129 [1841?]. 

ESCHSCHOLTZ, J. F. 

1829. In T. Thon, Entomologisches Archiv., vol. 2, p. 32. 
Fall, H. C. 

1934. Journ. New York Ent. Soc, vol. 42, p. 24. 
Fattig, p. W. 

1951. The Elateridae or click beetles of Georgia. Emory Univ. Mus. 

BuU., no. 10, 25 pp. 
Fenton 

1926. Journ. Econ. Ent., vol. 19, p. 502. 
Gtllenhal, L. 

1817. In Schoenherr, Synonymia insectorum, oder: Versuch einer Syno- 
nymie aller bisher bekannten Insecten . . ., Appendix, vol. 3, 
p. 138. 
Herbst, J. F. M. 

1806. Natursystem aller bekannten in- und auslandischen Insecten, . . ., 
vol. 10, 285 pp. 
Horn, George H. 

1874a. Trans. Amer. Phil. Soc, n. s., vol. 10, p. 473. 
1874b. Trans. Amer. Ent. Soc, vol. 5, p. 23. 

1879. [Communication on the Elateridae.] Trans. Amer. Ent. Soc, 
vol. 7, pp. xiv-xvi. 
Hyslop, J. A. 

1915. U.S. Dept. Agric Bur. Ent. Bull., no. 156, p. 17. 

1916. U.S. Dept. Agric. Farmers' BuU., vol. 725, p. 5. 

1921. Genotypes of the elaterid beetles of the world. Proc U.S. Nat 
Mus., vol. 58, pp. 621-680. 
Jeuniaux, Charles 

1955. Sur les Elateroides palearctiques. Bull. Ann. Soc. Roy. Ent. Belgi- 
que, vol. 91, pp. 230-237. 
Jewett, H. H. 

1946. Identification of some larval Elateridae found in Kentucky. Ken 
tucky Agric. Exp. Sta. Bull., no. 489, 40 pp. 
Kirby, W. 

1837. Insects. Pt. 4 in Richardson, Fauna Boreali-Americana, xxxix + 325 
pp., 8 col. pis. 
Knull, J. N. 

1959. Two new species of Melanotus (Coleoptera: Elateridae). Ohio 

Journ. Sci., vol. 59, no. 5, pp. 280-281. 
1962. A new Melanotus from the Chiricahua Mountains of Arizona (Cole- 
optera: Elateridae). Ohio Journ. Sci., vol. 62, no. 1, p. 34. 
Lane, M. C. 

1952. List of Elateridae of British Columbia. Proc Ent. Soc. British 

Columbia, vol. 48, pp. 65-67 [1951]. 
LeConte, J. L. 

1853. Revision of the Elateridae of the United States. Trans. Amer. 

Phil. Soc, n. s., vol. 10, pp. 405-508. 
1866. Additions to the coleopterous fauna of the United States, 1. Proc. 

Acad. Nat. Sci. Philadelphia, vol. 18, pp. 361-394. 



NO. 3568 MELANOTUS — QUATE AND THOMPSON 73 

Leng, C. W. 

1918. Notes on some changes in the Hst of Coleoptera. Journ. New York 

Ent. Soc, vol. 26, pp. 201-211. 
1920. Catalogue of the Coleoptera of America, north of Mexico, x + 470 pp. 
[Elateridae: pp. 166-175.] 
Leng, C. W., and Mutchler, A. J. 

1927. Supplement to Catalogue of Coleoptera of America, north of Mexico. 

[Elateridae: p. 29.] 
1933. Second and third supplements to Catalogue of Coleoptera of America, 
north of Mexico. [Elateridae: pp. 27, 83-85.] 
McClure, H. E. 

1933. Ent. News, vol. 44, p. 145. 
Melsheimer, F. E. 

1846. Descriptions of a new species of Coleoptera of the United States. 
Proc. Acad. Nat. Sci. Philadelphia, vol. 2, pp. 134-160. 
Paykull, G. 

1800. Insecta. Vol. 3 in Fauna svecica. 
PuTZEYs, Jules 

1908. Ann. Soc. Ent. Belgique, vol. 52, p. 294. 
Redtenbacfer, L. 

1849. Fauna Austriaca, xxviii + 883 pp. 
Say, Thomas 

1823. Descriptions of coleopterous insects collected in the late expedition 
to the Rocky Mountains, performed by order of Mr. Calhoun, 
Secretary of War, under the command of Major Long. Journ. 
Acad. Nat. Sci. Philadelphia, vol. 3, pp. 139-216. 
1825. Descriptions of new American species of the genera Buprestis, 
Trachys and Elater. Ann. Lye. Nat. Hist. New York, vol. 1, 
pp. 249-268. 
1839. Descriptions of new North American insects and observations on 
some already described. Trans. Amer. Phil. Soc, n. s., vol. 6, 
pp. 155-190 [1836]. 

SCHENKLING, S. 

1927. Coleopterorum catalogus, Elateridae, 2. No. 88 in Junk and 
Schenkling, pp. 265-636. 

SCHWARZ, 0. 

1892. Revision der palaarktischen Arten der Elateriden-Gattung Melanotus, 
Esch. Deutsche Ent. Zeitschr., pp. 145-164. 
Severin, H. C. 

1949. The wireworms (Elateridae) of South Dakota. Tech. Bull. South 
Dakota Agric. Exp. Sta., no. 8, 18 pp. 
Snodgrass, R. E. 

1957. A revised interpretation of the external reproductive organs of male 

insects. Smithsonian Misc. Coll., vol. 135, no. 6, pp. 1-60. 
Stephens, J. F. 

1830. Mandibuta, vol. 3. Illustrations of British Entomology. 
Srivastava 

1958. Annals Zool., vol. 2, p. 87. 
Stone, M. W., and Rowland, A. F. 

1944. Life history of the wireworm Melanotus longulus (Lee.) in southern 
California. U.S. Dept. Agric. Tech. BuU. 858, 30 pp. 



74 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Thomas, C. A. 

1941. The Elateridae of Pennsylvania. Journ. New York Ent. Soc, vol. 
49, pp. 233-263. 
Van Dyke, E. C. 

1932. Miscellaneous studies in the Elateridae and related families of 
Coleoptera. Proc. California Acad. Sci., ser. 4, vol. 20, no. 9, 
pp. 291-465. 
Van Zwaluwenburg, R. H. 

1921. Melanotus hyslopi, new species (Coleoptera). Proc. Ent. Soc. 

Washington, vol. 23, pp. 210-211. 

1922. External anatomy of the elaterid genus Melanotus (Coleoptera), 

with remarks on the taxonomic value of certain characters. Proc. 
Ent. Soc. Washingt,on, vol. 24, pp. 12-29. 
Weber, Friedrich 

1801. Observationeb entomologicae, continentes novorura quae condidit 
generum characteres, . . . xii+116 pp. 
Westwood, J. O. 

1840. An introduction to the modern classification of insects . . ., vol. 2. 



MELANOTUS — QUATE AND THOMPSON 



75 




Figure 4.-Male genitalia of Melanotus species: a-d, similis; e, spadix; f-h, decumanus; i, 
castanipes; ], communis; k, indistindus; I, dielrichi; m, verberans; n, miscdlus; o, omissus. 
(bcale lines=0.5 mm.) 



76 



PROCEEDINGS OF THE NATIONAL MUSEUM 



VOL. 121 




Figure 5 -Male genitalia of M,lanot.i species: a, lam b, c, tilosmj d, ,, "Pf""''' '• 
Sagittarius; n, corticinus. (Scale lines=O.S mm.) 



MELANOTUS — QUATE AND THOMPSON 



77 




Figure 6.— Male genitalia of Melanotus species: a, prasinus; b, testaceus; c, trapezoideus; d, 
tenax; e, f, macer; g, difficilis; h, parallelus; i, infaustus; j, americanus; k, cribriventris; /, 
perUnax; m, beameri. _:^^{Scak lines=0.5 mm.) 



78 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 7— Male genitalia of Melanolus species: a, concisus; b, obscuratus; c, lanceaius; d, 
hamatus; e, longulus longulus;/, longulus oregonensis; g, gradatus; h, insiptens; i, chiricahuae; 
j, cribricollis; k, leonardi; I, taenicollis. (Scale lines=0.5 mm.) 



MELANOTUS — QUATE AND THOMPSON 



79 




^Tul ^-"^^""^'^ genitalia of Melanolus species: a, hypothetical species with structures 
labe led; b, stmilis, "normal"; c, similis, "spiny"; d, decumanus; e, spadix; f, castanipes. 
(bcale hnes= 1.0 mm and 0.1 mm.) 



80 



PROCEEDINGS OF THE NATIONAL MUSEUM 

h 




Figure 9.— Female genitalia of Melanotus species: a,h, communis; c, miscellus; d,e, dietrichi^ 
f,g, indistindus; h, land; i, verberans; j, opacicoUis; k, emissus ( ?) ; /, pUosus. (Scale Imes- 
1.0 and 0.5 mm.) 



MELANOTUS — QUATE AND THOMPSON 



81 




Figure 10— Female genitalia of Melanotus species: a, ignohilis; h, depressus; c, Sagittarius; 
d, cribulosus; e, morosus; f, hyslopi; g,h, prasinus; i, corticinus; j, piceatus; k, pertinax; 
I, macer. (Scale lines= 1.0 and 0.05 mm.) 



82 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figure 11— Female genitalia of Melanotus species: a, tenax; b, trapewideus; c,d, teslaceus; 
e,f, americanus; g, parallelus; h, cribriventris; i,j, infaustus {>); k, gradatus; I, obscuratus. 
(Scale lines^O.S and 0.05 mm.) 



MELANOTUS — QUATE AND THOMPSON 



83 




Figure 12.— Female genitalia of Melanotus species: a, concisus (?); b, lancealus; c,d, longulus 
longulus; e,f, longulus oregonensis; g, insipiens; h,i, leonardi; j, taenicollis; k, chiricahuae; 
l,m ,cnbrjcollis. (Scale lines=0.5 and 0.05 mm.) 



us. GOVERNMENT PRINTING OFFICE: 1967 



PROC. US. NAT. MUS. VOL. 121 



QUATE AND THOM PSON— PLATE t 




K^i»i^- 




^^^^ 






Heads of Melanotus species: a, similis; b, decumanus; c, castanipes; d, communis; e, vc-rberans; 
f,lanei; g, pilosus; h, emissus; i, opacicolhs; j, depressus : k, Sagittarius; I, crihricollis; m, 
tenax; n, par-allelus; n, 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. McFadden ^ 



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 representmg 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 withm the family 
and between it and other closely related familes. 

Any attempt to establish subfamilial and generic lunits 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 mil have to be 
made in keys and definitions of taxa. 

The keys have been prepared chiefly for identification of last 
mstar larvae. If earher mstars are known, they either have been 

1 Modified from a Ph. D. dissertation submitted to the University of Alberta 
E(hnonton, Canada. ' 

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



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 wdsh 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) — AMNH 

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, S. 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) — DEI 

Illinois Natural History Survey Collection, Urbana, 111, (L. K. Gloyd) — INHM 

Louisiana State University, University Park, La. (H. V. Daly) — LSU 

M. W. McFadden Collection— MWM 

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) — UMICH 

University of Missouri, Columbia, Mo. (W. R. Enns) — UMO 

United States National Museum, Washington, D.C. (W. W. Wirth)— USNM 

Washington State Universitj^, 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. 



^'^- ^^"^ 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 calciiun 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 Resume 

Frorn 1720 to the present, nimierous 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 distmct periods. 

The first period (1720-1868) was devoted to the description of 
local faimas.^ In most mstances larvae were associated mth 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 (18G9), was 
characterized by the incorporation of biological features and the 



4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i2j 

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 imder 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 follo\ving discussion comprises a review of the literature on 
the biology of the Stratiomyidae along the lines of a general life 
history, A\dth 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 fife 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 

imder laboratory conditions and that the pupal stage may last from 
2 weeks to 5 months. May (1961), also working with 77. illucens, 
gave more precise figm-es for the duration of the various stages in 
the life cycle and, in addition, was able to complete the life cycle 
successfully. 

Eggs. — ^Eggs usually are laid in early spring by females that have 
ove^^vintered as larvae. When first laid, the eggs are pale yellow but 
become progressively darker as the hatchmg 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- 
Limd (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-Limd (1943) also mentions that the eggs of 
Stratiomyidae are difficult to distinguish from those of Tabanidae. 

Larvae. — Newly hatched larvae were noted by Irwin-Smith (1921) 
to molt as soon as they emerged from the egg. Active feeding com- 
menced unmediately. As stated previously, the duration of the larval 
state is unlmown 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 then- feeding habits (James, 1960), but there is 
considerable variation in habitat selection by species within the 
subfamiUes. 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 

Pupae. — The 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. 

Adults. — 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 respuatory system, the head capsule, and the integument. 



NO. 3509 SOLDIER FLY LARVAE — McFADDEN 7 

While the quahty 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 unkno^\Ti manner. Richards (1951), on the contrary, is of the 
opinion that the calcium deposits are laid do^vn on the external sur- 
face of the integument as a result of carbon dioxide diffusing outward 
through the mtegument 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 wiU 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 

'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 shoA\Ti 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 foimd 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 \\ithout legs or 
prolegs. 

Abdomen. — 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 spu-acular 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 sprues 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-Lmid, 1943; Lenz, 
1923) have reported that the hooks serve as anchoi-s 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 fimctionally 
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 fimction 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 wdth long hydrofuge setae, which enable the 
larvae to remain suspended from the surface of water for indefinite 
periods of time while the spu-acles are open to the atmosphere. When 
submerged, these setae also aid respiration by enclosing an air bubble 
trapped m 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 wdth 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 
(Chu'omyzinae) . 

As previously indicated, the environment in which the larvae live, 
their feeding habits, and the type of mouthparts are closely aUied 
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 11 

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 

E. 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 requii'ements 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 PROCEEDESrGS 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 Hermetiini. The mouthparts of 
the latter are very similar to those of the Sarginae and both groups 
live in the same type of enAoroment— 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): 



SOLDIER FLY LARVAE — McFADDEN 



13 



James 
1. 

2. Chiromyzinae — Altermetoponia 

3. Beridinae — Allognosta, Actina, 

Beris, Exodontha 
Sarginae — Sargus, Ptedicus, 

Microchrysa, Chloromyia,* 

Merosargus 
Cyphomyiinae — Cyphomyia, 

Dicyphoma 
Hermetiinae — Hermetia 



4. 



\ 9- 
i 10. 



Clitellariinae — Adoxomyia, 
Dieuryneura, Brachycara,* 
Euryneura,* Oxycera* 
Euparyphus, Caloparyphus, 
Akronia,* Nemotelus 

Stratiomyinae — Stratiomys, 
Hoplitomyia, * Labostigmina, * 
Anoplodonta,* Hedriodiscus, 
Odontomyia, Nothomyia, * 
Myxosargus 

Pachygastrinae — Neopachygaster, 
Eupachygaster, Pachygaster, 
Zabrachia, Berkshiria 



McFadden 

1. Xylomyinae — Xylomya, Solva 

2. Chiromyzinae — as in James 

3. Beridinae — as in James 

4. Sarginae — as in James 



5. Cyphomyiinae — see under 

Clitellariinae 

6. Hermetiinae — see under 

Clitellariinae 

7. Clitellariinae — Dieuryneura 

Cyphomyiini — Cyphomyia, 
Adoxomyia, Dicyphoma 
Hermetiini — Hermetia 

8. Stratiomyinae — Stratiomys, 

Hedriodiscus, Odontomyia, 
Myxosargus, Euparyphus, 
Caloparyphus, Aochletus 

9. Nemotelinae — Nemotelus 

10. Pachygastrinae — as in James 



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. Al- 
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 aSoZw 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 Chtellariinae. 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 Adoxomyia. 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 Adoxomyia. 

A second tribe, the Hermetiini (Hennetiinae 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 clitellariiue genera Adoxomyia, 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 tmce 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 wdth 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 



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 Division CYCLORRHAPHA 

No free cephaloskeleton within the 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 

in last larval skin 4 

Posterior spiracles visible, situated on apical, penultimate or antepenultimate 
segment, body not shagreened or visibly striated; pupa free 5 

4. Body 12-segmented, cylindrical, not shagreened, usually longitudinally 

striated, abdomen with a girdle of pseudopods on each segment; head 

retractUe; spiracular fissure vertical TABANIDAE 

Body 11-segmented, bristly, surface finely shagreened, without pseudopods; 
head not retractile; spiracular fissure transverse, sometimes rather small; 
peripneustic or amphipneustic STRATIOMYIDAE 

5. Posterior spiracles situated on penultimate or antepenultimate segment. 

THEREVIDAE, SCENOPINIDAE, MYDAIDAE, ASILIDAE, 

BOMBYLIIDAE 
Posterior spiracles situated on apical segment 6 

6. Last abdominal segment obliquely truncate and with projecting 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. 

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 integimaent (see p. 7). 

Family diagnosis. — Head elongate and narrow except for ocular 
prominences; not retractile; antennae usually located anterolateraUy 
but varying among species; setae on dorsum of head variable; body 
11-segmented; prothoracic segment mth functional spiracles, that 
segment ^^ider than the head; segment posterior to prothorax either 
parallel mth the lateral margins of the prothorax or tapering in a 
posterior dh-ection from the second abdominal segment as in Stratiomys 
(fig. 105); posterior spu'acles located m a transverse cleft on last 
abdominal segment; with or mthout 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 

apex STRATIOMYINAE 

Last abdominal segment with coronet of plumose or pinnate setae absent 
from apex 2 

2. Last abdominal segment with a coronet of plumose or pinnate setae on 

dorsum NEMOTELINAE 

Last abdominal segment lacking a coronet of plumose 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 mesothoracic segments with normal shagreened pattern on dorsum; 

anus not bordered anteriorly by teeth 4 

4. Mouthparts highly sclerotized and foreshortened, mandibles well developed, 

no setae or bristles present on mouthparts (fig. 15) ; recorded from San 

Francisco area only CHIROMYZINAE 

Mouthparts with a different combination of characters than those listed 
above; occurring in San Francisco area or elsewhere 5 

5. Dorsal row of transverse setae surrounded by smaller setae; mouthparts as 

in figure 16; uniformly colored larvae BERIDINAE 

Dorsal row of transverse setae not surrounded by smaller setae; mouthparts 
different than above; larvae uniformly colored or with a 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 imder 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 segment with a transverse row 
of tubercles on dorsum Solva Walker 

Prothoracic segment of larva lacking cleft appearance of margin; no tubercles pres- 
ent on dorsum of apical abdominal segment Xylomya Rondaiii 

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 

Figures 149, 150 

Description.— Pupariiim: length (head capsule lacking) 15.2 mm; 
width 4.1 mm. Other characters as given in key to genera. 

Biology. — Larvae of this species were removed from a hole in a 
sycamore tree. 

Material examined. — Virginia: Alexandria, Oct. 14, 1951, W. W. 
Wu-th, 1 puparium (USNM). 

Xylomya species 

Figures 7, 11, 25 
Description. — Mature larva: length 13 mm; wddth 3.2 mm. Other 
characters as given in key to genera. 

Biology. — 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 

Figures 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, pointing 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 sph-acle on the antero- 
lateral surface; spiracle small, golden yellow, slightly elevated; last 
segment roimded at apex and entirely smooth. Length 6.0 mm; 
diameter of thorax 1.85 mm; diameter of abdomen 1.75 mm. 

Biology. — Larvae have been collected from the follo^ving 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 foimd the larva 
of Solva to be predaceous on the larvae of Euxesta, 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 mth 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 imtil 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 ia June, July, and August. 

Material examined.— Illinois: Rock Island, Apr. 5, 1932, C. O. 
Mohr, 2 larvae, INHM. Pennsjdvania: 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 imder 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 owti as Hennig (1952) has done. Because 
of a scarcity of specimens iti 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 



NO. 3569 SOLDIER FLY LARVAE — McFADDEN 19 

australis Macqiiart, the only published information available on the 
larvae of this subfamily. 

The subfamily is represented in North America only by Altermeto- 
ponia nibriceps (Macquart). 

SuBFAMiLiAL CHARACTERS. — As giveu ill key to subfamilies. 

Habitat of larvae. — The only species of Chiromyzinae for which 
habitat data are available is A. rubriceps. 

Genus Altermetoponia Miller, 1945 

Figures 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 bro^vn, darkest at apex. 

Biology. — 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 Beridiaae 

Of the 5 genera that represent this subfamily in North America, 
the larvae of only 2 are known: Allognosfa and Exodontha; however, 
larvae of 2 of the remaining genera are laiown from other geographical 
areas and descriptions of these species have been included. 

SuBFAMiLiAL CHARACTERS. — As givcn in the key to subfamihes. 

Habitat of larvae. — The larvae in this subfamily are terrestrial 
but may be associated wath semiaquatic envh'onments; 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 foimd in decaying leaves. 
In addition to being 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, greater than 10 mm in length Exodontha Rondani 

Larva smaller, less than 10 mm in length 2 

2. Abdominal segments with lateral margin bilobed, the anterior smaller than 

the posterior one Allognosta Osten Sacken 

Abdominal segments with lateral margin smooth 3 

3. Last abdominal segment broadly indented at distal end and fringed with fine 

setae 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 Juscitarsis (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. 

Biology. — 'Little is known of the biology and life history of this 
species. Malloch's (1917) specimen was obtained from woods near 
Urbana, 111., 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, 
ace. 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 
FuUer (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. — Matiu-e hirva (?) : 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 segmeiits 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. 

Biology. — 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 21 

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 yeUow; body of larva extremely flattened 
dorsoventraUy; prothoracic spiracle elevated but relatively incon- 
spicuous; body segments from second thoracic to seventh abdominal 
with a distinct papilla at intersegmental fold. 

Biology. — 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 Exodontha luteipes (Williston) have 
been found. 

Generic characters. — As given in key to genera. 

Exodontha luteipes (Williston), 1885 

Figures 29, 30, 35 

Description. — 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. 

Biology. — 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, in the laboratory. 

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, Alicrochrysa, and Merosargus. 

The keys to the genera of this subfamily wiU work for mature 
larvae only, but, where earlier instars are known, descriptions of 
these forms are included. 

SuBFAMiLiAL CHARACTERS. — As givcn 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) Ptecticus Loew 

Venter of sixth abdominal segment with a shorter sternal patch; sternal patch 
lacking on venter of fifth abdominal segment 2 

2. Dorsolateral margin of segments 1-10 with a series of large, 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 

to tubercle 4 

4. Eye prominence anterior to protruding lateral tubercle . Merosargus Loew 
Eye prominence on protruding lateral tubercle 5 

5. Width of head at tubercle not more than 0.5 mm . . . Microchrysa 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. 



NO. 3569 



SOLDIER FLY LARVAE — ^McFADDEN 23 



Biology.— 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 

sessile 2 

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 

Figures 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. 

Biology. — 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. Vu-ginia: Alexandria, June 24, 1951, W. W. Wirth, 
15 larvae (early instar), from under horse dung, USNM. 

Sargus {Sargus) decorus Say, 1824 

Figures 55, 58, 59 

Description. — Mature larva: length 4.0-4.2 mm, mean 4.1 mm; 
\\ddth 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. 

Biology. — Larvae of this species have been collected from rotting 
leaves and cow manure. 



24 PROCEEDINGS OF THE NATIONAL IVIUSEUM vol. 121 

Material 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 

Figures 48, 54, 57 

Description. — Mature larvae: length 9 mm; width 3 mm. Other 
characters as given in key to species. 

Instar no. ?: length 6.5 mm; width 2.4 mm. Differs from mature 
larva since it lacks the vittae. 

Biology. — ^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 

Figures 17, 62-64 

Description. — Instar no. ?: length 6.5-7.4 mm, mean 7.04 mm; 
width 1.7-2.0 mm, mean 1.84 mm. Similar to S. decorus in this stage 
but much smaller. 

Biology. — 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. trivittatas Say have been collected north of Mexico. 
Generic characters. — -As given in key to genera. 

Ptecticus trivittatus (Say), 1829 

Figures 43, 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 25 

Biology. — 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 CaroUna: 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 sackenii Williston, 1885 

Description. — Mature larva: length 11 mm; width 4.5 mm. 

Biology. — No data available. 

Material examined.— Mexico: No other data, 2 larvae collected 
by A. Herrana, AMNH. 

Discussion. — 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 

Figures 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 -wddth of head at eye prominence to 
width at neck 1.32; venter of sixth abdominal segment mth the setae 
on each side of the sternal patch reduced; anal spines lacking. 

Biology. — 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 

piiparia 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 Locw, 1855 

Of the 2 species listed by James (pers. comm.) for this genus, only 
the larvae of Alerosargus 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) ; mdth 2.3 mm; head capsule similar to that of Sargus 
lucens but differing in key characters. 

Biology. — A puparium of this species was collected from debris at 
the base of a sqidrrePs nest. 

Material examined. — -Maryland: Linnieville, Jan. 1, 1914, R. C. 
Shannon, 1 pupariiun \^dth head capsule detached, USNM. 

Subfamily Clitellariinae 

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 A\dth the genus 
Adoxomyia to form the tribe Clitellariini and have reduced the sub- 
family Hermetiinae to tribal status. 

The genus Dieunjneura 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. 35G9 SOLDIER FLY LARVAE — McFADDEN 27 

range as far north as Oregon and Washington on the west coast and a 
third species, subulata, is found throughout many of the Eastern 
States. Fourteen species have been collected from the southwest as 
adults, but the genus Adoxomyia, which contains 11 of the 14 species, 
is known only from 3 puparia that lack the head capsule. 

Habitat of larvae. — All 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 bodj^ axis. 

Cyphomyia Wiedemann 

Median seta no longer than setae on either side of it (fig. 78) ; all 3 setae set 

on a Une diagonal to the body axis Dicyphoma James 

Genus Cyphomyia Wiedemann, 1819 

James (pers, comm.) recognizes 2 species in this genus: Cyphomyia 
bicarinata WUliston and C. marginata Loew. Larvae have been col- 
lected for both species, but because the specimens of C. 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 

FiGUE,Es 152-154 

Description. — Puparium: length 20.1 mm; width 2.9 mm; other 
characters as given in key to genera. 

Biology. — 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 fu'st 7 
abdominal segments somewhat elevated with low transverse ridges oc- 
cupying approximately the anterior and posterior foiu"th or fifth of the 
segment; ventrally these areas are only slightly elevated. 



28 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Biology. — 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. pilosissima from maguey (Agave species). 
No other data are available. 

Genus Dicyphoma James, 1937 

Only 1 species, Dicyphoma schae_fferi (Coquillett) , occm's 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. 

Dicyphoma schaefferi (Coquillett), 1904 

Figures 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. 

Biology. — 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, 1 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 2.8 mm. Other characters as given in key to genera. 

Biology. — No data available for A. rustica but larvae of A. clari- 
pennis have been collected from decaying cacti. 



N"- '^'"' SOLDIER FLY LARVAE — McFADDEN 29 

Material examined. — Washington: Tampico, Mar. 1, 193], A. R. 
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 hetninopla (Wiedemann), 1819 

Figures 74, 77, 80 

Description.— Mature larva: the characters that appear to be of 
generic significance are given m the key to genera. Specific characters 
are unimportant since the species does not occur within the geographic 
range covered in this paper. 

Biology.— Larvae of this species were collected from Kaempferia 
species, a tuberous-rooted plant. It has also been reported from 
papaya by Brimetti (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 
illucens (Linnaeus), has been recorded from as far north and east as 
Maryland and Delaware. 

Tribal characters. — As given in key to subfamilies. 

Habitat of 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 

on lateral margins 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 
median line o 

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 
leg group 3 



30 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

3. Labrum barely extending beyond ocular lobes; point not drawn out, blunt 

appearance concinna Williston 

Labrum extending beyond ocular lobes, end portion drawn out into a taper- 
ing point 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 

Figures 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 papUla 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. 

Biology. — 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. illucens, 
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. Fm-man, 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: FayetteviUe, 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 Michclbachcr, 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 cidl 
pile, UD; Thompsonville, Oct. 24, 1957, Al.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 manm-e, 
LSU. Maryland: Beltsville, Aug. 9, 1957, J. C. Hwang, 4 larvae 
from tiu-key manure, USNM. Missouri: St, Louis, Jidy 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 rottmg 
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 manm-e, 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 

Figures 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. 

Biology. — 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. 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Hermetia species 

Figures 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. 

Biology.— Same as noted for H. concinna. 

Material examined. — Three larvae with same data as for H. 
concinna. 

Discussion. — 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 

Figures 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. 

Biology. — 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 
Dteuryneura obscura (Coquillett), 1902 

Figures 16, 24, 27, 32 

Description. — Matm-e larva: length 14.9 mm; width 4.0-4.3 mm, 
mean 4.1 mm; head and body segments dark brown. 

Biology. — 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. 



NO. 3569 SOLDIER FLY LARVAE — McFADDEN 33 

SuBFAMiLL\L 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-flowing brooks to ocean shores. Although 
larvae in the genus Stratiomys 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 

(fig. 122) 5 

Seventh abdominal segment without curved sclerotized hooks on venter . 2 

2. Antenna located at apex of ocular lobe (fig. 126) 4 

Antenna not located at apex of ocular lobe . Caloparyphus James (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 

seventh present 5 

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 edge of lower 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 7 

Antenna at apex of ocular lobe; prothoracic spiracles neither elevated nor 
stalked 8 

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- 
tenna Euparyphus Gerstaecker 

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 Geoffrey, 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 m key to species 

Key to Species of Stratiomys Geoffrey 

(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 

that segment discaloides Curran 

Length of last abdominal segment greater than 2 times the basal width of 

that segment adelpha Steyskal, badia Walker, 

barbata Loew, meigenii Wiedemann, norma Wiedemann, normula Loew. 

Stratiomys discaloides Curran, 1922 

Figure 119 

Description.— 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. 

Biology.— 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.— Puparimn: length 31.3-41.0 mm; width 4.9-6.0 mm; 
color varymg from grey brown to dark brown; markmgs same as for 
S. discaloides. Other characters as given in key to species. 

Biology.— See biology section in this paper. 

Material examined. —S'fraiiomys 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 imder 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 meigenii Wiedemann: 
Kansas: Manhatten, Feb. 8, 1936, H. H. Schwardt, 2 puparia, WSU. 
Stratiomys norma Wiedemann: Wisconsin: T9N Knapps Creek, 
Richland Comity, R2W, June 10, 1954, R. H. Jones, 4 puparia, WSU. 



^°- ^"5^^ SOLDIER FLY LARVAE — McFADDEN 35 

Stratiomys normula Loew: Louisiana: Baton Rouge (?), 2 puparia 
LSU. 

Genus Myxosargus Brauer, 1882 

Four species of Myxosargus occur in North America but only the 
puparium of Myxosargus nigricomis Green is known. 
Generic characters.— As given in key to genera. 

Myxosargus nigricomis Green, 1918 

Figures 88, 92, 93 

Description.— 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. 

Biology. — No data are available for this species. 

Material 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 Euparyphus, only the larvae of E. limhrocutris Adams have 
been collected. Aochletus has been elevated to full generic status. 

Generic characters.— As given m key to genera. 

Euparyphus limhrocutris Adams, 1903 

Figures 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. 

Penultunate 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 sphacles are elevated as high as their basal diameter. 

Biology.— No data are available for this species. 

Material examined.— Washmgton: O'SuUivan Dam, Grant 
Coimty, May 7, 1955, J. A. Quist, 2 matm-e 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 Euparyphus 
is so great that I feel full generic status should be given to Aochletus. 



VOL. 121 



36 PROCEEDINGS OF THE NATIONAL MUSEUM 

Of the 3 species listed by James (pers. comm.) for this taxon, larvae 
have been coUected for the followmg 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 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 m 
key to species. 

Biology.— No data are available for this species. 

Material examined.— California: Topanga Canyon, Los Angeles 
Coimty, May 10, 1953, 3 puparia, WSU; Sespe Creek, Ventura 
County, June 15, 1948, W. W. Wu-th, 1 puparium, USNM. 

Aochletus brevicornis Loew, 1866 

Figures 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 ha key to 

species. 

Biology.— Larvae have been collected from springs m YeUowstone 
National Park. No mention is given regardmg the type of sprmg 
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, YeUowstone National Park, 
June 20, 1956, J. R. Miu-phy, 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- 
cidt 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 Euparyphus. 

James (pers. comm.) lists 11 species for this genus in North America. 
Associated larvae or puparia have been collected for 5 species, m- 
cluding 1 unidentified species. 

Generic characters.— As given in key to genera. 



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 

segment 3 

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 than 4 . 4 

4. Three setae in mesothoracic leg group tetraspilus (Loew) 

Five setae in mesothoracic leg group crotch! (Osten Sacken) 

Caloparyphus species 

Figures 112, 115, 118 

Description. — 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. 

Biology. — 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 

FiGUEEs 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. 

Biology. — No data are available for this species. 

Material examined. — California: Sespe Creek, Ventura County, 
Jime 15, 1948, W. W. Wirth, 20 larvae, USNM. 

Discussion. — Accurate measurements of length were unobtainable 
due to improper preservation of the specimens. 

Caloparyphus major (Hine), 1901 

Figures 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. 

Biology. — 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 
Figures 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 

Biology. — 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 

Figures 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. 

Biology. ^ — 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. vertehratus (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 

Biology. — James (pers. comm.) reported that larvae of this species 
live among floating vegetation in small streams. They feed on micro- 
organisms, algae, and the soft parts of plants. 

Material examined. — Locality (?) ; Jidy 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. 3569 SOLDIER FLY LARVAE — McFADDEN 39 

Subgenus Catasina Enderlein, 1914 

Odontomyia (Catasina) pubescens (Day), 1882 

Figures 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. 

Biology. — 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; Bilberry 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 

Figures 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. 

Biology.- — 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 

vittae on apical segment cincta Olivier 

Vittae in different pattern; body segments with fine white pubescence. 

communis James 



40 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i2i 

Odontomyia (Odontomyia) occidentalis James, 1936 

Figures 135, 136, 138 

Description. — Puparium: length 20 mm; width 3.5 mm; other 
characters as given in key to species. 

Biology. — 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 

Figures 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. 

Biology. — 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 

Figures 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. 

Biology. — ^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 Nemoteliiiae 

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 givcii in key to subfamilies. 



^'"- 3^«9 SOLDIER FLY LARVAE — McFADDEN 41 

Genus Nemotelus GeoiTroy, 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 

Biology.— Larvae of this species were collected from a salt spring 
(probably from the margin) 

Material examined.— Missouri: Petersburg, June 1, 1955, P. J. 
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 
Plaques centralis Hanson 

Larva 5 mm or more in length; dorsum of body segments with vittae and plaques 
(fig. 121) canadensis Loew 

Nemotelus (Camptopelta) centralis Hanson, 1958 

Figures 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. 

Biology.— No data are available for this species. 

Material examined.— Michigan: Cheboygan County, Aug. 17, 
1957, W. J. Hanson, 3 larvae, lab reared, KU 

Nem,otelus (Cam,ptopelta) canadensis Loew, 1863 

Figures 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-r.?.3— G6 4 



42 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Biology. — Larvae of this species have been collected from under 
cow mannre 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 resume 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 Eupachygaster. 

I have found variation in the measurements given by Ki-aft and 
Cook as compared with those taken by myself. In one species, 
Eupachygaster henshawi MaUoch, 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. 

SuBFAMiLTAL CHARACTERS. — As given in key to subfamilies. 

Habitat of larvae. — Pachygastrine larvae have been found under 
the bark of both deciduous and coniferous trees. Oviposition usually 
occm's 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 Ki'aft and Cook are: apple, 
quaking aspen, dwarf elm, American elm, cottonwood, hickory, white 
pine, Douglas fir, Engelmann spruce, and 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 i 
beneath the bark. MaUoch (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) 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 alhistylum, 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 alhistylum Johnson, 1914 

Figure 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. 

Biology — Larvae of B. alhistylum have been collected from under 
the bark of poplar (Populus deltoid es) 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. — Eight 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. 

pHcatum Kraft and Cook 



44 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Zabrachia politum Coquillett, 1901 

Figure 145 

Description. — Mature larva: length 4.3 mm; width 1.0 mm; other 
characters as given in key to species. 

Biology — No data are available for this species. 

Material examined. — No larvae of this species were examined. 

Zabrachia plicatum Kraft and Cook, 1961 

Figure 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. 

Biology. — Larvae have been collected from beneath the bark of 
Pinus contorta, P. ponderosa, Picea engelmanni, and Pseudotsuga 
mucronata. 

Material examined. — North Carolina: Kaleigh, 1941-1942, 9 
larvae from fallen pine, NCSC. 

Discussion. — 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 

Description. — 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. 

Biology. — Larvae were taken from under bark of Pinus ponderosa. 

Material examined. — No larvae of this species were examined. 

Discussion.— AU data presented for this species are taken directly 
from Kraft and Cook (1961). 



Nt>. 3569 SOLDIER FLY LARVAE — McFADDEN 45 

Neopachygaster maeulicornis (Hine), 1902 

Figures 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 m thoracic leg group same as in A^. occidentalis. 

Biology. — 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. 

Discussion. — 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. maeulicornis. 
Biology. — 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. maeulicornis; tufts of thoracic leg setae with the 
middle setae longest, inner seta next longest. 

Biology. — No data are available for this species. 

Material examined. — No larvae of this species were examined. 

Discussion.— 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 

rows punctifer Malloch 

Setae in transverse rows with the outermost setae much shorter than the 
others in the row 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 equal length 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. 

Biology. — Larvae have been collected from imder the bark of 
Carya, Pojpulus, and an unknown species of Pinus. 

Material 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 

Figure 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 

Biology. — 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. E,. Dodge, 5 larvae from under 
willow bark, NCSC. 

Eupachygaster henshatvi Malloch, 1917 

Figure 144 

Description. — Mature Larva: length (4.95 mm) 5.7-6.9 mm, J 
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 

Biology. — Larvae have been collected from under the bark of 
apple, elm, and oak trees. 

Material examined. — North Carolina: Aubm'ii, 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 Mcigcn, 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 

Figure 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. 

Biology. — 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). 



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221-533—66- 



58 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 1. — Habitats of stratiomyid larvae occurring in America north of Mexico 
(genera arranged by subfamily as given on p. 13) 



Genus 


Habitat 


Reference 


Xylomya 


Terrestrial; in crotch of tree 


New record 


Solva 


Terrestrial; under bark of 


Townsend, 1893; Malloch, 




trees; in rotted logs 


1917; Johannsen, 1922; 
Greene, 1926; Peterson, 
1951;Hennig, 1952 


Alter meto'ponia 


Terrestrial; in sod 


Irwin-Smith, 1920 


Allognosta 


Terrestrial; in decaying 


Malloch, 1917; Johannsen, 




organic material 


1922 


Actina 


Terrestrial; in decaying 
plant and animal material 


Fuller, 1934 


Beris 


Terrestrial; in decaying 


Williston, 1908; de Meijere, 




leaves; under bark of 


1916; Lenz, 1923 




fallen trees; in moss? 




Exodontha 


Terrestrial; in rotten wood 
under large rocks 


New record 


Sargus 


Terrestrial; in decaying 


Westwood, 1840; Lundbeck, 




plant and animal material; 


1907; Williston, 1908; 




in excrement 


Malloch, 1917; Johannsen, 
1922; Peterson, 1951; 
Hennig, 1952 


Pteciicus 


Terrestrial; in decaying 


Lindner, 1928; Hennig, 




organic material 


1952 


Microchrysa 


Terrestrial; in decaying 


Lundbeck, 1907; Malloch, 




organic material; in 


1917; Johannsen, 1922; 




garden soil; in excrement 


Seguy, 1926; Hennig, 
1952 


Merosargus 


Terrestrial; in debris at 
base of squirrel's nest 


New record 


Chloromyia 


Terrestrial; in decaying 


Brauer, 1883; Cornelius, 




organic material 


1860; Lundbeck, 1907; 
Seguy, 1926 


Cyphomyia 


Terrestrial; in decaying 
plant material 


James, 1957 


Dicyphoma 


Terrestrial; in decaying 
plant material 


James, 1962 


Adoxomyia 


Terrestrial; in decaying 
plant material 


New record 


Hermetia 


Terrestrial; in decaying 


Williston, 1908; Malloch, 




organic material; in 


1917; Johannsen, 1922; 




excrement 


Copello, 1926; James, 
1935; James, 1497; 
James, 1957 


Dieuryneura 


Terrestrial; in decaying 
plant material 


New record 


Oxycera 


Aquatic; on margins of 


Heeger, 1856; Lundbeck, 




lakes, ponds, and streams 


1907; Johannsen, 1922; 
Lenz, 1923; Johannsen, 
1935; Wesenberg-Lund, 1943 



SOLDIER FLY LARVAE — McFADDEN 
Table 1. — Continued 



59 



Genus 


Habitat 


Reference 


Euparyphus 


Aquatic; on margins of 


Johannsen, 1922; 




aquatic environments; 


Johannsen, 1935; 




usually associated with 


Wesenberg-Lund, 1943; 




mossy conditions 


Peterson, 1951; James, 
1960 


Caloparyphus 


Aquatic; in bog or swamp 


Quist (thesis) ; James, 




areas; usually associated 


1960 




with mossy conditions 




Stratiomys 


Aquatic; on margins of 


Hart, 1895; Johnson, 




aquatic environments; 


1895; Miall, 1895; 




usually associated with 


Lundbeck, 1907; 




Typha spp. ; occasionally in 


Williston, 1908; 




hot springs or saline 


MaUoch, 1917; 




habitats 


Johannsen, 1922, 1935; 
Peterson, 1951; 
James, 1960b 


Hedriodiscus 


Aquatic; on margins of lakes 
and ponds 


James, 1960b 


Odontomyia 


Aquatic; in much the same 


Hart, 1895; Johson, 1895; 




habitat as given for 


Lunbeck, 1907; Williston, 




Stratiomys 


1908; MaUoch, 1917; 
Johannsen, 1922, 1935; 
Peterson, 1951; James, 
1960b 


Myxosargus 


Aquatic; no specific data 
available 


New record 


Nemotelus 


Aquatic; under debris at 


Hahday, 1857a; Lunbeck, 




margins of lakes and 


1907; Malloch, 1917; 




ponds; frequently in 


Johannsen, 1922; Lenz, 




saline habitats 


1923; Johannsen, 1935; 
Wesenberg-Lund, 1943; 
Hanson, 1958; James, 
1960 


Berkshiria 


Terrestrial; under bark of 


Kraft and Cook, 1961; 




trees 


Cook, 1953 


Zabrachia 


Terrestrial; under bark of 


Malloch, 1915; Malloch, 




coniferous trees 


1917; Kraft and Cook, 
1961 


Neopachygaster 


Terrestrial; under bark of 


Malloch, 1917; Cook, 1953; 




trees 


Kraft and Cook, 1961 


Eupachygaster 


Terrestrial; under bark of 


Malloch, 1917; Kraft and 




trees 


Cook, 1961 


Pachygaster 


Terrestrial; in tree holes; in 
crotch debris and in hollow 


Kraft and Cook, 1961 




trees 





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 * 


Odontomyia species 


New record 


Genus species 


Stratiomys species 






Chloromyia formosa 


Lundbeck, 1907 


Eulophidae 






Tetrastichus species 


Microchrysa polita 


Lundbeck, 1907 


Chalcidae 






Chalcis barbara 


Stratiomys norma 


Hart, 1895 


Chalcis microgaster 


Odontomyia cincta; 
Odontomyia species; 
Odontomyia vertebrata; egg 
mass of Odontomyia species 


Hart, 1895 


Ichneumonidae 






New genus, new species 2 


Stratiomys species 


New record 


Genus, new species 


Nemotelus species 


Lundbeck, 1907 


Genus, species 


Sargus species 


Lundbeck, 1907 



> Taxonomlc status as given by B. D. Burks, U.S. National Museum. 
2 Taxonomic status as given by L. H. WaliJely, U.S. National Museum. 

Table 3. — Ecological and morphological specializations in stratiomyid larvae 



Subfamily 


Environment 


Nutrition 


Mouthparts 


Xylomyinae 


terrestrial- 


micropantophagous 


cylindrical brushes 




arboreal 




present 


Chiromyzinae 


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 


cyUndrical brushes 
absent 


Nemotelinae 


terrestrial 


micropantophagous 


cylindrical brushes 
absent 


Pachygastrinae 


terrestrial- 


micropantophagous 


cyUndrical brushes 




arboreal 




present 



SOLDIER FLY LARVAE — McFADDEN 



61 



0=0-1 

A=2-3 
■ = 4-5 

# = 6-7 


D 

c 

1 




a 

c 
'n 

E 


i. 
's: 
U 


c 

'S- 

m 




c 




b 


(U 

D 

C 

E 
^-> 


<u 

D 
C 



E 
<u 

Z 


0) 

a 

c 

a 
en 
>. 

JZ 



s. 


Xylomyinae WM ^ 








▲ 


■ 








Chiromyzinae 


m 





A 


■ 


A 


A 


A 


Beridinae 




Ma 


A 


A 





A 


Sarginae 






■ 





■ 








Clitellariinae 










■ 


# 


A 





Slratlomyinae 












■1- 


# 


Nemotelinae 














m^ 


Pachygastrinae 
















■ 



Figure 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. 



62 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 2. — Proposed phylogeny for the eight stratiomyld subfamilies. 







Figures 3-13. — 3, Odontomyia species, head, dorsal view (after Cook, 1949; A= antenna, 
B=mandibular-maxillary complex, C=labrum, D=eye, E=clypeu8, F= ocular lobe, G= 
frontal suture); 4, Odontoviyia species, head, ventral view (after Cook, 1949; H= maxillary 
palp, 1= palatum, J=prementum, K = submentum, L= pharynx); 5, Odontomyia species, 
head, lateral view (after Cook, 1949; M= pestle, N=labrum, 0= 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 





Figures 14-24. — Dorsal viev/: 14, Soha pallipes, mandibular-maxillary complex; 15, Aller- 
metoponia rubriceps, mouthparts; 16, Dieuryneura obscura, mouthparts; 17, Sargus cuprarius 
mouthparts; 18, Neopachygaster maculicornis, mandibular-maxillary complex; 19, Cypho- 
myia hicarinata, labium and mandibular-maxillary complex; 20, Stratiomys norma, 
mandibular-maxillary complex (after Malloch, 1917); 21, Hermetia illucens, mouthparts 
(labrum removed); 22, Xylcxmya species, first abdominal segment; 23, Beris vallata, apical 
segment; 24, D. obscura, apical segment. 



SOLDIER FLY LARVAE — McFADDEN 



65 






f T t 1' ^ ,^ 




41 



Figures 25^1. — Dorsal view: 25, Altermeloponia rubriceps, head; 26, 5ifru vallata, head; 
27, Dieuryneura ohscura, head; 28, 5. vallata, first abdominal segment; 29, Exodontha 
luteipes, head; 30, £. luteipes, apical segment; 31, Allognosta fuscitarsis, head; 32, Z). 
obscura, first abdominal segment; 33, Altermetoponia rubriceps, apical segments; 34, 
A. rubriceps, first abdominal segment; 35, E. luteipes, first abdominal segment; 36, y^. 
fuscitarsis, first abdominal segment; 37, A. fuscitarsis, apical segment; 38, A. fuscitarsis, 
outline of prothoracic segment; 39, D. obscura, apical segment; 40, Aclina incisuralis, 
head; 41, A. incisuralis, first abdominal segment. 








53 






\ / 



57 




^ V 



60 





64 

Figures 42-64. — Dorsal view: 42, Microchrysa polita, head; 43, Pteclicus trivittaUis, head; 
44, M. polita, first abdominal segment; 45, Actina incisuralis, apical segment; 46, il/. 
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. 




Figures 65-82. — Dorsal view: 65, Ilermetia illucens, head; 66, H. concinna, head; 67, //. 
aurata, head; 68, Hermetia species, apical segment; 69, H. concinna, first abdominal seg- 
ment; 70, Hermetia species, head; 71, //. 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, A. heminopla, head; 81, D. 
schaefferi, apical segment; 82, D. schaefferi, head. 




; I 



I 1, 



-J- 



84 




85 















^ ^ t. ; , /• 




103 



Figures 83-104. — Dorsal view: 83, Cyphomyia pilosissima, head; 84, C. pilosissima, first 
abdominal segment; 85, C. pilosissima, apical segment; 86, Odontomyia puhescens, head; 
87, Hedriodiscus vertehratus, head; 88, Myxosargus nigricornis, apical segment; 89, 0. 
pubescens, first abdominal segment; 90, 0. puhescens, apical segment; 91, H. vertehratus, 
apical segment; 92, M. nigricornis, first abdominal segment; 93, M. nigricornis, head; 
94, Euparyphus limbrocutris, head; 95, E. limhrocutris, apical segment; 96, Aochletus 
brevicornis, first abdominal segment; 97, A. brevicornis, head; 98, 0. virgo, first abdominal 
segment; 99, E. limbrocutris, first abdominal segment; 100, Nemotelus centralis, head; 
101, A. brevicornis, apical segment; 102, 0. virgo, apical segment; 103, A'^. centralis, first 
abdominal segment; 104, N. centralis, apical segment. 



SOLDIER FLY LARVAE — McFADDEN 



69 



105 




25 

1.13 





Figures 105-119.— Dorsal view: 105, Stratiomys norma; 106, 5. norma, head; 107, 5. 
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. 




120 










136 





7/ V\- 



129 




( ^;^ 



138 



J39 



Figures 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. crotcki, head; 126, Odontomyia cincta, head; 127, C. amplus, head; 128, C. amplus, 
first abdominal segment; 129, C. crotchi, first abdominal segment; 130, 0. cincta, first 
abdominal segment; 131, C. amplus, apical segment; 132, C. crotchi, apical segment; 
133, 0. cincta, apical segment; 134, N. canadensis, head; 135, 0. occidentalis, head; 136 
0. occidentalis, apical segment; 137, 0. communis, apical segment; 138, 0. occidentalis, 
first abdominal segment; 139, 0. communis, first abdominal segment. 



SOLDIER FLY LARVAE — McFADDEN 



71 




Figures 140-148. — 140, Eupachygaster fusca, dorsal view on right; 141, Berkshiria albistylum, 
dorsal view on right; 142, Zabrachia plicalum, dorsal view on right; 143, E. pundifer, 
abdominal terga; 144, E. henshawi, abdominal sterna; 145, Z. poliium, terminal segments; 
146, Neopachygaster maculicornis, dorsal view on right; 147, A'', occidentalis, dorsal view 
on right; 148, Pachygaster pulchra, dorsal view on right. (After Kraft and Cook, 1961.) 



72 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 







150 



152 




.•o \ / o. 



153 







Figures 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. marghiata, first ab- 
dominal segment, dorsal view; 154, C. marginata, apical segment, ventral view; 155, 
Adoxomyia rustica, first abdominal segment, dorsal view; 156, A. rustica, apical segment, 
ventral view. 



U.S. GOVERNMENT PRINTING OFFICE: I9S7 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION • WASHINGTON, D.C. 

Volume 121 1967 Number 3570 



REVISION OF THE FAMILY PANDARIDAE 
(COPEPODA: CALIGOIDA)i 



By Roger Cressey 

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 

' Modified from a Ph. D. dissertation submitted to Boston University, Boston, 

Mass. 



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 encoiu"agement 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 5 reduced. Egg strings 
consisting of long strings of eggs arranged in linear series. 

Male, — Body caligiform, without dorsal plates. First thoracic 



NO. 3570 PANDARIDAE — CRESSET 3 

segment fused ^\dth cephalon. Thoracic segments 2-4 free. Oral 
area generally as in female. Legs 1-4 biramose, rami always with long 
plumose setae. Legs 5 and 6 present. Abdomen 1- or 2-segmented. 
Caudal rami large. 

Discussion. — 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 apertiu-e, 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 (MuUer). 

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: 
Pandanis, 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 GROUP I 

Dorsal thoracic plates, if present, on segment 4 only; 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 

areas and short spines GROUP II 

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 2 

2. Abdomen and caudal rami hidden dorsally Perissopus 

Abdomen and caudal rami visible dorsally Pandarus 

3. Plates of segment 2 large, overlapping those of segment 3 4 

Plates of segment 2 reduced and lateral to those of segment 3 5 

4. Maxilliped with spatulate tip Pannosus 

Maxilliped with pointed tip Phyllothyreus 

5. Maxilliped with pointed tip, genital segment less than }^ body length. 

Gangliopus 
Maxilliped with spatulate tip, genital segment at least Y^ body length. 

Pseudopandarus 



PANDARIDAE — CRESSET 



GENERA OP GROUP II FEMALES 



1. Abdomen 2-segmented 2 

Abdomen 1-segmented 3 

2. Fourth leg lamelliform Dinemoura 

Fourth leg not lamelliform Pagina 

3. Fourth leg lamelliform Echthrogaleus 

Fourth leg not lamelliform 4 

4. Abdomen with large dorsal plate Demoleus 

Abdomen without dorsal plate 5 

5. Exopods of legs 1-3 3-segmented Paranesippus 

Exopods of legs 1-3 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. 

Female. — ^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 maxilhped ; 
pads also located on ventral sm'face of posterior corners of cephalon. 
Claw of maxilliped spatulate. Legs 1-4 biramose, rami of legs 1-3 
2-segmented. Rami of leg 4 1-segmented. Leg 5 present. Egg 
strings long and straight. 

Male. — 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. 

Discussion. — 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 ttie posterior edge 

of the plate of segment 3 BICOLOR GROUP. 2 

Dorsal plate of thoracic segment 2 extending well beyond posterior edge of 
plate of segment 3 CRANCHII GROUP. 4 

2. Cephalon only Ys of total body length; caudal rami small, scarcely visible 

dorsally bicolor 

Cephalon about }^ of total body length; caudal rami easily visible dorsally . 3 

3. Caudal rami long, about 3 times as long as dorsal abdominal plate and extend- 

ing well beyond it; posterior edge of plate of segment 4 without conspicuous 

sinus niger 

Caudal rami only about 1^2 times as long as dorsal abdominal plate and 
extending only slightl}^ beyond it; posterior edge of plate of segment 4 with 
deep sinus carcharhini 

4. Caudal rami with the inner basal half expanded to broad lobe . . smithii 
Caudal rami without such a lobe 5 

5. Caudal rami not extending more than ji length of dorsal abdominal plate. 

satyrus 

Caudal rami extending at least to tip of dorsal abdominal plate or well 

beyond it 6 

6. Dorsal plates of thoracic segments 3 and 4 fused and with broad posterior 

sinus 7 

Dorsal plates of thoracic segments 3 and 4 divided by deep sinus into 2 
nearly separate lobes 9 

7. Dorsal abdominal plate subtriangular; eye spots separated in fully pigmented 

forms cranchii 

Dorsal abdominal plate subcircular; body may or may not be darkly pig- 
mented 8 

8. Most of body darkly pigmented but eye spots connected by median posterior 

line sinuatus 

Body only slightly pigmented with pigment usually couhned 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 

segment 4 zygaenae 

Thoracic plate of segment 2 extending to middle of thoracic plate of seg- 
ment 4 katoi 

Pandarus satyrus Dana, 1852 

Figures 1-26 
Pandarus satyrus Dana, 1852-3, p. 1368. — 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 — CRESSET 7 

Specimens studied. — All 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 
(L8 by 1.5 mm), longer than wide, extending beyond tips of rami. 
Caudal rami (fig. 2) long, slender, widest at base, tapering distally 
(990/i by 228ju), 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 B leg S leg i 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:0 0:0 1:0 0:0 VI 

seg. 2 VI III X 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 . 

Male. — Body form as in figure 16. Length (not including setae 
on caudal rami) 10.3 mm and width (measiu-ed 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 432)u long. Second segment 
576^1 long. Caudal ramus (fig. 18) 1050^ by 665m, bearing 4 long and 
2 short setae. Outermost long seta 1365/1 long, innermost 910m long, 
and 2 median setae 1500/i 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 S leg 4 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:1 0:1 1:1 0:1 1:1 0:1 

seg. 2 IV:3 3 IV:6 8 IV:5 6 IV:o 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 — CRESSET 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. 

Discussion. ^ — 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) . I have examined a gi-eat number of specimens 
of both of these species and conclude that both are valid species. The 
principal differences between them are discussed imder 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. — Kr0yer, 1837, 
p. 202.— Steenstrup and Lutken, 1861, p. 390.— Rathbun, 1884, p. 488; 
1886, p. 317.— Beneden, 1892, p. 221.— Wilson, 1907, p. 403; 1908, p. 453; 
1932, p. 435; 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 studied. — Eighteen 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 (Miiller and 
Henle) (30°49'N, 64°02'W; 57°44'N, 65°42'W). Two coUections 
from Eulamia obscura (Lesueur) (34°45'N, 73°41'W; 36°42'N, 
70°00'W). Ten collections from Eulamia jioridanus (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 galapagemis (14°3'N, 92°8'W) and Sphyrna 
zygaena (Linnaeus) off Formosa. Also 1 collection from P. longimanus 
from Dm-ban, 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. 

Female. — ^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 xiew 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 maxilHped (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. cranchii 
(fig. 30) with patch of more rugose spines along outer edge than that 
of P. satyrus. Leg 5 (fig. 31) as in P. satyrus. Eggs and egg strings as 
in P. satyrus. 

Color as in P. satyrus. 

Male. — ^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. 
cranchii (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 
caudal rami. 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 haK 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 hea^^ier in P. cranchii. 

Pandurus 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 hke other mem- 
bers of the genus is sometimes found in large clusters on the fins. 
It is world^vide in distribution. 

Shiino (1954) has described and figured both sexes of this species 
under the name of P. satyrus. 



NO. 3570 PANDARIDAE — CRESSET 11 

Pandarus smithii Rathbun, 1886 

Figures 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. 139.— Brian, 1944, p. 202.— Barnard, 1948, p. 249; 1955, 
p. 259.— Causey, 1955, p. 6.— Shiino, 1959b, p. 353. 

Pandarus luguhris Heller, 1808, p. 205. — Basset t-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 Isurus oxyrhynchus 
Rafinesque (42°18'N, 64°02'W; 35°00'N, 70°00'W). Two collections 
from Eulamia obscurus (Lesuem-) (36°42'N, 70°00'W; 34''45'N, 
73°41'W). Three collections from Eulamia jioridanus 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 Isurus oxyrhynchus (16''13'N, 
63°29'E) and Eulamia jioridanus (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) (Tres 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 
Sphyma zygaena (Linnaeus) off the coast of Brazil. 

Female. — 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) L78 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 I leg Z leg S leg 4 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:0 0:0 1:0 0:0 VI I or 

seg. 2 VI III X IV VI I or II 

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. 

Male. — 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. 
cranchii. 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. 

Discussion. — Pandarus smithii is a widely distributed copepod 
found on a number of hosts and often occurring with Pandarus cranchii 
on carcharinid sharks. P. smithii can be easily distinguished from 
the other species of the cranchii 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 Carvallio 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. 
smithii. 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 

FiGUEES 46-66 

Specimens studied. — 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 — CRESSET 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. 

Female. — Body form as in figure 46. Total length (based on an 
average of 5 specimens) 7.5 mm. Greatest width (measured at widest 
part of cephalon) 3.8 mm. Cephalon widest posteriorly narrowed 
anteriorly, measuring 3.5 by 3.8 mm. 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 7.2 mm). 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 490^ long bearing 25 spines and setae. Spines and setae 
armed as in the figure. Second segment 168/n 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 fu'st 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 g leg S leg i 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:1 0:1 1:1 0:1 1:1 0:1 

seg. 2 IV:3 3 IV:6 8 IV:5 6 IV:5 5 

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 segment and abdomen, consisting of a 
spineUke process with stout plumose seta near base. 



14 PROCEEDINGS OF THE NATIONAL MUSEUM 

Color in life cream and generally devoid of pigmentation. 
Legs 1-4 bii'amose, with spine and setal formula as follows: 





leg 1 
exp. end. 


legg 
exp. end. 


legs 
exp. end. 


leai 
exp. end 


seg. 1 


1:0 0:0 


1:0 0:0 


1:0 0:0 


VI 


seg. 2 


VI III 


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 attaclunent 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. 
Egg strings long and of usual generic type. 

Color in life creamy yellow with light brown pigmentation as indi- 
cated in figure 46. 

Male. — Body form as in figm'e 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 360/i long. Second segment 612;li 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 (264:576/x) (see fig. 59). Collected with this mature 
male was also a young male (7.1 mm) with spermatophores developing 



NO. 3570 PANDARIDAE — CRESSET 15 

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 (408:420/^) (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. floridanus the 



Table 1. — Comparison of the length of the dorsal abdominal plates with the 
length of dorsally exposed caudal ramus in some specimens of Pandarus flori- 
danus with P. sinvatus (CHML=Cape Haze Marine Laboratory, Fla.) 



Specimens 


Plate 


Ramus 


P. floridanus 






ex Carcharodon carcharias (avg. of 4) Dennis, Mass. 


702m 


450m 


" (avg. of 2) USNM 107299 


810m 


576m 


" " (1 spec.) USNM 101876 


720m 


540m 


ex Lamna cornubica (1 spec.) USNM 32765 


720m 


360m 


P. sinuatus 






ex Negaprion brevirostris (avg. of 2) CHML 


495m 


468m 


ex Carcharinus leucas (avg. of 2) CHML 


621m 


630m 



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 showmg 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 
cranchii 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 

Figures 67, 68 

Pandarus sinuatus Say, 1817, p. 436. — 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. Ill; 1951, p. 139.— Pearse, 1952a, 
p. 27; 1952b, p. 213.— 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 studied. — Two collections from Carcharinus leucas 
Miiller and Henle from Sarasota, Fla. From the same locaUty 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. 

Female. — Body form as in figure 67. Total length (based on an 
average of 5 specimens) 6.5 mm. 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. 

Male. — As in P. floridanus. 

Discussion. — 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 — CRESSET 17 

Pandarus katoi, new species 

Figures 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 (10$ 9 and 4 0^0^) in alcohol 
deposited in the U.S. National Museum. Fourteen paratypes (109 9 
and 4cf cf) 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. 

Female. — ^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 
2.3 mm. 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) 720^1 long and 270/x wide bearing 4 spines and 2 plimiose 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 600^ long and bearing 13 naked setae (fig. 72). Second 
antenna (fig. 73) 3-segmented. Terminal segment Avith 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 as in figm*e. 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: 



legl 
exp. end. 


legs 
exp. 


end. 


legs 
exp. end. 


leg 4 
exp. end. 


1:0 0:0 


1:0 


0:0 


1:0 0:0 


VII I 


VI III 


X 


IV 


VI II 





seg. 1 
seg. 2 

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 wdth 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 
spmes 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 witli 
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 wdth light brown pigmentation as in 
figure 69. 

Male. — Body form as in figure 83, Total length, based on 5 speci- 
mens, 7.4 mm. 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 300/x long. Second segment 
410/i 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 (204:516/i) (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: 



legl 
exp. end. 


legs 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


IV 3 


111:6 8 


IV:5 6 


IV: 5 5 



seg. 1 
seg. 2 

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 — CRESSET 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. 

Female. — 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) 3.9 mm. Cephalon somewhat truncated 
and measming 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. katoi except that they are generally held parallel to each other. 

Oral area and aU 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. katoi. 

Color in preserved material creamy white except for some light 
brown pigmentation in mature females as indicated in figure 97. 

Male. — 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. 

Discussion.— 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 sHme" 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 
segment 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.— Kr0yer, 1837, p. 202; 1838, p. 34; 1863, p. 261.— 
Milne-Edwards, 1840, p. 470.— Baird, 1850, p. 288.— Beneden, 1851a, p. 94; 
1861, p. 148.— Norman, 1868, p. 301.— Olsson, 1868, p. 21.— Richiardi, 1880, 
p. 149.— Cams, 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.ll.— Scott, T., 
1900, p. 157.— Scott, A., 1904, p. 40.— Norman and Scott, T., 1906, p. 211.— 
Wilson, 1907, p. 400; 1932, p. 436; 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 — CRESSET 21 

Pandarus boscii Leach, 1816, p. 406; 1819, p. 535. — Gu6rin-Meneville, 1829- 
1843, p. 41. — Burmeister, 1833, p. 331.— Desmarest, 1825, p. 339. — Kr0yer, 
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 Sgualus 
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. 

Female.^ — 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) 3.8 mm. Cephalon only about one- third 
body length, measuring 3.6 by 3.8 mm. 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 
exp. end. 


leg 2 
exp. end. 


leg S 
exp. end. 


leg 4 
exp. end. 


1:0 0:0 


1:0 0:0 


1:0 0:0 


VI I 


VI III 


VIII V 


VI II 





seg. 1 
seg. 2 

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. 

Male. — 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. izi 

Spine and setal formula (based on T. and A. Scott's illustrations) 
are as follows : 

leg 1 leg 2 leg 3 leg i 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:1 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 cranchii 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 

Figures 109-118 
Pandarus niger Kirtesinghe, 1950, p. 83. 

Specimens studied. — 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 (measirred at the widest part of the cephalon) 3.8 mm. 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 ai'ea as in other members of genus. First antenna (fig. Ill) 
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. hicolor. Fhst maxilla (fig. 113) with 
group of short spines on basal thhd 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 i leg 2 leg S leg 4 

exp. eud. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:0 0:0 1:0 0:0 VII II 

seg. 2 VI III X VII VI III 

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. 

Male. — '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. 

Remarks. — 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 seem to 
be preferred hosts. 

This copepod is closely related to P. hicolor, but the natiu-e of the 
caudal ramus separates it easily from all other members of the genus. 
Kirtesinghe (1950) notes the relationship between this species and 
P. hicolor; 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 Be, Madagascar, from Carcharinus leucas, C. sorrah, and 
C. limhatus. A single collection from Durban, South Africa, from C. 
leucas. 



24 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Female. — 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 featm-es have been figured 
for the sake of comparison. Caudal rami (fig. 120) intermediate 
between that of P. bicolor and P. niger, measurmg 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 bii'amose, with spine and setal formula as follows : 

leg 1 leg Z leg 3 leg i 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:0 0:0 1:0 00: VIII I 

seg. 2 VI III X 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. 

Male. — Unknown. 

Remarks.— This species described by Ho (1960) from Formosa has 
been collected by the author in Nosy Be, 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 w^estern 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 commvmication 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 — CRESSET 25 

Phyllothereus Norman, 1903, p. 368. 
Parapandarus Wilson, 1924a, p. 7. 

Female. — Frontal plate distinctly separate. Fii-st 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-segmented, those of leg 4 1-segmented. All legs without plumose 
setae. Leg 5 present. Egg sacs long and straight. 

Male. — 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 
synonjTny 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 Phyllo'phora 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 Laminifera. 
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-juradoi 
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 (MUne-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.^ — ^Tliree 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, claAV 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- 
hped (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 
exp. end. 


leg 2 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


1:0 0:0 


1:0 0:0 


1:0 0:0 


VIII III 


VII III 


VII VII 


VIII IV 





seg. 1 
seg. 2 

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 
eggs. 

Color whitish yellow. 

Male. — 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 figm-e 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 bu-amose and not as highly modified as in female. Spine 
and setal formula as follows: 



NO. 3570 


PANDARIDAE — CRI 


:SSEY 






leg 1 
exp. end. 


legs 
exp. end. 


legs 
exp. end. 


Ugi 
exp. end. 


seg. 1 


1:0 0:0 


1:0 0:1 


1:0 0:1 


1:1 0:1 


seg. 2 


IV:3 3 


IV :G 8 


IV:5 6 


IV:5 5 



27 



Leg 5 (fig. 137) located at beginning of distal third of genital segment 
and consisting of a ventral lobe bearing 1 outer pluniose 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 theii' 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 weU. 
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.] 

Female.— Frontal plate distinctly separate. Fu'st 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. Fu'st 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. 

Male. — 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 tetrapturi 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). 

Female. — 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: 





exp. 


leg 1 
end. 


legs 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 


0:0 


1:0 0:0 


1:0 0:0 


VII II 


seg. 2 


VII 


III 


X VI 


VIII III 





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. 

Male.— 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 — CRESSET 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 
exp. end. 


legi 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


Beg. 2 


IV:3 3 


IV:6 8 


IV:5 6 


IV:5 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. 

Yy.'E.uh.uK^.—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 tetrapteri. 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, 1951a, p. 1245. [Refers to P. longus only.] 

Female. — 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 
1-segmented. Leg 5 present and consisting of a free segment. Egg 
sacs long and straight. 

Male. — 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 plmnose 
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- 
duced into long pointed processes gracilis 

Abdomen partially visible in dorsal view, genital segment with posterior corners 
rounded and not greatly produced longus 

Pseudopandarus gracilis Kirtesinghe, 1950 

Figures 147-150 

Pseudopandarus gracilis Kirtesinghe, 1950, p. 84. 
Pseudopandarus scyllii Yamaguti and Yamasu, 1959, p. 124. 

vSpecimens studied. — Paratypes on loan from P. Kirtesinghe. 
6 females ex Scoliodon palasorrah, Nosy Be, Madagascar. 

Female. — 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^ bu-amose and with spine and setal formula as follows: 





exp. 


legl 
end. 


legi 
exp. end. 


legs 
exp. end. 


legi 
exp. end 


seg. 1 


1:0 


0:0 


1:0 0:0 


1:0 0:0 


V:3 2 


seg. 2 


VI 


3 


IV:5 7 


IV:4 3 





Sm-face 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. 

Male. — Unknown. 

Remarks. ^ — Pseudopandartts gracilis was described in 1950 by P. 
Kirtesinghe and was not reported again until 1959 by Yamaguti 
and Yamasu. They determined their species to be new, based 
primarily on the structure of the tip of the maxilliped. Unfortmiately, 
Kirtesinghe's figure of this was misleading. I was 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. Theu" 
figure shows no articulation between segments. I 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 3. The plates of segments 2 and 3 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 sm'face of small 
inshore sharks {Carcharinus and Triakis). 

Pseudopandarus longus (Gnananiiithu, 1951) 

Figures 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 Dm'ban, South 
Africa. 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Female. — 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 pliunose. 

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. Fu'st 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 maxiUa (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 Kii-tesinghe and P. longus (Gnanamuthu) 
shows the following common features: the aiTangement 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 Dm^ban, South Africa). So far, it has been reported 
only from carcharinid sharks. 

Pannosus, new genus 

Type-species: Gangliopus japonicus (Shiino, 1960). 

Female. — 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 basaUy. 
Abdomen 1-segmented. Caudal rami held laterally on abdomen. 
Adhesion pads of cephalon well developed. First antenna 2-seg- 



NO- 3570 PANDARIDAE — CRESSEY 33 

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. 

Male. — Unlcnown. 

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- 
liopus. After a redescription of the generic diagnosis of this genus, 
it became apparent that this new copepod did not belong to that 
genus. Suice I have had no material of this species to work with, 
no further diagnosis of Shiino's species can be made. I have removed 
this species from the genus Ganglioims for the foUowmg reasons: 
(1) Shimo'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 Lutken, 1861, p. 393. [Type-species: P. dentatus.] 
Chlamys Beneden, 1892a, p. 227. 
Achtheinus Wilson, 1908, p. 450. 

Female.— 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 coveHng 
anterior portion of genital segment. Abdomen 1-segmented and 
attached ventrally to genital segment. Caudal rami rudimentary. 
Fu-st antenna 2-segmented. Second antenna armed with teeth at tip. 
Mouth parts of typical pandarid type. Adhesion pads present. 
Maxdhped 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. Egg strings long and straight. 

Male.— 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^ 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 ^ol. 121 

endopod unmodified. Abdomen 2-segmented. Caudal rami attached 
distally to abdomen. . . 

Discussion.— This genus is represented by two species, Ferissopus 
dentatus Steenstrup and Liitken 1861 and P. ohlongatus (WHson, 1908) 
A discussion of the history of the synonomy of these species is mcluded 
in the species descriptions. 

Perissopus ohlongatus (Wilson, 1908) 

Achtheinus ohlongatus 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. 11. 
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. Smce then, 10 species assigned to this 
genus have been described. A comparison of the description and 
figures of these species show that they should be included m the genus 
Perissopus. This is apparent when one compares the appendages of 
the two groups. In both, the second antenna has a hoodlike process 
bearinc^ spines. The maxillipeds of each bears a reduced claw and the 
basal s^'egment is in the form of a flattened pad. Legs 1-4 are much 
reduced in both and are smiilar in form. 

I have grouped all species described in the genus Achtheinus 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— & single species 
with variation in body form. I have examined WHson's material of 
A ohlongatus, 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. galeorhini. i ■■ • to 

This species can be separated from P. dentatus on the basis ot z 
characters. In P. dentatus the posterior corners of the genital seg- 
ment are sharply angular whereas in P. ohlongatus they are rounded. 
In P. dentatus the endopods of legs 1-4 are unarmed. In P. oh- 
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 



PANDARIDAE— CRESSET 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. 
Chlamjjs incisus Beneden, 1892a, p. 227. 
Perissopus crenatus Leigh-Sharpe, 1930, p. 7. 
Perissopus manuelensis Gnanamuthu, 1951a, p. 1252. 
Perissopus travancosiensis Kurian, 1955, p. 108. 
Perissopus serratus Heegaard, 1962, p. 175. 

Specimen studied.— Five females and 1 male ex Carcharinus 
milberti 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 
Dm'ban, South Africa. 

All descriptions and figures refer to specimens from Carcharinus 
milberti unless otherwise stated. 

Female.— 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 spme near base. Tip of claw (fig. 166) with curious hooded 
appearance. Mouth tube of usual type, labium fringed at tip (fig. 



legl 
exp. end. 


leg 2 
exp. end. 


legs 
exp. end. 


leg A 
exp. end. 


1:0 0:0 


1:0 0:0 


IV 


IV 


IV 


IV 







36 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

167). Mandible of usual form. Fii'st maxilla (fig. 168) consisting of 
broad basal lobe with short spine at tip and gi'oup of 3 short setae. 
Second maxilla with tip armed as in figiu-e 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 siu"face 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 : 

seg. 1 
seg. 2 

Legs 1-4 small and wealdy developed. Endopod of aU legs unarmed 
except for patches of spinules. Segmentation reduced in legs 3 and 4. 
Leg 5 (fig. 175) consistmg 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 imarmed lobes located near junction of 
abdomen and genital segment (see fig. 163). 

Egg strings long and straight. 

Color in life cream white without pigmentation. 

Male. — Body form as in figrn^e 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 pliunose 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 figm'e 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 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:1 0:1 1:1 0:1 1:1 0:1 

seg. 2 111:4 3 111:6 8 11:6 6 111:5 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. AU setae plumose. Leg 5 (fig. 



NO. 3570 PANDARIDAE — CRESSET 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 maculipinnis, 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 fh'st time. It was attached to a 
female when found. 



38 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Genus Dinemoura Latreille, 1829 

Caligus Mliller, 1785, p. 132. [Refers to C. productus only.] 

Dinemoura Latreille, 1829, p. 197. [Type-species: D. produda.] 

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 iV". gracilis only.] 

Female. — Frontal plate distinctly separate. Tlioracic segments 
2-4 free. Dorsal tlioracic 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 setides at base of terminal claw. Maxilliped with 
small terminal claw. 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 in 
form of broad lamellae. Egg strings long and straight or folded. 

Male. — 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. 

DiscussioN.^ — -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 Hterature over the generic 
name of this group. In 1829 Latreille separated Caligus productus 
Mliller 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 oral area 2 

Adhesion pads absent or reduced on oral area 3 

2. Genital segment about K body length. Dorsal plates of segment 4 wider 

than long latifolia 

Genital segment about jz body length. Dorsal plates of segment 4 longer 
than wide producta 



NO- 3"o PANDARIDAE — CRESSET 39 

3. Posterior corners of genital segment not greath^ produced and rounded, ferox 
Posterior corners of genital segment greatly produced and truncated. 

discrepans 
Dinemoura producta (Miiller, 1785) 

Figures 190, 191 

C aligns productus Miiller, 1785, p. 132. 

Dinemoura product.— Latreille, 1829, p. 197.— Kr0yer, 1837, p. 202.— Scott, T., 
1901, p. 124.— Scott, T., and Scott, A., 1913, p. 86.— Norman and Scott, T., 
1906, p. 211.— Norman and Brady, 1910, p. 404.— Page, 1923, p. 281.— 
Pesta, 1934. p. 27. — Oorde and Schuurmana 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 
Vaissicre, 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.— 'Kr0yer, 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 OO'N, 70 00' W), Prionace 
glauca (Linnaeus) (39 32'N, 28 02'W), Carcharodon carcharias (Lin- 
naeus) off Boothbay Harbor, Maine, Cetorhinus maximiis (Gunnerus) 
on loan from Zoological Museum in Amsterdam. 

Female— A good description of the female of this species has been 
given as recently as 1957 by Shiino. His description is of Dinematura 
affinis 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: 





legl 
exp. 


end. 


leg 2 
exp. 


end. 


legs 
exp. end. 


leg I, 
exp. end. 


seg. 1 


1:0 


0:0 


1:1 


0:1 


1:1 0:1 


V 


seg. 2 


IV:3 


3 


1:1 


0:2 


1:1 0:2 




seg. 3 


- 


- 


111:5 


6 


111:5 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. 

Male. — The male of this species was first described by Wilson 
(1923). Because I had no material of the male of this genus, I 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. 

Remarks. — 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 literatiue previously (mainly 
Wilson, 1907) but I have, in addition to these, placed D. qffinis, 
Thomsen, 1949, in synonymy with D. producta. Shiino (1957) de- 
scribed and illustrated D. affinis 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 
hums. It is probably worldwide in distribution. 



NO. 3570 PANDARIDAE — CRESSET 41 

Dinemoura ferox (Kr0yer, 1838) 

Figures 192, 193 

Dinematura ferox Kr0yer, 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). 

Female. — 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 Kr0yer (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. Fu'st 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 
exp. end. 


leg 2 
exp. end. 


legs 
exp. end. 


leg 4 
exp. end. 


sag. 


1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


IV IV 


seg. 


2 


IV:3 3 


1:1 0:2 


1:1 0:2 




seg. 


3 




111:5 6 


111:5 4 





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. 

Male. — Hansen (1932) described a copepod identified by Kr0yer 
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 stiU remains poorly known and 
perhaps even yet to be described. 

Discussion. — Dinemoura ferox 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 So7nniosus microcephalus 
(Bloch and Schneider), reported by Wilson (1920), and Centrophorus 
squamosus MiiUer 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 LtJtken, 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; 1932, 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 Isurus oxyrinchus Rafi- 
nesqueinthe 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). 

Female. — 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 
figm*e 195. 

Since the female of this species has been weU described and figured 
by Yamaguti (1936) and Shiino (1954), I wiU only discuss saUent 
featiu-es. Adhesion pads associated with first and second antenna 
and maxiUiped. Oral appendages similar to those of D. producta. 

Legs 1-4 biramose with spine and setal formula as follows : 







leg 1 


legg 


legs 


leg 4 






exp. end. 


exp. end. 


exp. end. 


exp. end 


seg. 


1 


1:0 0:0 


1:1 0:1 


1:1 0:0 


V 


seg. 


2 


IV:3 3 


1:1 0:2 


1:1 0:2 




seg. 


3 




111:5 6 


111:5 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, spatiilate process as belonging to leg 5 and the curved, 
clawlike process at the junction of the genital segment and abdomen 
as leg 6. 

Male. — 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 figiu-e this 
appendage. 

Discussion. — This is a fairly common species occurring on the 
body surface of a number of pelagic sharks of the genera Isurus, 
Lamna, and Carcharodon. These 3 genera of sharks compose the 
family Isuridae (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. — Eighteen females and 13 males from the body 
surface of Alopias mdpinus (Bonneterre) from the Indian Ocean 
(9°24'N, 54°58'E). Holotype female (USNM 113592), allotype 
male, and 10 paratypes (59 9, 5cfcf) deposited in alcohol in the 
U.S. National Museum, 10 paratypes (59 9, 5cf cf) 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 Be, Madagascar; a single 
collection from Alopias superciliosus from the Pacific Ocean (0°38'N, 
124°23'W). 

Female. — 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 : 







legl 
exp. end. 


leg$ 
exp. 


end. 


legi 
exp. 


end. 


legi 
exp. end. 


seg. 


1 


1:0 0:0 


1:1 


0:1 


1:1 


0:0 


4 


seg. 


2 


IV:3 3 


1:1 


0:2 


IV:6 


0:2 




seg. 


3 




111:5 






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. 

Male. — Body form as in figm-e 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 — CRESSET 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 less 1-4 as follows: 





eg 1 
exp. 


end. 




leg 2 
exp. end. 


legs 
exp. end. 


legi 
exp. end 


seg. 1 


1:0 


0:0 




1:1 0:1 


1:1 0:0 


1:0 0:1 


seg. 2 


IV:3 


3 




1:1 0:2 


IV: 6 0:2 


VI III 


seg. 3 




III: 


5 


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 tliroughout 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 membei's 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. paradoxus.] 

Female. — 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 maxilhpeds. 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. 

Male. — Body fonn 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. IVIales 
of this genus are poorly known. 

Discussion. — 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. D. 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 

strings hidden latus 

Demoleus heptatus (Otto, 1821) 

Figures 218-237 

Caligus heptatus. — Otto, 1821, p. 15. 
Caligus paradoxus 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 — CRESSET 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 heptaiv^. — DoUfus, 1943, p. 1. — Yamaguti, 1963, p. 115. 

Specimens studied, — Eleven females and 1 male USNM 60465 
from Hexanchus species from Monterey Bay, California. 

Female. — 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 Hne 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 as in 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 
exp. end. 


leg S 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


seg. 2 


IV:3 3 


IV:5 8 


IV:5 6 


IV:5 5 



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. 



48 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Egg strings folded to form 3 strands. 

Male. — 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 
knoAvn 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 acutiinnnis. 

Female. — 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 
1 -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 Z leg S leg i 

exp. end. exp. end. exp. end. exp. end. 

seg. 1 1:0 0:0 1:1 0:1 1:1 0:1 1:1 0:1 

seg. 2 IV:3 3 IV:5 8 IV:5 5 111:5 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. 

Male.^ — -Unknown. 

Discussion. — 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, 19G4, p. 285. [Type-species: P. tunica.] 

Female. — -Fi'ontal 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. 

Male. — 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 Be, Madagascar, and the remaining paratypes 
in the author's collection. 

Other specimens studied. — A single collection from Alopias 
superciliosus from Nosy Be, Madagascar; a single collection from the 
same host in the Pacific (0°38'N, 124°23'W). 

Female. — 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 comers 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 
1.5 mm. 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 — CRESSET 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. Fom' 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 25 1 . 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 
heavOy 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 : 





Ugl 
exp. end. 


eg i 
exp. end. 


legs 
exp. end. 


leg A 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:i 


seg. 2 


111:4 3 


1:1 0:2 


1:1 0:2 


1:1 0:2 


seg. 3 


- 


111:5 6 


111:5 4 


111:5 3 



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 hahs. 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, 

Male. — 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. Fom'th 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 sliown 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 a chela at tip. When chela is closed, 
claw of last segment fits into a bifm'cation 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 siu-face 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 — CRESSET 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- 
lilie 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 unhke all other known genera of this family in having 
the rami of legs 2-4 3-segmented. 

Genus Echthrogaleus Steenstrup and Liitken, 1861 

Dmemowra.— Guerin-Meneville, 1837, pi. 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: E. coleop- 

tratus.] 
Pandarus.— Thomson, 1889, p. 363. [Refers to P. armatus only.] 

Female. — 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. 

Male. — 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. E. 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. m 

Key to Females of Genus Echthrogaleus 

1. Posterior border of dorsal thoracic plate smooth. Leg 5 concealed in dorsal 

view coleoptratus 

Posterior border of dorsal thoracic plate serrate 2 

2. Leg 5 visible in dorsal view. Dorsal thoracic plate covering not more than 

H genital segment denticulatus 

Leg 5 concealed in dorsal view. Dorsal thoracic plate covering at least % 
genital segment torpedinis 

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. 13; 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. 139.— 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 lutkenii Norman, 1868, p. 300. 

Echthrogaleus perspicax Olsson, 1868, p. 18. — Wilson, 1907, p. 457. 

Echthrogaleus lutkenii. — 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 glaiica 
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 
literatm'e. The female has been well described and figm-ed (most 
recently by Shiino, 1954) ; however, the male is poorly known and 
will be dealt with here in more detail. 

Female. — 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). Fii-st maxilla as in figure 269. Adhesion 
pads present but small. 

Leg 1-4 biramose with spine and setal formula as follows: 





leg 1 
exp. end. 


legZ 
exp. end. 


legs 
exp. end. 


leg A 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


X IV 


seg. 2 


IV:3 3 


1:0 8 


1:1 6 




seg. 3 




111:5 


111:5 





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. 

Male. — 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: 





legl 
exp. end. 


legs 
exp. end. 


legs 
exp. end. 


leni 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


seg. 2 


IV:3 3 


1:1 8 


1:1 11:6 


1:1 5 


seg. 3 




111:5 


111:5 


11:5 



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 coleo'ptratus is a widely distributed 
species of copepod and has been reported from a wide variety of 
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 Nakamiira from Majimga, Madagascar. Three collections 
from Alopias vulpinus from the Indian Ocean Cl6°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°37'W; 11°15'N, 113°26'W). 

Female. — 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 re(;ently redescribed and figm^ed 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: 





legl 
esp. 


end. 


legs 
exp. 


end. 


legs 
exp. end. 


legi 
exp. end 


seg. 1 


1:0 


0:0 


1:1 


0:1 


1:1 0:1 


X VI 


seg. 2 


IV: 3 


3 


1:1 


0:2 


1:1 6 




seg. 3 






IV:4 


6 


111:5 





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. 

Male. — 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 218/x long. Second segment 360^ long. 
Caudal rami long, measuring 720/x by 144ju (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: 





I 
exp. 


end. 


leg 2 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg, 1 


1:0 


0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


seg. 2 


IV:3 


3 


1:1 8 


1:1 11:6 


1:1 5 


seg 3 






111:5 


111:5 


111:4 



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 

Figures 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. 

Female. — 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 6.2 mm). Dorsal thoracic plates on segment 4. These 
plates very conspicvious 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 inner 
margins. 

Oral area w^ith adhesion pads reduced. Pad associated with maxU- 
liped in form of a posteriorly directed process. Oral appendages like 
those of E. denticulatus. 

Legs 1-4 (fig. 293) biramose with spine and setal formula as follows : 





legl 
exp. end. 


legi 
exp. 


end. 


legs 
exp. end. 


legi ^ 
exp. end 


seg. 1 


1:0 0:0 


1:1 


0:1 


1:1 0:1 


IX IV 


seg. 2 


IV:3 3 


1:1 


7 


1:1 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 E. denticalatus. 

Color in preserved specimens cream tan, devoid of heavy pigmen- 
tation. 

Egg strings long and straight. 

Male. — Unknown. 

Discussion. — This species seems closely related to E. 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 — CRESSET 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: A''. orientaUs.] 
Nogagus. — Beneden, 1892b, p. 246. [Refers to N. augustatus only.] 

Female. — 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 bu-amose. 
Rami of legs 1-3 2-segmented, those of leg 4 1-segmented. Leg 5 
reduced to 1 or 2 setae. Leg 6 absent. Egg strings straight. 

Male. — 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 Nesijj-pus that are now certainly not members of this genus: 
Nesippus curticaudis Dana, 1852, N. borealis Steenstrup and Lutken, 
1861, and A^. 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 

1. Endopod of leg 4 unarmed 2 

Endopod of leg 4 with long se'lac 3 

2. Genital segment conspicuously narrowed anteriorly', caudal rami with promi- 

nent setae crypturus 



60 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Genital segment not conspicuously narrowed anteriorly, caudal rami with 

small reduced setae tigris 

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 Thomson, 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 
Be, 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 (measm'ed 
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 pliunose. 

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. 



3. 3570 PANDARIDAE — CRESSET 61 

Legs 1-4 biramose with spine and setal formula as follows: 





leg 1 
exp. end. 


legH 
exp. end. 


legs 
exp. end. 


leg J, 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:0 0:1 


V:4 4 


seg. 2 


IV:3 3 


IV:5 7 


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. 

Male.— 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 
gUl 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 shoidd 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 occullus 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 Carcharinus galapagensis. From 
Nosy Be Madagascar: Sphyrna lewini (4 collections), Carcharinus 
maculipinnis (8 collections), C. leucas (4 collections), Scoliodon 
palasorrah (1 collection), and Galeocerdo cuvier (1 collection). 

Female. — Body form as in figure 305. Total length 6.8 mm (based 
on an average of 5 specimens). Greatest width 3.5 mm (measm*ed 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 
exp. 


end. 


leg 2 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 


0:0 


1:1 0:1 


1:1 0:1 


V 


Beg. 2 


IV:3 


3 


IV:5 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 joint. 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. 

Egg strings long and straight. 

Color in life cream. 

Male. — ^Body form as in figure 316. Total length 5.4 mm (based 
on an average of 2 specimens). Greatest w^idth 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 sub terminal. Each ramus joined dis tally 
to abdomen. 

Oral area with adhesion pads as m 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) 
bii-amose with spine and setal formula as follows: 





le 
exp. 


end. 


legSi 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 


0:0 


1:0 0:1 


1:1 0:1 


IV:4 4 


seg. 2 


IV:3 


3 


IV:5 7 


IV:4 4 





Leg 5 (fig. 324) located near outer distal corner of genital segment 

I 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. 

I The copepod was originally described by Heller in 1865 and has 
since been redescribed as A^. occidtus, N. costatiis, and A^. gracilis by 
C. B. Wilson (1924; 1924b; 1935). I have examined Wilson's types 
of A'', gracilis and determined this to be the same as N. cryptarus. 
Nesipjpus occultus from the pubhshed description also appears to be 
synonymous. Nesippus costatus was described from an immature 
female and was collected together with the material described as 
A^. occultus by Wilson. It appears to be merely a young female of 
the type described as N. 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 
maxiiUped should separate this sex from other species of the genus. 

Nesippus tigris, new species 

Figures 325-345 

Specimens studied. — Twelve females and 4 males from Galeocerdo 
cuvier from Sarasota, Fla. Holotype female, allotype male, 3 para- 
types (299, Icf) 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 Be, 
Madagascar (699). 

Female. — 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 segment 
with a median sinus posteriorly. Abdomen (fig. 326) 1-segmented. 
Caudal rami (see fig. 326) 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-segmented. 

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 du'ected 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 maxUla (fig. 331) with a short terminal claw 
bearing rows of fringe. The penidtimate 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 bu'amose with spine and setal formula as follows: 





legl 
exp. end. 


leg 2 
exp. end. 


legs 
exp. end. 


legi 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


IV 


seg. 2 


IV:3 3 


IV:5 7 


IV:4 4 





Leg 1 (fig. 334) with both rami 2-segmented. Setae constricted 
near their bases as in A'^. 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 Avith 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-segmented. Exopod with 4 terminal 
spines. Endopod unarmed. Leg 5 a single seta located on ventral 
sm'face of genital segment near posterior corner (see fig. 326). 

Egg strings long and straight. 

Color in life cream. 

Male. — Body form as in figure 338. Total length 5.7 mm (based 



NO- 3570 PANDARIDAE — CRESSET 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 hau's. 

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: 

^^Ol legs legs Ug jt 

exp. end. exp. end. exp. end. exp. end. 

seg 1 1:0 0:0 1:1 0:1 1:1 0:1 IV-3 4 

seg2 IV :3 3 IV:5 7 IV:5 4 

Legs 1 and 2 as in female. Leg 3 ^vith a modification on last endopod 
segment as in figm-e 343. This is consistent Avith group II males, 
although this modification is much reduced in this species. Leg 
4 (fig. 344) mth 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 Be 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 N. crypturus but can be separated from it on the basis of the 
caudal^ rami and by the fact that in A^. crypturus the genital seg- 
ment is considerably narrowed anteriorly whereas in A^. 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 Be, Madagascar. 

Female.— Body form as in figm-e 346. Total length 3.8 mm (based 
on an average of 2 spechnens). Greatest width 1.5 mm (measured at 
widest part of cephalon). Cephalon rounded, slightly wider than 
long (1.2 by 1.5 mm). Thoracic segments 2 and 3 indistinctly divided 

221-534—67 5 



QQ 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) . Fhst 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 m other 
species of genus. Mouth tube and mandible of usual pandarid type. 
First maxilla (fig. 351) with a short spme and a small patch of setules 
near base of the claw. Claw with rows of frmge as in figm^e. Maxil- 
liped (fig. 352) bearmg a blunt terminal claw opposed by a crater-like 
area on opposite segment. 

Legs 1-4 bkamose with spine and setal formula as follows : 

leal leg 2 les S ^^^ ^ . 

exp. end. exp. end. exp end. exp. end. 

seg. 1 1:0 0:0 1:1 0:1 1:1 0:1 IV:3 4 

seg. 2 IV:3 3 IV:5 7 IV:4 4 

Leg 1 (fig. 353) with both rami 2-segmented. Fhst exopod segment 
wiSi an outer spme. Last exopod segment with 4 outer spines and 
3 mner setae. Fhst endopod segment unarmed. Last endopod 
segment with 3 terminal setae. Leg 2 (fig. 354) with both rami 
2-segmented. First exopod segment with an outer spine and an 
mner seta. Last exopod segment with 4 outer spines and 5 inner seta. 
Fu-st endopod segment with an inner seta. Last endopod segment 
with 7 terminal setae. Leg 3 (fig. 355) with each ramus 2-segmented. 
Fhst segment of the exopod with an outer spine and an inner seta. 
Last exopod segment with 4 outer spines and 4 mner setae. Fhst 
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 pomt. 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. 

Discussion.— This copepod has been collected only from Bhyn- 
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 — CRESSET 67 

This species can be separated from A^^. orientalis by the lack of 
the dorsal plate on fourth segment. It can be separated from A^. 
crypturus by the nature of adhesion pads of the cephalon and the 
nature of the fourth leg of the female. It differs from A^. tigris by 
the nature of the fourth thoracic segment (wide in A^. tigris, narrow 
in N. vespa) and also by the nature of the caudal rami. 

Genus Paranesippiis 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 Acanthidium 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 : 





legl 
exp. end. 


leg S 
exp. end. 


legs 
exp. end. 


leg 4 
exp. end. 


seg. 1 


1:0 0:0 


1:1 0:1 


1:1 0:1 


1:1 0:1 


seg. 2 


IV:3 3 


1:1 0:2 


1:1 0:2 


1:1 5 


seg. 3 




111:5 6 


111:4 4 


111:4 



It is interesting to note that the egg strings remain recm'ved and 
hidden beneath the genital segment as in some other members of group 
II {Demoleus 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 

era], 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 configm'ation 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, Dinemoum, 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. 



PAND ARID AE — CRE SSE Y 



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 








Mspd 


Mxpd 


corner of 
cephalon 


Pandarus 


+ 


+ 


+ 





+ 


Pseudopandarus 


+ 


+ 


+ 


— 


+ 


Perissopus 


+ 


+ 


+ 


— 


— 


Paranesippus 


+ 


+ 


+ 


— 


— 


Pannosus 


+ 


+ 


+ 


— 


— 


Nesippus 


hooklike 


+ 


+ 


— 


— 


Dinemoura (part) 


double 


+ 


+ 


— 


— 


Demoleus 


+ 


+ 


+ 


+ 


— 


Pagina 


+ 


— 


— 


— 


— 


Echthrogaleus 


+ 


+ 


hooklike 


— 


— 


Phyllothereus 


— 


— 


— 


— 


— 


Dinemoura (part) 


— 


— 


— 


— 


— 


Gangliopus 


— 


— 


+ 




" 



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, ^vithin 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 dm'ing 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 siu-face 
of the host attaches by cementing the maxilliped to the host. The 2 
genera of group I found on the giUs 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 3 setae. 

Leg 2: In group I the exopod of all species is 2-segmented. In 
group II 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, ^^'ith only 2) have 3- 
segmented exopods. Copepods of group I have a 2-segmented 
exopod except Perissopus, which has only 1. 



NO. 3570 PANDARIDAE — CRESSET 71 

Leg 4: In many genera of both groups this leg in the female is 
lamelliform {Pandareus, Phyllotherus, Echthrogaleus, and Dinemourd). 
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 w^here 
only the smaller ones were present that these might be erroneously 
considered as matiu-e. 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 Avas also present (see figs. 
59, 60). 



72 



PROCEEDINGS OF THE NATIONAL MUSEUM 



A List of Sharks Examined with the Pandarid Copepods Recovered 

from Them 

(Number in parenthesis indicates collections made) 



Hexanchus species 

Denioleus paradoxus (1) 

Isurus oxyrhynchus 

Pandarus smithii (3) 
Dinemoura produda (1) 
Dinemoura latifolia (3) 
Echthrogaleus denticalatus (3) 

Lamna nasus 

Pandarus floridanus (1) 
Echthrogaleus coleoptratus (1) 

Lamna ditropis 

Echtrogaleus coleoptratus (1) 

Carcharodon carcharias 
Pandarus smithii (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 (3) 

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 cranchii (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 smithii (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 pxjriformis (4) 

Dinemoura producta (1) 
Galeocerdo cuvier 

Pandarus cranchii (2) 

Nesippus tigris (2) 

Nesippus orientalis (2) 

Nesippus crypturus (2) 



PANDARIDAE — CRE SSE Y 



73 



Negaprion breviroslris 

Pandarus sinuatus (1) 

Galeorhinus species 

Pandarus niger (1) 
Pandarus car char ini (1) 

ScoUodon palasorrah 

Pseudopandarus gracilis (1) 
Nesippus orientalis (1) 
Nesippus cryplurus (1) 

Rhizoprionodon acutus 

Pseudopandarus longus (1) 

Sphyrna mokarran 

Nesippus cryplurus (1) 



Sphyrna lewini 

Nesippus orienlalis (4) 
Nesippus cryplurus (4) 

Sphyrna zygaenae 

Pandarus cranchii (1) 
Pandarus smithii (1) 
Pandarus zygaenae (3) 
Nesippus orienlalis (1) 

Squalus aculipinnus 
Demoleus lalus (1) 

Squalus acanlhias 

Pandarus bicolor (1) 

Rhynchobalus djeddensis 
Nesippus nana (1) 



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Copepoda. Monogr. 29b (vol. 123) in Weber, Siboga-Expeditie, 
40 pp. 
1934b. A third list of parasitic Copepoda of Plymouth with notes. Para- 
sitology, vol. 26, no. 1, pp. 112-113. 
Lewis, A. G. 

1963. Life history of the caligid copepod Lepeophtheirus dissimulatus Wilson, 
1905 (Crustacea Caligoida). Pacific Sci., vol. 17, no. 2, pp. 195- 
242. 
Markewitsch [Markevich], a. p. 

1957. Parasitic Copepoda of fish of USSR, 259 pp. [In Russian; reference 
not seen.] 
Marukawa, H. 

1925. Illustrated encyclopedia of the fauna of Japan. [Reference not 

seen.] 
1947. Revised edition of illustrated encyclopedia of the fauna of Japan, 
exclusive of Insecta. [Reference not seen.] 
Matthews, L., and Parker, H, W. 

1950. Notes on the anatomy and biology of the basking shark. Proc. 
Zool. Soc. London, vol. 120, pp. 535-576, 15 figs., 8 pis. 
McClendon, J. F. 

1906. On the development of parasitic copepods, 1 and 2. Biol. Bull., vol. 

12, no. 1, pp. 37-52; no. 2, pp. 53-88. 

1907. The spermatogenesis of Pandarus sinuatus Say. Biol. Bull., vol. 13, 

pp. 114-119. 
1910. Further studies on the gametogenesis of Pandarus sinuatus. Arch. 

Zellf., vol. 5, no. 2, pp. 229-234, 1 illustr., 1 tbl. 
MiERS, E. J. 

1880. On a small collection of Crustacea made by Edward Whymper, Esq., 

chiefly in the N. Greenland Seas. Journ. Linn. Soc, vol. 15, 

pp. 59-73. 
Milne-Edwards, H. 

1840, Histoire naturelle des Crustacea, comprenant I'anatomie, la physiologie 

et la classification de ces animaux, 638 pp., 42 pis. 



78 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

MoNOD, T., and Dollfus, R. 

1938. Pandarins pen connus (Generes Phyllothyreus Norman 1903 et 
Gangliopus Gerstaecker 1854). Ann. Parasit. Hum. Comp., vol. 
16, no. 3, pp. 196-209. 
MULLER, O. F. 

1785. Entomostraca, sou Insecta testacea quae in aquis Daniae et Norvegiae 
reperit, descripsit, et inconibus illustravit, 134 pp. [Reference not 
seen.] 
NoRDMANN, Alexander von 

1832. Mikrographische Beitrage zur Naturgeschichte der wirbellosen 
Thiere, xvi+150 pp., 10 pis. 
Norman, A. M. 

1868. Last report on dredging among the Shetland Isles. In Report of the 
British Association for the Advancement of Science for 1868, 
p. 301. 
1903. New generic names for some Entomostraca and Cirripedia. Ann. 
Mag. Nat. Hist., ser. 7, vol. 11, pp. 367-369. 
Norman, A. M., and Brady, G. S. 

1910. The Crustacea of Northumberland and Durham. Trans. Nat. Hist. 
Soc. Northumberland, Durham and Newcastel-upon-Tyne, n. s., 
vol. 3, no. 2, pp. 252-417. 
Norman, A. M., and Scott, T. 

1906. Crustacea of Devon and Cornwall, 232 pp. 
NuSfES-Ruivo, L. 

1956. Copepodes parasitas de peixes dos mares de Angola. Anais Junta 
Investig. Ultramar., vol. 9, no. 2 (1954), pp. 9-45. 
Olsson, p. 

1868-69. Prodromus faunae copepodorum parasitantium Scandinaviae. 
Acta Univ. Lundensis, no. 8, 49 pp., 2 pis. 
OoRDE DE Lint, G. M. van, and Schuurmans Stekhoven, J. H., Jr. 

1930. Copepoda parasitca. Tierw. Nord-Ostee (Gimpe und Wagler), 
no. 31, pp. 73-198. 
Otto, A. W. 

1821. Conspectus animalium quorundam maritimorum nondum editorum 
. . ., 20 pp. 
Otto, A. W. 

1828. Beschreibung einiger neuen, in den Jahren 1818 und 1819 im mittel- 
landischen Meer gefundener Crustaceen. Nov. Act. Acad. 

Leopold-Carol., vol. 14, no. 1, pp. 331-354, pis. 20-22. 
Pearse, a. S. 

1952a. Parasitic Crustacea from the Texas coast. Inst. Mar. Sci. Port 

Arkansas, Texas, vol. 2, no. 2, pp. 5-42. 
1952b. Parasitic crustaceans from Alligator Harbor, Florida. Quart. Journ. 
Florida Acad. Sci., vol. 15, no. 4, pp. 187-243. 
Pearson, J. 

1905. A list of the marine copepoda of Ireland, 1: Litoral forms and fish 
parasites. Rep. Sea and Inl. Fish. Ireland (1904), pt. 2, pp. 143- 
170. 
Pesta, Otto 

1934. Krebstiere oder Crustacea, 1: Ruderfiisser oder Copepoda, 6: 
Caligoida. Pt. 29 in Dahl, Die Tierwelt Deutschlands, 68 pp. 



PANDARIDAE — CRESSEY 



79 



Rathbun, 

1884. 



1886. 



1887. 



POCHE, F. 

1902. Bemerkimgen zu der Arbeit des Hcrrn 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 Chondr acanthus. Proc. U.S. Nat. Mus., vol. 9, pp. 310-324. 

Description of new species of parasitic copepods belonging to the 
genera Trebius, Perissopus, and Lernanthropus. Proc. U.S. Nat. 
Mus., vol. 10, pp. 559-571. 

RiCHIARDI, S. 

1880. Contribution alia fauna d'ltalia: Catalogo sistematico dei crostacei 

che vivono sul corpo degli animali aquatici. In Catalogo degli 

Espositori, pp. 147-152. 
Rose, M., and Vaissiere, R. 

1952a. Catalogue pr^liminaire des cop6podes de I'Afrique du Nord, 1. 

Soc. Hist. Nat. Afrique Nord, vol. 43, pp. 113-136. 
1952b. Catalogue pr^liminaire des copepodes de I'Afrique du Nord, 2. 

Soc. Hist. Nat. Afrique Nord, vol. 43, pp. 164-176. 
Catalogue preliminaire des copepodes de I'Afrique du Nord, 3. 

Soc. Hist. Nat. Afrique Nord, vol. 44, pp. 83-99. 



1953. 



Bull. 



Bull. 



Bull. 



Say, T. 
1817. 

Scott, A. 
1904. 

1929. 

Scott, T. 
1900. 



An account of the Crustacea of the United States. 
Nat. Sci. Philadelphia, vol. 1, 458 pp. 



Journ. Acad. 



Some parasites found on fishes in the Irish Sea. Trans. Biol. Soc. 

Liverpool, vol. 18, pp. 33-45. 
The copepod parasites of Irish Sea Fishes. Rep. Lancahire Sea-Fish. 

Lab., no. 37, pp. 81-118. 



18th Ann. Rep. Fish. 



1901. 



1902. 



1904. 



Notes on some crustacean parasites of fishes. 

Bd. Scotland, pp. 144-188. 
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. 
Rep. Fish. Bd. Scotland, pp. 275-278. 
Scott, T., and Scott, A. 

1913. The British Copepoda, 1: Copepoda parasitic on fishes. Ray Soc. 
London, vol. 2, 72 pis., 256 pp. 
Shen, C. J., and Wang, K. N. 

1958. A new parasitic copepod, Achtheinus impcnderus (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 
summary.] 



80 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Shiino, S. 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 oflf Kesennuma, Miyagi Prefecture. Rep. Fac. Fish. Pref. 
Univ. Mie, vol. 2, no. 3, pp. 359-375. 
1959a. Ostpazifische parasitierende Copepoden. Rep. Fac. Fish. Pref. 

Univ. Mie, vol. 3, no. 2, pp. 267-333. 
1959b. Sammlung der parasitischen Copepoden in der Prafekturuniversitat 
von Mie. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 2, pp. 
334-374. 
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, 
Japan. Rep. Fac. Fish. Pref. Univ. Mie, vol. 3, no. 3, pp. 471-500. 
Smith, S. I. 

1874. Invertebrate animals of Vineyard Sound. Rep. Comm. Fish and 
Fisher. 1871 and 1872, 478 pp., 38 pis. 
Stebbing, T. R. R. 

1910. General catalogue of South African Crustacea. Ann. South African 
Mus., vol. 6, no. 5, pp. 281-593. 
Steenstrup, J. J. S., and LiItken, 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. 
Stephenson, K. H. 

1940. Parasitic and semiparasitic Copepoda. Zool. Iceland, vol. 3, no. 34, 
pp. 1-24. 
Stuardo, J., and Fagetti, E. 

1961. Copepodos parasitos chilenos, 1: Una lista de las especies conocidas 
y descripcion de tres especies nuevas. Rev. Chilena Hist. Nat., 
vol. 55, pp. 55-82. 
Thomsen, R. 

1949. Copepods parasitos de los pesces marinos des Uruguay. Comm. 
Zool. Mus. Hist. Nat. Montevideo, vol. 3, no. 54, pp. 1-41. 
Thomsen, G. M. 

1889. Parasitic Copepoda of New Zealand. Trans. New Zealand Inst., 
vol. 22, pp. 353-376. 
Valle, a. D. 

1880. Crostacei parassiti dei pesci del mare Adriatico. Boll. Soc. Adriat. 
Sci. Nat., vol. 6, pp. 55-90. 
Wilson, C. B. 

1905. New species of parasitic copepods from Massachusetts coast. Proc. 
Biol. Soc. 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, pis. 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. 



NO. 3570 PANDARIDAE — CRESSEY 81 

Wilson, C. B. 

1911. North American parasitic copcpods: 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. CoU., vol. 91, no. 3298, art. 19, 

pp. 1-9. 
1935b. Parasitic copepods from the Pacific Coast. American Midi. Nat., 

vol. 16, no. 5, pp. 776-797. 
1936. Parasitic copepods from the Dry Tortugas. In Papers from the 

Tortugas Laboratory. Carnegie Inst. Washington Publ. 452, 

vol. 29, no. 12, pp. 327-347. 
Yamaguti, S. 

1936. Parasitic copepoda from fishes of Japan, 3: Caligoida, 2, 21 pp. 
1963. Parasitic copepoda and Branchiura of fishes, 390 pp. 
Yamaguti, 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 




Figures 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). 



PANDARIDAE — CRESSET 



83 




Figures 9-15. — Pandarus satyrus, female: 9, second maxilla (D); 10, maxilliped (D); 11, 
leg 1 (D); 12, leg 2 (D)j 13, leg 3 (F); 14, leg 4 (F); 15, leg 5 (G). 



84 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figures 16-23— P and ar us 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). 



PANDARID AE — CRE SSE Y 



85 




Figures 24-30.~Pandarus satyrus, male: 24, leg 4 (F); 25, leg 5 (G); 26 leg 6 (E). P. 
cranckii, 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 




Figures 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). 



PANDARIDAE — CRESSET 



87 




Figures 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 



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Figures 44-S0.—Pandarus smithii, male: 44, leg 2 (D); 45, leg 5 (G). P. floridanus, 
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). 



P ANDARID AE — CRE SSE Y 



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- 



90 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 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). 



PANDARIDAE CRESSEY 



91 




Figures 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 




Figures 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). 



PANDARIDAE — CRESSET 



93 




Figures 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 




Figures 911-99.— Pandarus katoi, new species, male: 93, leg 3 (D); 94, leg 4 (D); 95, leg 5 
(E); 96, leg 6 (E). P. zygaenae, female: 97, dorsal (A); 98, maxilliped (G); male: 99, 
dorsal (B). 



PANDARIDAE — CRESSET 

100 



95 




Figures lOO-m.-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 




Figures 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). 



PANDARID AE — CRE SSE Y 



97 




Figures 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. m 

\ 



126 




130 



Figures 124-130. — Pandarus carcharini, female: 124, leg 4 (F). Phyllother/rus 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 
(F). 



PANDARIDAE — CRESSET 



99 




Figures Ul-Ul.—Phyllothereus cornutus, female: 131, leg 1 (F)- 132 leg 2 (C)- 133 
leg 3 (C); 134, leg 4 (C); 135, area of leg 5 (F). Male: 136, dorsal (A); 137, leg 5 (G)'. 



100 



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Figures 13S-U3 —Phyllotlureus cornutus, male: 138, area of leg 6 (G). Gangliopus 
pyriforniis, 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). 



PANDARIDAE — CRESSEY 



101 




Figures li4r-lS0.—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 




Figures lSl-\S6.—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). 



PANDARIDAE — CRESSET 



103 




157 



Figures lS7-l63.—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). 



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Figures \6^m.-Perissopus deniatus, 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). 



PANDARIDAE — CRESSEY 



105 




Figures 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— t— 8 



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PROCEEDINGS OF THE NATIONAL MUSEUM 



178 180 



177 




Figures l77-l&2.—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). 



PANDARID AE — CRE SSEY 



107 




Figures 183-188.— PmV/o^uj- dentatus, male, 183, leg 3 (E); 184, leg 4 (E); 185, leg 5 (H); 
186, leg 6 (H). Female: 187, dorsal (C); 188, dorsal (C). 



108 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 189-191. — Perissopus dentatus, female: 189, dorsal (C). Dinemoura produda, 
female: 190, dorsal (A); 191, oral area (B). 



PAND ARID AE — CRE SSE Y 



109 




Figures 192-193.— Dinemoura ferox, female: 192, dorsal (J); 193, ventral fj). 



no 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figures 194-196. — Dinemoura lalifolia, female: 194, dorsal (A); 195, oral area (B); 196, 

area of legs 5 and 6 (F). 



P AND ARID AE — CRE SSE Y 



111 




t Figures 197-200. — Dinemoura discrepans, new species, female: 197, dorsal (A); 198, ventral 
(A); 199, first antenna (D); 200, second antenna (F). 



112 



PROCEEDINGS OF THE NATIONAL MUSEUM 



201 




Figures 20\-2Q^. -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 
Tecond mTxiua (i); 205, maxillipcd (F); 206, leg 1 (F); 207. leg 2 (F); 208, leg 3 (F). 



PANDARIDAE — CRESSET 



113 




Figures 209-216. — Dinemoura discrepans, new species, female: 209, area of legs 5 and 6 
(F); 210, egg string (no scale) (lOX). Male: 211, dorsal (A); 212, genital segment, ventral 
(C); 213, endopod of leg 3 (G); 214, leg 4 (G); 215, leg 5 (H); 216, leg 6 (E). 



114 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figure 217-221.— Dinemoura disirepans, new species, male: 217, attachment area of 
spermatophore (E). Demoleus heptaius, female: 218, dorsal (A); 219, abdomen and 
caudal rami (C); 220, oral area (A); 221, process between maxillipeds (D). 



P ANDARIDAE — CRE SSE Y 



115 




Figures 221-21%.— Demoleus fuptatus, female: 222, first antenna (D); 223, second antenna 
(D); 224, first maxilla (G); 225, claw of second maxilla (G); 226, maxilHped (F); 227, 
leg 1 (D); 228, leg 2 (D). 



116 PROCEEDINGS OF THE NATIONAL MUSEUM 

232 




233 



Figures 229-m.-Demoleus heptatus, female: 229, leg 3 (D); 230, leg 4 (D); 231, leg 5 (G); 

232, leg 6 (G). 



P ANDARID AE — CRE SSE Y 



117 




Figures 235-242. — Demoleus heptaus, 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 




Figures 243-247. — Pagina tunica, female: 243, dorsal (A); 244, lateral (A); 245, caudal 
ramus (C); 246, oral area, ventral (D); 247, first antenna (E). 



J 



PANDARIDAE — CRESSET 



119 




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 (E). 



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PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figures 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). 



f 



PANDARIDAE — CRESSEY 



121 




Figures 260-267. — Pagjtia tunica, male: 260, dorsal (A); 261, genital segment and abdo- 
men, ventral (C); 262, second antenna (H); 263, maxllliped, 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 




269 



270 



Figures 268-27S.—EchthTOgaleus coleoptratus, female: 268, dorsal (A); 269, first maxilla 
(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). 



PANDARIDAE — CRESSET 



123 



I 




Figures 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). E. denticulatus, female: 281, dorsal 
(B); 282, second maxilla (G). 



124 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figures 2S3-290.—Echthrogaleus denticulaius, 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 (H). 



PANDARIDAE — CRE SSE Y 



125 




Figures 291-29S. —Eckthrogaleus 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. j2i 




Figures 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). 



PANDARID AE — CRE SSE Y 



127 




Figures 303-308.— Nesippus orievidis, female: 303, leg 3 (G); 304, leg 4 (G). A^. cryp- 
tuTus, 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 vol. 121 

315 



309 




Figures 309-3 IS. —Nesippus cryplurus, female: 309, first maxilla (E); 310, maxUliped 
(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 (I). 



PANDARIDAE — CRESSEY 

320 318 



129 




Figures 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, leg4(K);324, leg5(H). 



130 



PROCEEDINGS OF THE NATIONAL MUSEUM 

'7^ 




329 



Figures 32S-331. —Naippus 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). 



PANDARIDAE — CRESSET 



131 



) , 336 




Figures 332-339.— Nesippus tigris, new species, female: 332, maxiUiped (D); 333, adhesions 
pad at base of maxiUiped (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 




342 



Figures SiO-diS.—Nesippus tigris, new species, male: 340, maxlUiped (G); 341, tip of 
maxiUiped claw (H); 342, adhesion pad at base of maxiUiped (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). 



PANDARID AE — CRE SSE Y 



133 




Figures 349-356.-^m/./,«. 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). • s v y. 



U.S. GOVERNMENT PRfNTING OFFICE: 1967 



Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION . WASHINGTON, D.C. 




Volume 121 1967 



]N umber 3571 



SUPPLEMENTARY DESCRIPTIONS 

OF TWO MYODOCOPID OSTRACODS 

FROM THE RED SEA 



By Louis 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 spechnen 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 Natm-historisches Museum, Vienna, 
Austria: (1) One bottle, containing one vial with 3 males, with red 
label marked: ' 'Philomedes polae, n. sp. cf ; Rotes Meer: Gulf von Suez; 



1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Coll- Tola' 1895/8; Dr. Graf det." (2) One bottle with white label 
marked: 'Thilomedes sp. 9; Rotes Meer: Stat. 90; CoU. Tola' 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 "PMlomedes sp. 9 (A)." 

Smce 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 EupMlomedes 
arostratus Kornicker, recently described for the fii-st 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 pubhshed 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 hke 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. Muller, 1912 

Genus Euphilomedes Poulsen, 1962 

Type-species. — Euphilomedes nodosus Poulsen, 1962, by subsequent 
designation, Kornicker (1967). 

Euphilomedes polae (Graf, 1931) 

Figures 1-5 

Philomedes polae Graf, 1931, p. 37, fig. 5. 
Philomedes species. — Graf, 1931, p. 38, fig. 6. 

Lectotype. — Specimen designated as specimen B on slide. Natur- 
historisches Museum, Zoologische Abteilung, Vienna, Austria. 
Sex. — Male. 



RED SEA OSTRACODS — KORNICKER 




Figure \.—Euphilomedes polae, male, specimen C (except a and h, spec. B): ^, 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; e, medial view of part of shell posterior 
showing reticulations, normal pore canals and surface hairs; /, 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; /, 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 lonijitiide 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 
greatest height near middle, prominent rostrum and shallow rostral 
incisure (fig. la); anterior margin of rostrum and anteroventral 
margin of shell with scalloped outUne formed by cresent-like marginal 
denticulations; alate shield projecting laterally and anteroventrally 
from behind rostral incisure, partly covermg incism-e, flap strongly 
sclerotized anteriorly (figs. Ic,/). Sm-face of valves with kregular 
polygons; polygons with pebbly texture appearing as minute polygons 
under high magnification (figs. U,k). Posterior dorsal margin with 
linear hinge depressed below outline of shell (figs. 26,c); hinge not 
visible in lateral view except by transmitted fight. 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. la,i,j). Inner lamella broad with vestibule; no parallel 
striations or line of concrescence observed on inner lamella. Selvage 
with wide, corrugated, lamefiar prolongation having fringe of slender 
spines along outer margin. SLx long haks bearing secondary spines, 
forming row on inner lameUa behind rostrum (figs. lg,h); inner lamella 
ventral to rostral incisure with small plumose hair, followed by wide 
space and then about 5 plumose hau's (fig. 2a); about 30 haii's on 
postero ventral part of inner lamella (figs. 2d,e). 



RED SEA OSTRACODS — KORNICKER 




^jf fif' 



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; e, medial view posteroventral and 
posterodorsal margin showing medial hairs; /, lateral view of right 1st 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; {, protopodite and endopodite of 2nd antenna;;, medial 
view 1st 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,h,e,f,g,i; c,d,j,k; h,l.) 



6 PROCEEDESTGS OF THE NATIONAL MUSEUM vol. 121 

Hairs with either pointed or blunt ends distributed on rostral surface 
(fig. 16), some forming row near margin of shell; slender haii-s with 
pointed ends on posterior surface of shell, some forming row near pos- 
terior margin (fig. Id); 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. 2/): 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 hau-s and the 
more proximal bristle with short hairs distally, and 4 long ventral 
bristles sub terminally, 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. 2i). Exopodite (fig. 2g): 1st joint elongate; 2nd 
joint about K 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 K 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. 2h,i); 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-l, 3a, b) : No coxale endite observed. Basale 
(fig. 36): ventral margin with 3 short slender spinous bristles, and 2 



RED SEA OSTRACODS — KORNICKER 




Figure 3. — Euphiloviedes polae, male, specimen B: a, medial view of exopodite and 1st 
endopodite joint without distal bristles of left mandible; b, 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; e, endites and exopodite of 5th limb, all bristles not shown; 
/, 6th limb; g, 7th limb (1 bristle in distal group broken); h, distal end of 7th limb; i, frontal 
organ and median eye; ;', lateral view of right furcal lamella; k, copulatory organ; /, detail 
of distal tip of copulatory organ; wz, lateral eye. (Same scale, In microns: a,e,h,l; 
b,f, i, m; c, g; d,j, k.) 



8 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

long bristles with wreaths of long hau'S and short spines distaUy. 
Dorsal margin with 3 bristles, 1 near middle and 2 subtermmaUy. 
Medial sm-face with about 5 short bristles proximally near ventral 
corner and 1 longer bristle near ventral margin at middle of jomt. 
Exopodite reaching middle of 1st endopodite joint, with 2 subeqiial 
terminal bristles with short marginal spines (fig. 3a); tip of joint with 
blunt hirsute process bearing short spine. Endopodite: Fu-st jomt 
(fig. 2j) with 1 short spinous bristle and 3 long hirsute posterior 
bristles distally. Second joint (fig. 2k) anterior margin with proximal 
group 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. 2/) 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 
onsljlGS 

Sixth limb (fig. 3/): Fu-st 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, h): 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, I) : Long, slender, divided into 3 lobes, 
each with 2 annulated bristles; 1 lobe with large curved sclerotized tip. 

Furca (fig. 3j): 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.) 



RED SEA OSTRACODS — KORNICKER 




Figure 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; b, 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; /, medial 
view of valve near posterior showing large and small reticulations; g, m_edial 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; 
i, medial view of posterior and ventral marginal area of right valve showing distribution 
of hairs on inner lamella; ;, lateral view of left 1st antenna; k, 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, I.) 

221-535—66 2 



XO 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. 3i). 

Frontal organ (fig. 'Si): Elongate, 2-jointed with short spines on 
distal half of 2nd joint and subterminally on basal joint. 

Description of female. — Shell (figs. 4a-i): 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; iiTegular polj^gons on surface of shell subdivided by 
minute polygons larger than on male shell and visible under low 
magnification (fig. 4/). Inner lamella with parallel striations on 
an tero ventral part; 9 long hairs v>'ith secondary spines forming row 
on inner lamella behind rostrum (figs. 46,c) ; inner lamella below rostral 
incisure with small hair followed by wide space and then 5 hau's (fig. 
4e); posteroventral part of inner lamella with at least 21 hairs (fig. 4i); 
lateral surface of rostrum and anterior part of shell with numerous 
hairs with blunt ends (fig. 4^0- 

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. 4J): Fu'st and 2nd joints with siu'face hairs. 
Second joint with 3 bristles, 1 dorsal, 1 ventral, and 1 lateral. Third 
joint witli 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. 4Ar, I) : Protopodite subtriangular without 
transparent rim. Exopodite (fig.4Z) 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 natatorj^ hairs or marginal 
denticulations; 4th to 8th joints with bristles having natatorj^ 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. 4k,l) 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 wi.th small spine at lip and long stout 
terminal bristle with wreaths of long hairs near middle and short 
distal spines. 



RED SEA OSTRACODS — KORNICKER 



11 




Figure S.—Euphilomedes polae, female, specimen 1: a, medial view right mandible (surface 
hairs not shown); b, lateral view left m.axilla, all bristles not shown; c, posterior view left 
5th limb; d, medial view right 6th limb; e, 7th limb;/, distal end of 7th limb; g, distal end 
frontal organ; A, frontal organ and medial eye; i, medial view right lamella of fiirca; 
;, 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 

]\Iandible (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. 

Alaxilla (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; 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 v/ith 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 1st exopodite joint about 
same size as distal tip of 2nd exopodite joint. Distal anterior margin 
of 1st joint with 2 centrally located bristles. Large cm-ved 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. 

Furca (figs. 5i,j): 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 f ureal claws on 2 specimens is as follows: 

















claw number 
















1 


« 


s 


4 


5 


6 


7 


S 


9 


10 


// 


n 


13 


H 


15 16 


specimen 1 
































right 


P 


P 


s 


s 


s 


s 


P 


s 


s 


s 


s 


P 


S 


s 


S S 


left 


P 


P 


s 


s 


s 


s 


P 


s 


s 


s 


p 


s 


s 


s 




specimen 2 
































right 


P 


P 


s 


s 


s 


s 


P 


s 


s 


s 


s 


p 


s 


s 


S 


left 


P 


P 


s 


s 


s 


s 


P 


s 


s 


s 


s 


p 


s 


s 





Eyes: Large medial eye (fig. 5A); lateral eyes not observed. 

Frontal organ (figs. hg,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. 

Remarks. — 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 E. 
polae and E. arostratus are compared as follows: 



E. polae 


E. arostratus 


male female 


male female 


8-9 10-12 


7 9 


4 5 


4 5 



number of secondary furcal claws 

cleaning bristles in distal group of 7th limb 

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 E. 
arostratus. 



14 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Eiiphilomedes arostratiis Kornicker, 1967 

Figure 6 
Eiiphilomedes arostrala Kornicker, 1967, p. 2. 

HoLOTYPE.— USNM 112658. Sex: female. Male Atoll, Maldive 
Islands, Indian Ocean. 

Locality. — 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 
greatest height near middle, prominent rostrum and antennal sinus 
(figs. 6a, b, e); anterior margin of rostrum and anteroventral margin of 
shell with scalloped outHne formed by crescent-like marginal denticu- 
lations. Surface of valves with distinct irregular polygons (fig. 6^); 
polj^gons 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. 6A). 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. 66) ; inner lamella below rostrum with small hair foUowed 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: Fu-st joint with medial hairs. Second joint with 
medial hairs and, distall}^, a dorsal, ventral, and lateral bristle, all 
bristles bare or with short hau-s distally. Third joint with few medial 
hairs and 1 ventral and 2 dorsal bristles, all with short hairs, the 



RED SEA OSTRACODS — KORNICKER 



15 




Figure 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;/, endopodite of 
left 2nd antenna; g, lateral view protopodite, endopodite, and 1st 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 
scars are located below anterior hinge element; i, medial view right 6th limb (surface 
hairs not shown); /, distal part of 7th limb; k, 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,ej,h-j; c,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 hau's 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. Qf,g): Protopodite triangular without trans- 
parent rim (fig. 6g). Exopodite: first joint elongate; 2nd joint 
about Yi length of 1st; 3rd joint about % length of 1st; distal margins 
of joints 2-8 with comb of short spines; basal spines not observed. 
Fkst joint without bristle; bristle of 2nd joint about % length of 3rd 
joint (fig. 6^), 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. 6/) 3-jointed; 1st joint with 5 bare short 
bristles and 1 long distal bristle with wreaths of long hau's 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 haii-s 
and short spines distally. Dorsal margin with 3 bristles. Medial 
siu"face 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 blimt 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 annidated 
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 phimose bristles. Outer 
lobe of 3rd exopodite joint with 2 stout plumose bristles. 

Sixth limb (fig. Qi) : 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. Qj-l): 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 cm*ved 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 sm'face 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 concernmg 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 E. 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 1 1 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 

Baird, W. 

1850. The natural history of the British Entomostraca, 364 pp., 36 pis. 
[Ostracoda: pp. 138-182, pis. 18-23.] 
Graf, H. 

1931. Expedition S.M.S. "Pola" in das Rote Meer: Die Cypridimdae 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. 

MtJLLER, G. W. 

1912. Ostracoda. Ft. 31 in Das Tierreich, 434 pp., 92 figs. 

POULSEN, 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 3571 



SUPPLEMENTARY DESCRIPTIONS 

OF TWO MYODOCOPID OSTRACODS 

FROM THE RED SEA 



By Louis 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. cf ; Rotes Meer: Gulf von Suez; 

1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Coll: Tola' 1895/8; Dr. Graf det." (2) One bottle with white label 
marked: "Philomedes sp. 9; Rotes Meer: Stat. 90; Coll. Tola' 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 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 pubhshed 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 CaroHne Bartlett, who inked the fmal 
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 

Type-species. — Euphilomedes nodosus Poulsen, 1962, by subsequent 
designation, Kornicker (1967). 

Euphilomedes polae (Graf, 1931) 

Figures 1-5 

Philomedes polae Graf, 1931, p. 37, fig. 5. 
Philomedes species. — Graf, 1931, p. 38, fig. 6. 

Lectotype. — Specimen designated as specimen B on slide. Natur- 
historisches Museum, Zoologische Abteilung, Vienna, Austria. 
Sex. — Male. 



RED SEA OSTRACODS — KORNICKER 




Figure l.—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 
showmg surface hairs and some reticulations; e, medial view of part of shell posterior 
showing reticulations, normal pore canals and surface hairs; /, 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; ;, 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 Phlomedes 
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 aU, 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 coUected at 
Station 90. Data concernmg 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 
greatest height near middle, prominent rostrum and shallow rostral 
incisure (fig. la); anterior margin of rostrum and anteroventral 
margin of shell with scaUoped outhne formed by cresent-like marginal 
dentlculations; alate shield projecting laterally and anteroventrally 
from behind rostral incisure, partly covering incisure, flap strongly 
sclerotized anteriorly (figs. lc,f). Surface of valves with h-regular 
polygons; polygons with pebbly texture appearing as minute polygons 
under high magnification (figs. le,k). Posterior dorsal margin with 
Unear hinge depressed below outUne of shell (figs. 2b,c); hmge not 
visible m 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 hyahne spot situated some distance 
above others; 2 large and 4 smaller scars situated below anterior hmge 
element (figs. la,i,j). Inner lamella broad with vestibule; no parallel 
striations or Ime of concrescence observed on inner lamella. Selvage 
with wide, corrugated, lamellar prolongation having fringe of slender 
spines along outer margin. Six long hau'S bearing secondary spines, 
forming row on inner lamella behind rostrum (figs. lg,h); inner lamella 
ventral to rostral incisure with small plumose hair, foUowed by wide 
space and then about 5 plumose hah's (fig. 2a); about 30 hau-s on 
postero ventral part of inner lamella (figs. 2d,e). 



RED SEA OSTRACODS — KORNICKER 




Figure 2.—Euphiloviedes polae, male, specimen B (except c, spec. C) : a, medial view of 
antero ventral 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; e, medial view posteroventral and 
posterodorsal margin showing medial hairs; /, lateral view of right 1st 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; ;*, medial 
view 1st 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.) 



I 



6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Hairs with either pointed or blunt ends distributed on rostral sui'face 
(fig. lb), 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. Id); 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. 2/): 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 hau's. Foiu"th 
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. 2i). Exopodite (fig. 2g): 1st joint elongate; 2nd 
joint about Vz 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 ji 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. 2h,i); 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-l, 3a, b) : No coxale endite observed. Basale 
(fig. Sb): ventral margin with 3 short slender spinous bristles, and 2 



RED SEA OSTRACODS — KORNICKER 




Figure 3. — Euphilomedes polae, male, specimen B: a, medial view of exopodite and 1st 
endopodite joint without distal bristles of left mandible; b, lateral view of basale and 
exopodite of right mandible; c, maxilla, all bristles not shown; d, Sth limb, hairs not 
shown on all epipodial bristles; e, endites and exopodite of Sth limb, all bristles not shown; 
/, 6th limb; g, 7th limb (1 bristle in distal group broken); h, distal end of 7th limb; i, frontal 
organ and median eye; ;', 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,l; 
b,f,i,m; c, g; d,j,k.) 



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 siu-face 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. 2j) with 1 short spinous bristle and 3 long hirsute posterior 
bristles distally. Second joint (fig. 2k) anterior margin with proximal 
group 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. 21) 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. Sd, e) : Epipodial appendage with about 47 plumose 
bristles. Outer lobe of 3rd exopodite joint with 2 stout plumose 
bristles. 

Sixth limb (fig. 3/): 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. Sg, h): 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. 3^, I) : Long, slender, divided into 3 lobes, 
each with 2 annulated bristles; 1 lobe with large curved sclerotized tip. 

Furca (fig. 3j): 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.) 



RED SEA OSTRACODS — KORNICKER 




Figure 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; b, medial view of anterior end of left valve, sclerotized areas 
stippled; c, media] 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; /, 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; 
i, medial view of posterior and ventral marginal area of right valve showing distribution 
of hairs on inner lamella; /, lateral view of left 1st antenna; k, 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, I.) 



221-535—66- 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. i21 

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. 3i)- 

Frontal organ (fig. 3i): Elongate, 2-jointed with short spines on 
distal half of 2nd joint and subterminally on basal joint. 

Description of female. — Shell (figs. 4a-i): 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; UTegular polygons on surface of shell subdivided by 
minute polygons larger than on male shell and visible under low 
magnification (fig. 4/). Inner lamella with parallel striations on 
antero ventral part; 9 long hairs with secondary spines forming row 
on inner lamella behind rostrum (figs. 46, c) ; inner lamella below rostral 
incisure with small hair followed by wide space and then 5 hau's (fig. 
4e); posteroventral part of inner lamella with at least 21 hairs (fig. 4i); 
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. 4j) : 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. Si.\th 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. 4k, I) : Protopodite subtriangular without 
transparent rim. Exopodite (fig. 40 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 margmal denticulations; 9th joint with 2 short bare bristles and 
2 subequal long bristles with natatory hairs and marginal denticula- 
tions. Endopodite (figs. 4k,l) 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. 



RED SEA OSTRACODS — KORNICKER 



11 




Figure x~Euphilomedes polae, female, specim.en 1: a, medial view right mandible (surface 
hairs not shown); b, lateral view left maxilla, all bristles not shown; c, posterior view left 
5th limb; d, medial view right 6th limb; e, 7th limb;/, distal end of 7th limb; g, distal end 
frontal organ; h, frontal organ and medial eye; i, medial view right lamella of furca; 
;, 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 

Jvlandible (fig. 5a): Coxale endite large, bifurcate, with rows of 
spines. Basale^ ventral margin with 4 short bristles foUowed 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 hau-s near middle. Endopodite: distal end of 1st joint 
with 1 short and 3 long bristles ventraUy; anterior margin of 2nd joint 
with proximal group of 4 and distal group of 5 bristles; posterior margm 
with 2 subterminal bristles and termmally 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 hak along 
dorsal margin; coxa with short bare bristle on dorsal margm; 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. 5c): Epipodial appendage with about 49 bristles. 
Triangular tooth anterior to mam tooth of 1st exopodite joint about 
same size as distal tip of 2nd exopodite joint. Distal anterior margin 
of 1st 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 hu-sute. 

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. 

Furca (figs. 5i,j): 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 



^°- '"' 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 f ureal claws on 2 specimens is as follows: 

















claw number 
















/ 


t 


5 


4 


B 


6 


7 


8 


9 


10 


// 


Ig 


13 


u 


15 le 


specimen 1 
































right 


P 


P 


S 


s 


s 


s 


P 


S 


S 


s 


s 


p 


s 


s 


s s 


left 


P 


P 


s 


s 


s 


s 


P 


S 


S 


s 


p 


s 


s 


s 




specimen 2 
































right 


P 


P 


s 


s 


s 


s 


P 


s 


s 


s 


s 


p 


s 


s 


s 


left 


P 


P 


s 


s 


s 


s 


P 


S 


s 


s 


s 


p 


s 


s 





Eyes: Large medial eye (fig. 5A); lateral eyes not observed. 

Frontal organ (figs. 5g,h): 2-jointed with distal joint 3 times length 
of proximal joint; tip rounded; sm-face of 2nd joint and distal end of 
1st joint with fine spines. 

Remarks.— Extreme sexual dimorphism among some species of 
ostracods creates inherent difliculties 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 E. 
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 g 9-11* n 

*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 ll! 

CoM-pKmso^s. —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 E. 
arostratus. 



\ 



]^4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Euphilomedes arostratus Kornicker, 1967 

Figure 6 
Euphilomedes arostrata Kornicker, 1967, p. 2. 

HoLOTYPE. — USNM 112658. Sex: female. Male Atoll, Maldive 
Islands, Indian Ocean. 

Locality. — Gulf of Suez, Ked 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 
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-hke marginal denticu- 
lations. Surface of valves with distinct irregular polygons (fig. 6A); 
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. 6A). 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. 66); 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 hau-s and, distally, a dorsal, ventral, and lateral bristle, all 
bristles bare or with short hau's distally. Thu-d joint with few medial 
hairs and 1 ventral and 2 dorsal bristles, all with short hairs, the 



RED SEA OSTRACODS— KORNICKER 




Figure 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;/, endopodite of 
left 2nd antenna; g, lateral view protopodite, endopodite, and 1st 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 
scars are located below anterior hinge element; i, medial view right 6th limb (surface 
hairs not shown); ;, distal part of 7th limb; k, distal tip of 7th limb; /, detail of distal end 
of bristle of 7th hmb; m, lateral view left caudal lamella. (Same scale, in microns: 
o,e,},h-j; c,d,g,m; k,l.) 



\ 



VOL. 121 



16 PROCEEDINGS OF THE NATIONAL MUSEUM 

longer dorsal bristle also provided with wreaths of long l^airs ^^^J 
middle Fovu-th joint with 2 dorsal bristles, each with wreaths of 
Lg hah-s. and 4 long ventral bristles subdistaUy bristles bare or wi h 
hort hai;s distaUy. Fifth joint, inferred to be inserted ventraUy 
between 4th and 6th joints, bearing sensory bristle with broad base 
and numerous filaments. Sixth joint with single dorsal brise 
distally having short spines. End joint with 1 short dorsal bristle 
wHh short spfnes, 2 slender bristles with filaments, 2 bristles with 
filaments, and usual long, stout c- and f-bristles. .,, , , 

Second antenna (figs. 6/,^): Protopodite triangular without trans- 
narent rim (fig. 6^). Exopodite: first joint elongate; 2nd jomt 
about K length of 1st; 3rd joint about % length of 1st; distal margins 
of joints 2-8 with comb of short spines; basal spmes not observed^ 
First joint without bristle; bristle of 2nd joint about K length of 3rd 
joint (fig. 6y), bearing short marginal spines Bristles of pmts 3-8 
bearing natatory hairs, without marginal spines. End_ jomt w.th 4 
stout bristles with natatory hairs and 2 short slender bristles withou 
hairs. Endopodite (fig. 6/) 3-jointed; 1st joint with 5 bare short 
bristles and 1 long distal bristle with wreaths of long hairs near 
middle and short spines distaUy ; 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 obsciu-ed with the result that presence or absence 
of spine at tip could not be ascertained). _ 

Aiandible : No coxale endite observed. Basale : ventral margm with 
4 short slender bristles and 2 long bristles with wreaths of long hairs 
and short spines distaUy. Dorsal margin with 3 bristles. Medial 
surface with about 5 short bristles proximaUy near ventral corner and 
1 longer bristle near ventral margin at middle of joint. Exopodite 
reaching middle of 1st endopodite joint, with 2 s^bequal terminal 
bristles; tip of joint with blunt hh-sute process Endopodite: f^rst 
joint wi^th 1 short and 3 long ventral bristles distally. Second jom 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 
sm-face of basale and 1st and 2nd joints of endopodite provided with 

^""Maxilla: Ve'iy smaU. It seems to have a structm-e typical of the 
genus, but because of its smaU size I have not attempted to describe 
it fully. 



NO. 3571 RED SEA OSTRACODS — KORNICKER 17 

Fifth limb: Epipodial appendage with 45 phimose bristles. Outer 
lobe of 3rd exopodite joint with 2 stout plumose bristles. 

Sixth Umb (fig. Qi) : 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. Qj~l) : 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 spmes. Two long ciu'ved pegs opposing 
comb. 

Copulatory organ : Long slender, divided into 3 lobes with bristles. 

Fiu-ca (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 hmb as contrasted to 3 in this position 
on the male of E. 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 1 1 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 

Baird, W. 

1850. The natural history of the British Entomostraca, 364 pp., 36 pis. 
[Ostracoda: pp. 138-182, pis. 18-23.] 
Graf, 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. 

MtJLLER, G. W. 

1912. Ostracoda. Pt. 31 in Das Tierreich, 434 pp., 92 figs. 
POULSEN, 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 Roger 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 Bruun and also at 
Nosy Be, 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 Eudactylinidae, 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 

1 



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 oiAlopias milpinus at station 115; from 1 specimen of 
Carcharodon carcharias 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 Isurus oxyrinchus Rafines- 
que at station 282; from 1 specimen of Carcharinus floridanus at 
station 144. 

Habitat: Body sm'face 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 maculijnnnis (Poey) 
from Nosy Be, Madagascar; from 2 specimens of Carcharinus leucas 
(Valenciennes) from Nosy Be 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. 



^°- ^"^ INDIAN OCEAN COPEPODS — CRESSET 3 

Pseudopandarus gracilis Kiitesinghe, 1950 

Collections: From 1 specimen of Scoliodon palasormh (Bleeker) 
from Nosy Be; 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 tjutjot at the 
same location; from 1 specimen of Carcharinus obesus and Rhizopri- 
onodon acutus both from Durban. 

Habitat: Body surface. 

This copepod was origmaUy 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. 

Cangliopus pyriformis Gerstaecker, 1854 

CoUections: 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) 

CoUections: 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) 

CoUections: 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 

CoUections: From 3 specimens of Alopias mdpinus at stations 106, 
109 110, 115, 288, and 289; from 2 specimens of Isurus oxyrinchus 
Kafinesque 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 Lutken, 1861) 

Collections: From 2 specimens of Isurus oxyrinchus at stations 
123 139, 296, and 310; from 1 specimen of Isurus 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 Be 
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 
Be and Majunga, Madagascar, and station 119. 
Habitat: Body sm^face (usually ventral). 

Nesippus orientalis Heller, 1868 

Collections: From 6 specimens of Carcharinus maculipinnis, 4 
specimens of Sphyrna lewini (Griffith), 3 specimens of Carcharinus 
leucas, 1 specimen of Galeocerdo cuvier (Peron and LeSueui-), 1 speci- 
men of Rhizoprionodon acutus, 1 specimen of Scoliodon palasorrah, aU 
from Nosy Be; from 1 specimen of Sphyrna mokarran (Riippell) at 
station 409; from 1 spechnen 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 specmiens of Carcharinis maculipinnis, 5 
spechnens of Sphyrna lewini, 5 specimens of Carcharinus leucas, 1 
specimen of Galeocerdo cuvier, all from Nosy Be; from 1 specunen of 
Carcharinus longimanus at station 291 ; from 1 spechnen of Carcharinus 
leucas at station 409; from 1 spechnen of Sphyrna mokarran at station 
409; from 1 spechnen of Carcharinus species { floridanus?) at station 

412. 

Habitat: Roof of mouth and gill arches. 

Nesippus species 

Collections: From 1 spechnen of Galeocerdo cuvier from Nosy Be; 
from 1 specunen of the same host caught off Comores Island (Zaudsi). 

Habitat: Nasal cavities. 

This new species is described in detail in my paper (1967) re\asing 
the Pandaridae. 



NO. 3572 INDIAN OCEAN COPEPODS — CRESSET 5 

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 Be. 
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 Vaissiere, 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 Be; 
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 Isurus species caught off East 
London, South Africa; from 1 specimen of Isurus 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 Isurus oxyrinchus at stations 296 
and 310; from 2 specimens of Isurus 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 Be. 

Habitat: GUI filaments. 

Nemesis robusta (Beneden, 1851) 

Collections: From 1 specimen of Sphyrna mokarran at station 409; 
from 2 specimens of Sphyrna leioini and 4 specimens of Carcharinus 
leucas from Nosy Be. 

Habitat: Gill filaments. 

Nemesis aggregatus, new species 

Figures 1, 2, 6-20 

Collections: From 5 specimens oi Alopias vulpinus at stations 109, 
1 10, 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. 

Female. — '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 
140/z, 103;i, and 140/i respectively. Caudal rami (fig. 7) about twice 
as long as wide (85/i by 40;u), 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 maxOla, in general, as in other members of genus. Tip armed 
as in figure 11. MaxUliped (fig. 12) in form of stout claw. 




Figures 1-2. — Nemesis aggregatus, new species, dorsal view: 1, female; 2, male. 




Figures 3-5. — Eudadylina pusilla, new species: 3, dorsal view, female. Eudadylina pollex, 
new species: 4, dorsal view, female. Kroyeria ge?nursa, new species: 5, dorsal view, 
female. 



INDIAN OCEAN COPEPODS — CRESSET 



Legs 1-4 (figs. 13-16) biramose, all rami 2-segmented. Spiiie and 
setal formula as follows: 



legl 
exo. end. 


legi 
exo. end. 


legs 
exo. end. 


leg i 
exo. end 


1:0 0:1 


1:1 0:1 


1:1 0:1 


1:1 0:1 


III 11 


VII V 


VII IV 


VIII IV 



seg. 1 
seg. 2 

Leg 1 with endopod turned so that inner edge faces exopod. First 
segment of exopod curved toward endopod forming chela-like struc- 




FiGURES 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 PROCEEDESTGS 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 imiseriate, long. 

Male. — Body form as in figure 2. Total length 4.0 mm. 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: 



exo. 


leg i 
end. 


Ugg 
exo. end. 


leg S 
exo. end. 


legi 
exo. end. 


1:0 


0:1 


1:1 0:1 


1:1 0:1 


1:1 0:1 


III 


II 


IV:3 6 


111:4 5 


111:5 5 



seg. 1 
seg. 2 

Legs 2-4 (figs 17, 18, and 19) armed as in figures. Terminal endopod 
segment of leg 3 bearing a modified seta as sho\vn in figure 18. Ter- 
minal endopod segment of leg 4 also wdth 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 A^". pallida Wilson, A^. vermi 
Scott, N. robusta (van Beneden), A^. atlantica Wilson, A^^. pilosa Pearse, 
and N. macrocephalus Shiino by the nature of the spinules on segment 
2 of the second antenna. In A^". 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 spinules. Nemesis 
tiburo and A^. versicolor differ from A^. 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 foimd 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 m 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 
lewini; aU from Nosy Be. 

Habitat: Gill filaments. 



INDIAN OCEAN COPEPODS — CRESSEY 



9 



Eudactylina pusilla, new species 

Figures 3, 21-30 

Collection: 3 females from 1 specimen of Galeocerdo cuvier from 
Nosy Be. Holotype USNM 113299. Paratype USNM 113300. 
One specimen dissected for study. 

Female.— Body form as in figure 3. Dorsal surface of body 
covered with broad spinules. Total length 4.6 mm. Greatest mdth 
1.3 mm. Cephalon mder than long (1.3 by 0.8 mm), widest postero- 
laterally. First pedigerous thoracic segment fused with head. 




Figures 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 



Thoracic segments 3-5 free. Thoracic segment bearmg leg 5 free 
not combined to form genital segment. Genital segment (fig. 21) 
somewhat mder than long (0.8 by 0.7 mm). Abdomen 2-segmented. 
Caudal ramus (fig. 22) terminating in blunt process with accessory 
spme and 2 subterminal setae. 




Figures 23-30.--Eudactylina pusilla, new species, female: 23 first -"t.^'^-: f' ^^r'^. 
antenna; 25, mandible and first maxilla; 26, second maxilla; 27, max.lhped; 28, first leg, 
29, second leg; 30, fourth leg. 

First antenna (fig. 23) 5-segmented, second segment mth stout 
clawHke spine on outer distal corner. Other segments armed as 
in figure. Second antenna (fig. 24) 3-segmented, second segment 



NO. 3572 INDIAN OCEAN COPEPODS — CRESSET 11 

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) mth terminal claw, segment bearing claw \\dth patch of 
prominent spinules. Maxilliped (fig. 27) chelate, tip of movable 
process bearing broad scalelike spine opposed by process on preceding 
segment. 

Legs 1^ biramose, covered with stout spinules. Spine and setal 
formula as follows: 





exo. 


leg i 


end. 


exo. 


legi 


end. 


exo. 


legs 


end. 


leg 4 
e.xo. end. 


seg. 1 


1:0 




0:0 


1:0 




0:0 


1:0 




0:0 


1:0 0:0 


seg. 2 


1:0 




0:0 


1:0 




0:0 


1:0 




0:0 


1:0 0:0 


seg. 3 


3 




2 


II 




I 


III 




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 
figiu'e. 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 4 same as leg 3. Leg 5 (see fig. 21) broad lobe with 3 distal 
setae and 1 basal seta. Lobe covered Avith spinules. Leg 6 (see 
fig. 21) reduced to single short spme on genital segment at area of egg 
string attachment. 

Egg strings uniseriate. 

Male.^ — ^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. 

Female. — Body form as in figure 4. Dorsal siu-face of body 
smooth. Total length 3.6 mm. Greatest width 0.8 mm. 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 



measuring 360/x, 600/x, and 960/x, respectively. Thoracic segment 
bearing leg 5 free, not combined to form genital segment. Genital 
segment somewhat Avider than long (300m by 340^). Abdomen 2- 
segmented, segments measuring 141^ and llSju, respectively. Caudal 
rami (fig. 31) about twice as long as wide (132^ by 75^), bearing 3 
termmal spines and 3 subterminal setae. Rami covered with spinules 
on dorsal surface. 




Figures 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 



^°- 3572 INDIAN OCEAN COPEPODS — CRESSET 13 

form of short claw bearing 3 spines. Second segment with 2 short 
setae on inner margin. Mandible (fig, 34) styhform 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 
exo. 


end. 


leg 2 
exo. 


end. 


legs 
exo. 


end. 


legi 
exo. 


end. 


seg. 


1 


1:0 


0:0 


1:0 


0:0 


1:0 


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 


III 


1 


III 





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 spinides on both rami. Leg 3 similar to leg 4 
except endopod last segment incompletely di\aded (fig. 40). Leg 
4 (fig. 41) endopod 2-segmented. Exopod 3-segmented. Exopods of 
legs 2-4 with few spinides 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 egg string attachment. 

Egg strings uniseriate. 

Male. — Unknown. 

Remarks. — This new species can be separated from all known 
species of the genus (except E. acanthii Scott, E. 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). E. 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 E. turgipes is 3-segmented. 
It is 2-segmented in E. pollex. 

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 Be; 
from 1 specimen of the same host from station 409; from 1 specimen 
of Carcharinus longimanus from station 291. 

Habitat: GiE filaments. 

This copepod has been reported from a wide range of hosts and is 
probably cosmopolitan in distribution. 



14 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Kroyeria dispar Wilson, 1935 

Collections: From 3 specimens of Galeocerdo cuvier from Nosy Be. 
Habitat: Gill filaments. 

The only previous report of this copepod is the original description 
from Puerto Rico from Squalus. 




Figures 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 Be. 

Habitat: Gill filaments. 

This copepod has been reported previously from off the south- 
east coast of the United States. The previously reported hosts 



INDIAN OCEAN COPEPODS — CRESSET 



15 



plus the new records here mdicate that this species may be restricted 
to sharks generally found only in shallow coastal waters. 




Figures 49-53. — Kroyeria gemursa, new species, female: 49, first leg; SO, second leg; 51, 
third leg; 52, fourth leg. Male: 53, genital segment, abdomen and caudal rami. 

Kroyeria gemursa^ new species 

Figures 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 

Female. — ^Body form as iii figure 5. Total length 9.7 mm. Great- 
est width 1.5 mm. Cephalon as mde 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 wdth posterior 
sclerotized ring at point of attachment of caudal rami (see fig. 43). 
Caudal rami (fig. 43) 3 times as long as wide (264pt by 94/i). Each 
ramus fringed along inner margin, bearing 4 terminal and 2 sub- 
terminal setae. Outer 2 terminal setae stout, plumose on mner 
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 mner 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, bhmt claw at tip. Maxilliped (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: 





exo. 


leg 1 
end. 


exo. 


leg 


2 
end. 


exo. 


eg 3 
end. 


legi 
exo. end. 


seg. 1 


1:1 


0:1 


1:1 




0:1 


1:1 


0:1 


1:1 0:1 


seg. 2 


0:1 


0:0 


1:1 




0:0 


1:1 


0:0 


1:1 0:1 


seg. 3 


11:4 


6 


11:5 




6 


11:4 


4 


11:4 3 



Leg 1 (fig. 49) exopod with fringe on outer margm of all 3 segments. 
Outer margin of last 2 endopod segments with row of many short 
spmes. 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. 

Male. — 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 3-segmented, end of abdomen not sclerotized as m female. 
Caudal rami nearly 6 times as long as wide (352/i by 6lAt). 



INDIAN OCEAN COPEPODS — CRESSEY 



17 




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 



Spermatopliores 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 Be 
and station 119. 

Habitat: Gill filaments. 

Copepods Parasitic on Sharks Examined 



Lamnidae 
Isurus oxyrinchus 

Pandarus smithii 

Echthrogaleus denticulatus 

Dinemoura latifolia 

Nemesis lamna 

Anthosoma crassum 
Isurus glaucus 

Dinemoura latifolia 
Isurus species 

Anthosoma crassum 

Nemesis lamna 
Lamna species 

Echthrogaleus colcoptratus 
Carcharodon carcharias 

Pandarus cranchii 

Nesippus orientalis 

Ne77iesis 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 



INDIAN OCEAN COPEPODS — CRESSET 



19 



Carcharinus obesus? 

Pseudopandarus longus 

Carcharinus leucas 

Pandarus carcharini 
Perissopus dentatus 
Nesippus orientalis 
Nesippus crypturus 
Alebion gracilis 
Paralebion elongatus 
Nemesis robusta 
Kroyeria gracilis 

Carcharinus m aculipinnis 
Pandarus carcharini 
Nesippus orientalis 
Nesippus crypturus 
Nemesis versicolor 
Eudactylina aspera 
Kroyeria spatulata 

Carcharinus limbatus 
Alebion alatus 

Carcharinus albirnarginatus 
Alebion gracilis 

Carcharinus spallanzani 
Pandarus niger 

Carcharinus tjutjot 

Pseudopandarus longus 
Perissopus dentatus 

Carcharinus sorrah 

Kroyeria spatulata 



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 



Literature Cited 

Abildgaard, p. C. 

1794. Beskrivelse over tvende nye Monoculi Linn., Caligi Mull. {Caligus 
crassus et oblongus). Skr. Nat. Hist. Selskabet-Kjobenhavn, vol. 
3., no. 2, pp. 46-54. 

BeNEDEN, p. J., VAN 

1851. Recherches sur quelques Crustacfes inf^rieurs. Ann. Sci. Nat., vol. 14, 
no. 3, pp. 71-131. 
Bere, R, 
1930. 



1936. 

Capart, a. 
1953. 



The parasitic copepods of the fish of the Passamaquoddy Region. 

Contr. Canadian Biol. Fish., vol. 5, no. 13, pp. 1-10. 
Parasitic copepods from the Gulf of Mexico fish. American Midi. 

Nat., vol. 17, no. 3, pp. 577-625. 

Quelques cop6podes parasites de poissons marins de la region de 
Dakar. Bull. Inst. Francaise Afrique Noire, vol. 15, no. 2, pp. 
647-671. 



20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Cressey, R. F. 

1963. A new genus of copepods (Caligoida, Pandaridae) from a thresher 
shark m Madagascar. Cah. O.R.S.T.O.M. Oceanogr., ser. Nosy 
Be, vol. 2, no. 6, pp. 285-297. 

1966. Bariaka alopiae n. gen. n. p. (Copepoda, Caligoida), a parasite on the 

Gills of the Thresher Shark in Madagascar. Bull. Mar. Sci., vol. 
16, no. 2, pp. 324-329. 

1967. A revision of the family Pandaridae. Proc. U.S. Nat. Mus., vol. 121, 

no. 3570, pp. 1-133. 
Dana, J. D. 

1852. Crustacea, pt. 2. Vol. 13 in United States exploring expedition . . . 
1838-1842 . . ., pp. 691-1618. 
Gerstaecker, a. 

1854. Beschreibungzweier neuer Siphonostomen-Gattungen. Arch. Naturg., 
vol. 20, no. 1, pp. 185-195. 
Gnanamuthu, C. P. 

1951. New copepod parasites of sharks. Ann. Mag. Nat. Hist., vol. 4, 
ser, 12, pp. 1236-1256. 
Gu:6rin-Meneville, F. E. 

1837. Crustaces. Vol. 26 in Iconographie du regne animal. 
Heller, C. 

1868. Crustaceen. No. 8 in vol. 2 of Zoologischer Teil in Reise der Oester- 
reichischen Fregatte Novara . . ., 280 pp., 25 pis. 
Ho, J. S. 

1963. On five species of Formosan parasitic copepods belonging to the sub- 
order Caligoida. Crustaceana, vol. 5, pt. 2, pp. 81-98. 

KiRTESINGHE, P. 

1950. Parasitic copepods of fish from Ceylon, 3. Parasitology, vol. 40, 

nos. 1 and 2, pp. 77-86. 
Leach, W. E. 

1819. Entomostraca. In vol. 14 of Dictlonaire des sciences naturelles, 
pp. 524-545. 
Milne-Edwards, H. 

1840. Histoire naturelle des crustaces comprenant I'anatomie, la physiologic 
et la classification de ces animaux, 638 pp. 
Pearse, a. S. 

1948. A second report on parasitic copepods collected at Beaufort, N.C. 
Journ. Elisha Mitchell Sci. Soc, vol. 64, no. 1, pp. 127-131. 

1951. Parasitic Crustacea from Bimini, Bahamas. Proc. U.S. Nat. Mus., 

vol. 101, no. 3280, pp. 341-372. 
Rathbun, R. 

1886. Descriptions of parasitic copepoda belonging to the genera Pandarus 
and Chondracanthus. Proc. U.S. Nat. Mus., vol. 9, pp. 310-324. 
Risso, A. 

1826. Histoire naturelle des principales de I'Europe Meridionale et parti- 
culiferement ce celles des environs de Nice et des Alpes Maratimes, 
vol. 5, 135 pp. 
Scott, A. 

1929. The copepod parasites of Irish Sea fishes. Rep. Lancashire Sea Fish 
Lab. 1929, pp. 81-118. 



NO. 3572 INDIAN OCEAN COPEPODS — CRESSEY 21 

Scott, T. 

1901. Notes on some parasites of fishes. 19th Ann. Rep. Fish. Bd. Scotland, 
pt. 3, pp. 120-153. 
Shiino, S. M. 

1957. Copepods parasitic on Japanese fishes, 15: Eudactylinidae and 
Dichelesthiidae. Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 3, 
pp. 392-410. 
Smith, S. 

1874. Invertebrate animals of Vineyard Sound and adjacent waters. 
Rep. Comm. Fish Fisheries 1871-1872, 478 pp., 38 pis. [Reference 
not seen.] 
Steenstrup, J. J. S., and LtJTKEN, 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. 341-432. 
Vaissiere, R. 

1959. Parasites de poissons de mer ouest-africains recoltes par J. Cadenat, 
2: Copepodes. Bull. Inst. Francai§ Afrique Noire, vol. 21, ser. 
A, no. 2, pp. 534-553. 
Wilson, C. B. 

1905. New species of parasitic copepods from the Massachusetts Coast. 

Proc. Biol. Soc. Washington, vol. 18, pp. 127-132. 
1911. North American parasitic copepods: Descriptions of new genera 

and species. Proc. U.S. Nat. Mus., vol. 39, pp. 625-634. 
1913. Crustacean parasites of West Indian fishes and land crabs, with 
descriptions of the new genera and species. Proc. U.S. Nat. 
Mus., vol. 44, pp. 189-277. 
1922. North American parasitic copepods belonging to the family Dichele- 
sthiidae. Proc. U.S. Nat. Mus., vol. 60, pp. 1-100. 
1932. The copepods of the Woods Hole Region, Massachusetts. U.S. 

Nat. Mus. Bull. 158, 635 pp. 
1935. New parasitic copepods. Smithsonian Misc. Coll., vol. 91, no. 19, 
pp. 1-9. 
Yamaguti, S. 

1963. Parasitic Copepoda and Branchiura of fishes, 1104 pp. 



b 




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 Arthur G. Humes and Ju-Shey Ho ^ 



At Nosy Be, in the northwestern part of Madagascar, several species 
of copepods (Humes and Frost, 1964) are known to be associated 
with members of the alcyonarian 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 Be 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'Oceanographie et des Peches at Nosy Be. 



1 Both authors: Department of Biology, Boston University, Boston, Mass. 

1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Family Lichomolgidae Kossmann, 1877 

Genus Lichomolgiis Thorell, 1860 

Lichomolgus organicus, ueyv species 

Figures 1-30 

Type material.- — ^207 99, 98 cf'cf, and 15 copepodids from a 
colony of Tubipora musica (Linnaeus), in 1 m, Pte. Ambarionaomby, 
Nosy Komba, near Nosy Be, Madagascar, collected June 8, 1964: 
holotype 9 (USNM 113387), allotype (USNM 113388), and 144 
paratypes (USNM 113389) (100 99 and 44 c^ cf ) deposited in the 
United States National Museum, 80 paratypes (50 99 and 30 d^cf) 
in the Museum of Comparative Zoology, Harvard University, and 
the remaining paratypes in the collection of A. G. Humes. 

Other specimens.— All from Tuhi/poi^a musica collected in 1964: 
8 99 and 8 cf cf , in 1 m, Pte. a la Fievre, Nosy Be, Jan. 5; 1 9, in 
15 cm, Tany Kely, a small island south of Nosy Be, Mar. 29; 4 99 
and 8 cfd^, in 1 m, Pte. Ambarionaomby, Mar. 31; 15 99 and 1 
cf, in 1 m, Pte. a la Fievre, Apr. 15; and 14 99, in 2 m, Pte. 
Ambarionaomby, Sept. 27. 

Female.- — 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. 

Segment of leg 5 (fig. 2) Avider than long, 61^ x 133)u, ■\\dth fifth 
legs borne laterally. Ventrally between this segment and the genital 
segment no apparent intersegmental sclerite. Genital segment (fig. 2) 
about as long as mde, 140^ x 142/i, 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 80/i. Areas of attachment of 
egg sacs located dorsolaterall}'" on posterior part of expansions. Each 
area (fig. 3) with 2 naked setae, 12 and 13^ in length; posterior to the 
setae a small rounded sclerotized protuberance. Three postgenital 
segments, each without ornamentation except for a few surficial 
haks (sensilla), 37/i x 67^t, 26/x x 55/x, and 28/i x 51^, respectively, from 
anterior to posterior. 

Caudal ramus (fig. 4) only a little elongated, 36/^ x 24/i 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 23/i, along outer edge to base of lateral seta 
23/i, and to base of outermost distal seta 33/i. Of the usual 6 setae, 
outer lateral seta naked and 66/x long; remaining 5 feathered: pedi- 
cellate dorsal seta 47/x, innermost distal seta 115ju, outermost distal 
seta 91 11, and the 2 long median terminal setae, both "pegged," 
169;u (outer) and 200m (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 egg 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: 28^1 (56ai along anterior margin), 95^, 25/x, 50m, 
50/i, 37m, and 22m, 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, 65m along its outer edge, 39m along its inner edge, and 27m 
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 recm-ved claws 45m and 34m long (measured along 
greatest 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 spinides 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-segmented, 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 



end 


0:1 


0:1 


1,5 


exp 


1:0 


1:1 


III,T,5 


end 


0:1 


0:2 


1,11,3 


exp 


1:0 


1:1 


111,1,5 


end 


0:1 


0:2 


1,11,2 


exp 


1:0 


1:1 


11,1,5 



4 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

setides along one side. A minute setiile 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 
22jLi 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 nimierals= setae): 

P 1 protopod 0:1 1:0 exp 1:0 1:1 111,1,4 

P 2 protopod 0:1 1:0 

P 3 protopod 0:1 1:0 

P 4 protopod 0:1 1:0 

end 0:1 II 

Inner seta on coxa of legs 1-3 long and plumose, but in leg 4 short 
{17 fi) 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 
24ju in length. Endopod of leg 4 shorter than exopod. First segment 
33m X 25m with inner distal plumose seta 41^ long. Second segment 
Qlfx long (including 2 terminal spinous processes) and 23^ wide in 
basal half and 19^ 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 21 fj., inner 39^ long, ratio about 1:1.86. 

Leg 5 (fig. 21) with free segment elongated and slightly arcuate in 
outline, 66m x 23m, with short stout spines along outer edge and extend- 
ing around to distal ventral siu"face; bearing terminally 2 naked setae 
35m and 37m 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 egg sac (see fig. 3). 

Color in life in transmitted light rather translucid, eye red. 

Male. — 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) 29/^ x 67/x. Genital segment longer than 
wide, 164/x x 122/i in greatest dimensions, and in dorsal view rather 
bottle shaped, its lateral borders gently rounded. No intersegmental 
sclerite ventrally between these 2 segments. Fom- postgenital seg- 
ments 18)u X 45)U, 19iJ, X 43ju, 15/x x 40ju, and 17/i x 43/x, respectively, 
from anterior to posterior. 

Caudal ramus (fig. 24) resembling that of female, but slightly less 
elongated, its greatest dimensions 31jLi x 21ju, 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 formida 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, 68;^ along outer 
edge, 52 IX along inner edge, and 15/i 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. Armatiu-e like that of female, but 2 claws more nearly 
equal in length (34/i and S5fi), 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 178^ in length (measiu-ed along axis), 
with narrow terminal lamella, smooth lamellar fringe along its concave 
sm*face, and 2 setae near its base, one on posterior surface 44)U long 
and finely barbed, other on anterior siu"face 17 ix 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. On last segment of this endopod 2 spines, outermost 
of setae in female being here replaced by a spine; outer spine 29)u long, 
inner spine 43/* 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)Li (outer) and 51ju (inner), in another 16/1 and 37)u, with an average 
of 17.5/1 and 44/i, ratio of 1:2.5. 



6 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Leg 5 (fig. 28) with free segment elongated and slender, SG/x x 9/i, 
with nearly parallel sides in dorsal view, and with 2 terminal setae 
unequal, outer Slju and inner 19/x 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 23^ and 25ju long. 

Spermatophore (fig. 30) inside body of male about 148iu x 68/i plus 
neck of 14ju. 

Color in life resembling that of female. 

Etymology. — The specific name organicus, from Latin, relating 
to musical instriunents, 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 L. 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 maxUla. In L. 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 111,1,5, 
and the free segment of leg 5 in the female is 2.2:1 with a basal ex- 
pansion. In L. 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 111,1,5 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 L. gigas Thompson and 
A. Scott (1903) there is one claw on the second antenna and the body 
is larger (2 mm) . In L. longipes (Sewell, 1 949) 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 L. 
rigidus Ummerkutty (1962) the formula for the last segment of the 
exopod of leg 4 is 111,1,5 and the body is larger (1.25 mm). In L. 
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 
(1.7 mm). 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 mth 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 cTcf from a colony of Tuhipora 
musica (Linnaeus), in 1 m, Pte. Ambarionaomby, Nosy Komba, near 
Nosy Be, Madagascar, collected June 8, 1964: holotype 9 (USNM 113391) , 
allotype (USNM 113392), and 89 para types (USNM 113393) (79 99 
and 10 cfcf ) deposited in the United States National Museum, 45 
paratypes (40 99 and 5 cfcf ) in the Museum of Comparative Zoology, 
Harvard University, and the remaining paratypes in the collection 
of A. G. Hiunes. 

Other specimens. — 'All from Tubipora musica collected in 1964: 
121 99 and 53 cf cT, in 1 m, Pte. a la Fievre, Nosy Be, Jan. 5; 9 99, in 
15 cm, Tany Kely, a small island south of Nosy Be, Mar. 29; 8 99, in 
1 m, Pte. Ambarionaomby, Nosy Komba, Mar. 31; 14 99 and 3 d^cf, 
in 1 m, Pte. a la Fievre, 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. 

Female. — Body (fig. 31) Avith prosome more pointed anteriorly 
than in L. 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) 46m x 107m. Genital segment slightly- 
wider than long, 110m x 115m, in dorsal view expanded laterally just 
behind its midregion, posterior to which segment is constricted, with 
width at posterior end of segment 60m. 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 25m x 53m, 21m x 48m, and 
23m X 46m, respectively, from anterior to posterior. 

Caudal ramus (fig. 33) only slightly longer than wide, 28 X 23m in 
greatest dimensions, ratio of length to width 1.22 : 1. Length along 
inner edge to innermost distal seta 19m, along outer edge to base of 
lateral seta 19m and to base of outermost distal seta 25m. 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 o\dgerous females broken. 

Rostral area (fig. 34) slightly more elongated posteriorly than in L. 
organicus. Segments of first antenna 23m (44m), 97m, 21m, 46m, 45m, 
31m, and 20m 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 
(63m along its outer edge, 39m along its inner edge, and 21m mde). 
Larger claw 37m long, and more slender claw 32m 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 27m long. 
Endopod of leg 4 (fig. 35) wuth the following dimensions: first segment 
33m X 25m with inner seta 40m long, second segment 60m x 21m (greatest 
width) and 18m (least width) with 2 terminal spines 25m (outer) 
and 54m (inner) in length, the ratio about 2.16:1. 

Leg 5 similar to that of L. organicus, dimensions of free segment 
about 63m X 24m. 

Leg 6 (represented by 2 setae near attachment of egg sacs) like 
that in previous species. 

Color in life resembling that of L. organicus. 



COPEPODS — HUMES AND HO 



Male. — 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 25^ x 66m. Genital segment 151/i x 117^. P'our 
postgenital segments 14^ x 45/i, 16/x x 42ix, 13m x 40m, and 17m x 42M,respec- 
tively, from anterior to posterior. Caudal ramus 25m x 21m, or 1.19:1. 

Ratio of length of prosome to that of urosomc 1.57:1. 

Table 1. — Comparison of certain features of Licliomolgus organicus and 

L. conjunctus 



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 



L. organicus 



L. conjunctus 



0.99 X 0.49 mm 
rounded 

2.1:1 

36m X 24m (1.5:1) 
lateral indentations near 

middle (fig. 2) 
last segment 27/x wide, 

claws 4.5m and 34m 
24m 



0.80 X 0.38 mm 
more pointed 

2.6:1 

28m X 2.3m (1.22:1) 
lateral indentations 

further back (fig. 32) 
last segment 21m wide, 

claws 37m and 32m 
27m 



21m and 39m (1:1.85) 25m and 54m (1:2.16) 



0.89 X 0..32 mm 
rounded 

31mx21m (1.48:1) 
17..5Mand 44m (1:2.5) 

36m X 9m (4:1) 



0.71x0.27 mm 
more pointed 

25mx21m (1.19:1) 
18m and 50m (1:2.8) 

28m X 8m (3.5:1) 



Rostral area like that of female. First antenna resembling that 
of L. organicus, with same formula for setae and aesthetes. Second 
antenna also resembling that species; last segment 73m along outer 
edge, 54m along inner edge, and 14m wide. Two claws (34m and 35m) 
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 

221-5:^7—67 2 



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 27/x and inner 40^ long. Two terminal spines on 
endopod of leg 4 18/i (outer) and 50m (inner), ratio of 1:2.8. 

Leg 5 resembling that of L. organicus, but free segment relatively 
slightly shorter, 28/i x 8)tx. 

Leg 6 similar to that in previous species, but 2 naked setae slightly 
longer (28m). 

Spermatophores, seen only inside body of male, resembling those of 
L. organicus. 

Color in life like that of female. 

Etymology. — 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 L. 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 L. organicus and L. conjunctus were recovered after washing 
unbroken colonies of Tubipora 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 1 m, Pte. Ambarionaomby, Nosy Komba, near Nosy Be, Madagas- 
car, collected Mar. 31, 1964: holotypic 9 (USNM 113395) and 4 
paratypic 99, (USNM 113396) deposited in the United States National 
Museum; remaining 3 99 (dissected) in the collection of A. G. Humes. 

Other material.' — Also from Tubipora musica: 1 9, in 1 m, Pte. 
Ambarionaomby, Nosy Komba, June 8, 1964. 

Female.' — 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 H 

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, 94m x 242/i, 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, 130m x 198m. 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 11m long, a spiniform seta 8m long, and 2 small spinous 

processes. On ventral surface of posterior half of genital segment a 

transverse row of prominent spinules. Three postgenital segments 

78m X 135m, 60m x 118m, and 91m x 91m, 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) 104m x 34m, or 3 tunes longer than wide. 
Minute basal outer spinule 3m long. Outer lateral seta 22m long and 
naked. Outermost terminal seta 41m long, naked, and composed of 
2 parts: a sclerotized proximal half and a more slender distal portion. 
Innermost terminal seta 57m long and haired along inner side. Two 
long terminal setae 209m (outer) and 462m (inner), both basally 
"pegged" and bearing lateral spinules. Pedicellate dorsal seta 
39m 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 sm-faces 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) 319m x 107m, 
extending a little beyond ends of caudal rami, Avith each egg about 40m 
in diameter. 

Rostral area (fig. 44) small and well sclerotized. Posterior to it, 
between bases of second antennae, a small circidar sclerotization! 
First antenna (fig. 45) 6-segmented and relatively short and robust, 
Its length about 143m. Lengths of segments measured along their 
posterior nonsetiferous margins: 15m (39m along anterior margin), 
19m, 28m, 18m, 13m, and 24m, respectively. First segment with 5 setae, 
proximal one lightly haired, distal one spiniform, with a row of 
scaldike 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 ju x 19ju in 
greatest 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 spinides 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 spinidose 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 maxUlipeds 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): 

P 1 protopod 0:1 1:1 

P2 protopod 0:1 1:0 

P3 protopod 0:1 1:0 

P4 protopod 0:1 1:0 

Free edges of all 4 intercoxal plates ^\ith rows of long slender 
setules. Inner margin of basis of leg 1 (fig. 54) bearing spinulose 



exp 


1:0 


1:1 


111,1,4 


end 


0:1 


0:1 


1,3 


exp 


1:0 


1:1 


111,1,5 


end 


0:1 


0:2 


11,1,3 


exp 


1:0 


1:1 


111,1,5 


end 


0:1 


0:2 


11,11,2 


exp 


1:0 


1:1 


11,1,5 


end 


0:1 


0:2 


11,11,1 



NO. 3573 COPEPODS — HUMES AND HO 13 

spine 25^1 long, with near its insertion an outer row of spinules and an 
inner spiniform process with slightly bifvu-cated tip. In legs 2-4 this 
margin more or less rounded and carrying only a row of haii'iike 
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 83/x x 53ju in greatest 
dimensions. Four elements borne on this segment consisting (from 
dorsal to ventral) of a finely barbed spine 39/x long, a naked seta 66^ 
long, a strong spinulose spine 44m long, and a short weakly barbed 
spine 23iJL in length. Along dorsal edge of segment a row of spinules. 
Seta borne on body dorsal to insertion of segment 40ju 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. 

Male.- — -Unknown. 

Etymology. — The specific name latipes, from Latin, meaning 
"ha\'ing 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.Jurcvfer 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 shoidd 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. 
furcifer. 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 (pi. 82, m). The maxUliped 
is 4-segmented, the last segment ornamented with a small fanlike 
fringe, instead of being 3-segmented as in Sars' figure (pi. 82, mp^). 
The armature of the last segment of the endopod of leg 1 is 1,3 
instead of 1,11,3 as in Sars' species. The free segment of leg 5 is 
expanded instead of being narrow as in H. jurcijer. 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. jurcijer. 

Hippomolgus cognatuSf new species 

Figures 59-69 

Type MATERiAL.^Holotype 9 (USNM 113397) from a colony of 
Tubipora musica (Linnaeus), in 1 m, Pte. Ambarionaomby, Nosy 
Komba, near Nosy Be, Madagascar, collected Mar. 31, 1964, 
deposited in the United States National Musemn. 

Other material. — 2 99 from Tubipora musica, in 1 m, Pte. a la 
Fievre, Nosy Be, Jan. 5, 1964, 1 9 (USNM 113398) in the United 
States National Museum, the other (dissected) in the private collection 
of A. G. Hmnes. 

Female.^ — Body (fig. 59) resembling that of H. latipes. Length 
(not including setae on caudal rami) 0.90 mm (0.82-0.95 imn) 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, 78m x 188)u. Genital segment (fig. 60) 
only slightly wider than long, 120^ in length, 132ju wide in its an- 
terior half, and 110m ^vide 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 22/x long, a spine 8jLt long, and 2 spinous proc- 
esses. Three postgenital segments 66/x x 101m, 60m x 91m, and 80m x 82m, 
respectively, from anterior to posterior. 

Caudal ramus (fig. 62) 64m x 31m in greatest dimensions, or 3 times 
longer than wide. Outer lateral seta 25m, outermost terminal seta 
39m, innermost terminal seta 88m, 2 long terminal setae 220m (outer) 
and 450m (inner), dorsal seta 33m long. 

Ratio of prosome to m"osome 1.17:1. 

In single ovigerous female collected egg sac (fig. 63) 352m x 138m, 
reaching nearly to ends of caudal rami, with each egg about 47m 
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 
bifm-cated; 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, 61m along inner 
and ventral margin, 51 /x along outer and dorsal margin, and 22ai 
wide. Armature from dorsal to ventral consisting of spine 21/i, 
seta 50m, spine 25/x, and spine 40m 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. 
Male.' — Unknown. 

Etymology. — 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.—^, cognatus differs from 
H. 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 1,4 instead of 1,11,3 as in 
Sars' species, and in having a relatively much shorter caudal ramus, 
3:1 instead of 13:1 as in R 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 
armatm-e 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 Tuhipora 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 

BocQUET, C, and Stock, J. H. 

1957. Copepodes parasites d'invertebres des cotes de France, 1: Sur deux 

genres de la famille des Clausidildae, commensaux de mollusques: 

Hersiliodes Canu et Conchyliurus nov. gen. Proc. Koninkl. 

Nederlandsche Akad. Wetensch. Amsterdam, ser. C, vol. 60, 

no. 2, pp. 212-222. 
Brady, G. S. 

1910. Die marinen Copepoden der Deutschen Siidpolar-Expedition 1901- 

1903, 1: liber die Copepoden der Stamme Harpacticoida, Cyclo- 

poida, Notodelphyoida und Caligoida. No. 5 of vol. 11 (Zool. 

vol. 3) in Deutsche SiJdpolar-Expedition 1901-1903, pp. 497-593. 

GURNEY, R. 

1927. Zoological results of the Cambridge expedition to the Suez Canal, 
1924, XXXIII: 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 alcyonarians 
and madreporarians in Madagascar. Cah. O.R.S.T.O.M. Oceanogr., 
S6r. Nosy-Be, 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. 
NiCHOLLS, 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. 0. 

1917. Clausidiidae, Lichomolgidae (part). Pts. 11 and 12 of vol. 6 (Co- 
pepoda, Cyclopoida) in An account of the Crustacea of Norway 
with short descriptions and figures of all the species, pp. 141-172. 
Sewell, R. B. S. 

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 invei'tebrates. Pubbl. Staz. 
Zool. Napoli, vol. 31, no. 1, pp. 59-75. 
Thompson, I. C, and Scott, A. 

1903. Report on the Copepoda collected by Professor Hcrdman, at Ceylon, 
in 1902. Rep. Gov. Ceylon Pearl Oyster Fish. Gulf of Manaar, 
suppl. rep., no. 7, pp. 227-307. 
Ummerkutty, 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. 




Figures l-%.-Luhomolgus organicus, new species, female: 1, dorsal (A); 2, urosome, dorsal 
(i5); 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, with two arrows indicating 
positions of aesthetes added in male, dorsal (F); 8, second antenna, posterior (F). 
(Capital letters=scale at which drawn; a '=first_ antenna, a"=second antenna, r=ros- 
tral area.) ' 

17 




Figures 9-\S.—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= maxilliped, Pi==leg 1.) 
18 




I 



Figures 19-27. —Lie homolgus 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 




29 








Figures 28-36. — Lichomolgus organkus, new species, male: 28, leg 5, dorsal (D); 29, leg 6, 
ventral (G); 30, spermatophore, inside male, ventral (F). Lichomolgus conjundus, new 
species, female: 31, dorsal (A); 32, urosome, dorsal (B); 33, caudal ramus, dorsal (C); 34, 
rostralarea,ventral(B);35,endopodof leg 4, anterior (F). Male: 36, dorsal (I). (Capital 
letters=scale at which drawn; a'=first antenna, a"== second antenna, r= rostral area.) 
20 




42 



41 



39 






Figures 37-Ai.—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 (I); 44, rostral area, ventral (F). (Capital letters-scale at which drawn- a '= first 
antenna, a "= second antenna, lm=labrum.) 



21 




Figures 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 




Figures S4r-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 




Figures 6\-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=maxilliped; Pi = leg 1, a' = first antenna, a" = second antenna, Im- 
labrum.) 



24 



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the United States 
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SMITHSONIAN INSTITUTION . WASHINGTON, D.C. 




Volume 121 19(37 



Number 3574 



COPEPOD CRUSTACEANS 

PARASITIC ON TELEOST FISHES 

OF THE HAWAIIAN ISLANDS ' 



By Alan 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 vnth 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 vnth 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- 
scribmg 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 Shimo (1963) describes Midias 
lobodes, Caligus coryphaenae, and BracUella thynni from fishes ex- 
ammed in t he 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 Oceanogi-aphy, University 
of British Columbia, Vancouver, Canada. 



2 PROCEEDINGS OF THE NATIONAL MUSEUM "^o^- ^^i 

(1958) lists, by family, the copepods taken from the stomachs of 
some parasite-pickmg fishes of the genus Labroides and (1961) the 
parasitic copepods taken from the Manini (Acanthurm tnostegus 

The study here reported has benefited from coUections 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, especiaUy by Drs. 
Bo^^^an, Mannmg, 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. 

Methods.— The external surface, gill cavities, buccal cavity, and 
nasal cavities of the teleost hosts were examined for parasitic copepods. 
Copepods coUected were kiUed in either 95 percent ethyl alcohol or 
10 percent formalin and later transferred to 95 percent ethyl alcohol. 
Specunens to be dra^^^l or dissected were placed m 85 percent lactic 
acid to clear and soften them, stamed with Chlorazol Black E dis- 
solved m 85 percent lactic acid, and then placed in benzyl alcohol tor 
final clearing and for dissection and drawing. 

Drawmo-s of the enthe anhnal were made from specnnens placed 
in benzyl ''alcohol and covered with a cover shp, supported so that 
the shape of the organism was not distorted. Both a camera lucida 
and a Bausch and Lomb Tri-Smiplex Micro-Projector were used m 
makmg the drawings. The appendages and processes were drawn_ m 
situ or were removed and mounted m either Hoyer's mountmg 
medium or a 1:1 mixture of Turtox's CMC-10 and CMC-S. Meas- 
urements were made with an ocular micrometer on specunens softened 
m lactic acid and held loosely in place by a cover shp supported 

by spacers. , i + i„ 

In the following figures the 9 and c^ signs are used separately 
under each drawing to mdicate a difference between the appendage 
or body part of the female and that of the male. The symbols are 
used together (9 cT) to mdicate the shnilarity 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 fine under the 
appropriate symbol. If only one sex is represented in the coUection, 
the symbols are not used. ■ j- 4. 

Terminology.— The term "cephalothorax" is used to mdicate a 
condition m which one or more of the thoracic segments are fused with 
the cephalon. The maxfiliped-bearmg 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 



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) m the Caligidea (Caligoidea of Yamaguti 1963) 
In the lernaeocerids, penneUids, 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 mimediately posterior to the cephalothorax. The term 
^ 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 contam eggs. 

With three exceptions, the terminology applied to the appendages 
and processes is the same as that in Lewis (1964a). 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 "maxiUipeds" refers 
to the modified pair of appendages on the first thoracic segment whUe 
the sternal furca" is a bifurcate, posteroventrally directed projection 
between and slightly posterior to the maxilliped bases of trebiids 
euryphorids, and caligids. The term "thoracic legs" designates the 
pair of semifohaceous or fohaceous appendages arising from some of 
the thoracic segments posterior to the first or maxiUiped-bearino- seg- 
ment, whOe the "caudal rami" are the pair of appendages at the 
posterior end of the body. 

The three exceptions to the original termmology have arisen from 
a recently completed study made by the author on the nature of the 
maxillae m the Cahgidea (ms in preparation). It appears that the 
setule- or seta-bearing node lateral to the mouth cone of this group 
of cahgoids is the maxiUule, but it seems that the spine or plate imme- 
diately behind this structure in the trebiids, eirgids, euryphorids, and 
cahgids IS an accessory formation and not part of the maxiUule. ' The 
name "maxiUule" 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 hnmediately behmd the maxiUule. The 
term "maxUlae" applies to the second pair of maxiUae with its asso- 
ciated pair of openmgs to the maxUlary glands. The term "post- 
antennal process" is used to designate the process-bearing nodules and 
the spmelike projection, if present, situated lateral and sli-htly 
posterior to the base of the antenna. 

To faciUtate the use of the thoracic leg tables, a hypothetical tho- 
racic leg IS shown in figure 1, giving the various thoracic leg armature 



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. , • ^u 

In the section entitled "Material" for each of the species the 
abbreviation USFWS means U.S. Fish and Wildhfe Service HMS 
means the USFWS vessel Hugh M. Smith, CHG the USFWS vessel 



P H dH nihrriH 



dmH 




Figure 1.— Hypothetical thoracic leg showing various armament components 



a: adhesion surface 
C: heavy fringing plumosities 
c: light fringing plumosities 
D: large denticulations 
d: small denticulations 
dH: large denticulated spine 
dm: denticulated membrane 
dmH: large spine with denticulated mem- 
brane 
fm: frilled membrane 
fmH: large spine with frilled membrane 
H: large spine 



IP: interpodal plate 

m: membrane 

P: plumose seta 

p: plumose setule 

P': naked seta 

p': naked setule 

pH: large plumose spine 

Q: seta plumose on one side and mem- 
branous on other 

qH: large spine plumose on one side and 
membranous on other 

rh: spinule 

s: solitary hairlike process 



h : small spine 

Table 1.— Armature of hypothetical thoracic leg shown in figure 1 





Surface 


Inter- 
podal 
Plate 

m 


Protopodite 


Exopodite 


Endopcdite 


Leg 


1 


2 


1 


2 


3 


1 


2 


3 




Outer 
Inner 


s,P 


fm.p 
m,a,s 


rh.dni, 
dnill 

c,P 


{mil 
c,P 


H.h.nill, 
mh.dll 

CQ.qH.pH 


d,D, 
C 

P 


c 
c,2P 


d,p,p',P', 
P 

c,p',P' 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 5 

Charles U. Gilbert, JRM the USFWS vessel J.R. Manning, and LL 
means a USFWS longiine 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 

Pseudotaeniacanthiis Yamaguti and Yamasu, 1959 

Diagnosis. — Body cyclopoid, cephalon fused with first 2 thoracic 
segments, shghtly 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 mthout large, posterior 
hooks at apex of Y. Maxillule 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. 

Pseiidotaeniacanthus puhi, new species 

Figures 2, 3 

Material. — 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 

Description. — Female cephalo thorax (fig. 2a) bell shaped, consist- 
mg of cephalon, maxilliped-bearing, and first pedigeroiis segments. 
Second pedigeroiis segment distinct from and slightly narrower than 
cephalo thorax; third pedigeroiis segment distinct from and slightly 
narrower than second; fom-th pedigerous segment distinct from and 
slightly narrower than thu'd, broadly rounded posteriorly. Fifth 
pedigerous segment approximately haK 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. 2/) subrectangular, short, with 2 long and 3 short terminal setae. 

Male prosome (fig. 26) 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. 2%) 5- or 6-segmented, proximal 
segment expanded on proximal posterior surface. Female and male 
antenna (figs. 2l, 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. Thu'd 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. 2j) rodhke, 2-parted. Fkst 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. 2i), arms of Y with 9-10 transverse rows of spinules giving 
brushlike appearance. Female and male maxillule (fig. 2j) situated 
immediately posterior to mandible base, consisting of pair of setae 
attached to platehke area of heavy sclerotization contiguous with 
posterior edge of platelike area of heavy sclerotization forming attach- 
ment surface for mandible. Maxilla (fig. 2k) represented by large. 

Figure 2. — Pseudotaeniacanthuspuhi,ne.vi%^&dQ%,doxs,a\ 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 fiist abdominal segment. Right fifth 
leg: e, posterior view. Ventral view: /, caudal ramus; g, female cephalothorax (mdbl= 
mandible, mxpd = maxilliped); h, male cephalothorax (mxpd = maxilliped); i, right antennule 
and heavily sclerotized postantennal process; ;', right mandible (mdbl), maxillule (ma-1), 
and paragnath (pg); k, maxilla; /, antenna; m, projection on proximal portion of third 
segment of antenna. 



COPEPOD CRUSTACEANS — LEWIS 







1 — > — I — I — 1 — I 

0.05 mm ' 



8 PROCEEDINGS OF THE NATIONAL MUSEUM ^o^- '-' 

heavily sclerotized, knoblike projection posterior and lateral to oral 
region. Knob with broad base, tapered to narrow distal region with 




O.OSmnn. 9^ 

e 

Yrov..3.^Pseudotaeniacanthus puhi, new species: ., male maxiUiped, ventral vJe- 
Thoracic legs: b, left first, anterior view; c, left, second, posterior view; d, right third 
posterior view; e, right fourth, posterior view. 

medially facing cuphke indentation, indentation bearing single, seta- 
like process. Female and male paragnath (fig. 2j) minute, situated 



COPEPOD CRUSTACEANS — LEWIS 



9 



immediately posterior to maxillule, consisting of single, jflabby, 
subcorneal 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 
Pseudotaeniacanthus puhi, new species 


Of 


Leg 


Surface 


Inter- 
pod al 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


1 


Outer 
Inner 


c 


P 
P 






P 


9P 
P 


c 
P 


C 
P 


C,3P 
2P' 


II 


Outer 
Inner 


D 


D 


P 
D 
P' 


D,pH* 
c 


D,pH* 
P 


D,pH*,D,2pH 
5P 


2D 
P 


2D 
2P 


2D,P,2D,2P 
2P 


III 


Outer 
Inner 


D 


D 

rh 


P 
D 


D,pH 
c 


D.pH 
P 


D,pH,D,2pH 
5P 


C,D 
P 


CD 
2P 


C, D,P,2D, 
2P 
2P 


IV 


Outer 
Inner 


D 


D 

rh 


P 
D 


D,pH 
e 


D,pH 
P 


D,pH,D,2pH 
5P 


2C 
P 


C,D 
P 


C,2D,2pH 
P 



*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 armatm^e 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 

Diagnosis. — 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 3- 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 OP 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- 
moiis, first with 2- or 3-segmented rami; second to foui'th with 3- 
segmented rami. Fifth legs uniramoiis, 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 woidd 
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 nioa, new species 

Figures 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 specmien of 0. 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 : 

Greatest length, excluding caudal setae 

Length of prosome 

Width of prosome 

Length of cephalothorax 

Length of genital segment 

Width of genital segment 

Length of egg sac (1 sac) 



mean 


range 


0. 76 


0. 70-0. 85 


0.57 


0. 52-0. 62 


0.40 


0. 34-0. 43 


0.33 


0. 30-0. 36 


0.07 


0. 06-0. 07 


0.13 


0. 12-0. 14 




0.55 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 11 

Description of female. — Body (figs. 4a, b) of general taenia- 
canthiform nature. Cephalothorax consisting of cephalon, maxilliped- 
bearing, and first pedigerous segments; second througli fourth 
pedigerous segments distinct. Frontal region (fig. 4e) narrow, un- 
armed except for small, slightly projecting rostrum. Cephalothorax 
projectmg 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 fom*th pedigerous segments decreasing in 
width, with ventraUy and posteriorly projecting extensions as on 
cephalothorax except not as well developed. Fourth pedigerous 
segment rounded posteriorly, extending over dorsal siu-face 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. 4/) 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 
pkunose 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 sm-face, 3 small, naked setules 
from posterior distal sm^face, 3 long, naked setules and one aes- 
thete (?) from anterior distal surface. Antenna (figs. 4,ry, 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 IK times the length of second, with large, elongate 
projection on inner surface, extending from distal end of segment 
proximaUy, 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. 



12 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 j 




COPEPOD CRUSTACEANS — LEWIS 



13 



Mandible (fig. 5a) 2-segmented, first segment slightly longer than sec- 
ond, second bearing 2 short, flattened, pectinate processes distally, 
ventralmost longest. 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 un- 
mediately posterior to mandible base, bearing 2 long, very lightly plumose 
setae, 1 shorter naked seta and 2 minute setules. Maxilla (figs. 5a, c) 
with one distinct segment although indication of second segment fused 
with cephalo thorax; armature of distinct segment consisting of 2 spikelike 
projections, one subterminal and very hghtly plumose, second terminal, 
finely denticulated along inner margin. Paragnath (fig. 5a) lobate, 
curved inward sharply, with slightly fuzzy distal end. 

Maxilliped (fig. 5d) 2-segmented 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 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 


C 


c,P 
2P,d 




2H 


2n,P 
6P 




P 


C 
7P 




II 


Outer 
Inner 


C 


c,P 
C 




dm,H 


dm, II 
P 


2dm,3H 
4P 


P,C 


P,C 


dm,3H 
3P 


III 


Outer 
Inner 


C 


C,P 
c,c 




dm,H 


dm,H 
P 


2dm,3n 
4P 


P,C 


P,C 


dm,3H 
2P 


IV 


Outer 
Inner 


C 


C,P 




dm,H 


dm,H 
P 


3H 
4P 


P,C 


dm 
P,C 


2dm,3H 


V 


Outer 




C,P* 




dm,2H, dm, P, dm, 
H,C* 













'Designation under protopodite and exopodite for convenience; association of segment uncertain. 

Figure 4. — Anchistrotos moa, new species, female: a, dorsal view; b, lateral view; c, genital 
segment, sixth legs, abdomen, and caudal rami, ventral view; d, fifth leg, ventral view; 
e, rostrum, frontal region and proximal segment of antennules, dorsal view;/, 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 




0.0 5 mm. 



NO. 3574 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 Diclielestliiidae 
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 
buamous, rami 1- or 2-segmented; third and fourth thoracic leg 
pau's rudimentary or absent; fifth pair absent. 

Diagnosis 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 

Material. — Thirteen adult females from the external surface of 
Mulloidichthys auriflamma Forskal collected at Kauai, Hawaii. One 

Figure 5. — Anchislrotos 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;/, left second; g, left third; h, right 
fourth. 



mean 


range 


0.67 


0. 61-0. 73 


0. 18 


0. 16-0. 20 


0.29 


0. 24-0. 33 


0.56 


0. 49-0. 62 


0.23 


0. 20-0. 25 


0. 44 


0. 36-0. 52 



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). 
Measurements.^ — (In mm) 10 females: 

Greatest length of body 

Length of cephalothorax 

Width of cephalothorax 

Length of trunk 

Width of trunk 

Length of egg strings (5 strings) 

Description of female. — Body (fig. 6a) divisible into two regions, 
Cephalothorax, including cephalon and maxilliped-bearing thoracic 
segment, forming first region, pedigeroiis 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 forifiing 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 siu-face. Caudal rami (fig. 66) small, each tipped 
by 2 small setules. 

Antennule (fig. 6c) 6-segmented, strongly compressed. First 
segment with 6 irregular naked setae from ventral margin; second 
segment with 3 setae; third w^ith 1 from ventral margin, 1 from distal, 
and third from dorsal surface; fom'th segment naked; fifth with one 
seta from dorsal surface; sixth with 7 setules distally. Antenna (fig. 
Qd) 2- or 3-segmented, basalmost portion squat and irregular, segment- 
like but questionable, with several knoblike articulation siu"faces 
received by proximal end of penultimate segment. Penultimate 

Figure 6. — Hatschekia breviramus, new species, female; a, dorsal view; ^, posterior region, 
left side showing caudal ramus (r) and anal indentation (a); c, right antennule, anterior 
view; d, left antenna, ventral view; e, membrane-like flap at base of antenna, ventral view; 
/, 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; i, distal end of maxilliped, ventral view. 



COPEPOD CRUSTACEANS — LEWIS 



17 




226-321—67 2 



18 



PROCEEDINGS OF THE NATIONAL MUSEUM 



segment elongate, broader proximally than distally, posterior and me- 
dial sm^aces rugose. Distalmost segment short, subspherical, fused 
with clawlike terminal process, with very fine, hairlike accessory 
process (not shown in figiu-e). Distinct flaplike projection present on 
heavily sclerotized cephalothoracic depression associated with an- 
tenna base (figs. Qd, e). Mandible (fig. 6/) appearing 3-parted although 
divisions indistinct; appendage rodlil^e, 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, nregularly pointed distally, with single 
setule from distal surface. 

MaxHliped (figs. 6^,^) 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 proxunally 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, mnermost 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 natm'e and 
armature of legs see figiu"es 7a, b, and table 4. 




0.03 mm 



003mm. 



Figure 7. — Hatsckekia breviramus, new species, female, thoracic legs: a, first; b, second. 



Discussion. — Hatsckekia 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, 



COPEPOD CRUSTACEANS — LEWIS 



19 



Table 4. — Armature of thoracic legs I and II of the female of Hatschekia brevira- 

mus, new species 



Leg 


Surface 


Interpodal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 2 


I 


Outer 
Inner 




P' 


P' 


P' 
P' 


P' 
P 




II 


Outer 
Inner 






P' 


P' 
P' 


p' 


P' 
2p' 



distinct principally in nature of armature of antennule and thoracic 
legs. 

2. H. ostracii Yamaguti, 1953: Similar in body shape and in 
the presence of the processes at the base of the antennae. H. 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 iJ. ostra- 
cii. The endopodite of the fu'st thoracic leg is 2-segmented in H. 
ostracii and 1-segmented in H. breviramus; H. ostracii also has a 
rudimentary thh*d 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. ostra-cii 
Yamaguti, 1953, has the projections at the base of the antennae that 
are present in H. breviramus. The maxilliped armature is also similar 
but the 2 species differ, most distinguishably in the armatm-e 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-segmented 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 Nordmanii, 1832 

Diagnosis. — 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, — 8 host records: 

location hosts reference 

Hawaiian Islands Unknown Nordmann, 1864 

Acanthurus dussumieri Lewis, 1964a 

A. mata 
A. olivaceus 

A. triostegus sandvicensis 
Ctenochaetus strigosus 
Naso hexacanihus 
N. unicornis 

Material. — 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. 

Description. — See Lewis, 1964a. 



Family Pennellidae 

Pennella Oken, 1816 

Diagnosis. — 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, smTOunded by one or more types of 
papillae. Two to thi'ee 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, cylmdrical 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 sm'face 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 length of body. Antennule 2- to 5- 
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 21 

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 

Figures 8, 9 

Pennella histiophori Thomson, 1889, p. 368, pi. 38, fig. 2. — Bassett-Smith, 1899, p. 

483.— Wilson, 1917, p. 113 (key).— Yamaguti, 1963, p. 208. 
Pennella zeylanical 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 


Histiophorns herschelli 


Thomson, 1889 


Ceylon 


H. gladius 


Kirtisinghe, 1932, 1964 



Material. — One complete, ovigerous female, 1 nonovigerous female 
without cephalothoracic papiUae, and 1 incomplete female (trunk and part 
of neck missing), in addition to part of larval exuvium, collected by D. 
W. Strasburg from external siuface of Makaira audax (PhHippi) at 
9°34'S, 136°45'W (USNM 112871). 

Remarks. — Although the host was collected far from the Hawaiian 
Islands, the wide distribution of the host and its common occm-rence 
around Hawaii suggested the inclusion of the copepod in this treatise. 

Measurements. — (In mm) 2 females: 

Total length (1 specimen) 

Length of cephalothorax, excluding papillae 

Width of cephalothorax 

Length of neck 

Width of neck 

Length of trunk 

Width of trunk 

Approximate length of abdominal plumosities 

Length of attachment horns 

Length of egg string (1 string) 178 

Description. — Female: Cephalothorax (figs. 8a, b) presumably 
consisting of cephalon and fu'st thoracic segment although maxillipeds 



172 






7,7 






8, 8 






79, 51 






3,2 






86, 89 






6, 5 






25, 28 






(17, 10), 


(12, 


12) 



22 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




O.Smm. 



Figure 8. — Pennella hisliophorP. Thomson, 1889, female: a, dorsal view; h, 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);/, cephalothorax, 
anterior view showing oral region; g, lateral view of some papillae in oral region. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 23 

not visible. Cephalotliorax with 2 large, heavily sclerotized, dac- 
tyliform attachment processes on posterior lateral sm-face, processes 
projecting laterally and slightly posteriorly. Anterior end of cepha- 
lothorax with 4 adjacent rows of soft, UTegular papillae (figs. 8/, g) 
running dorsoventraUy, 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 
thu'd 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. 8e). Caudal rami knobhke, 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. 96) 3-segmented although articulat- 
ing in doughnut-shaped structure distinct from cephalothoracic 
cuticle. Fu-st segment with 7 very lightly plumose setae on anterior 
surface, second segment more elongate than fu'st, with approximately 
1 1 very lightly plumose setae on anterior and anterior dorsal surface in 
addition to 1 elongate, aesthete-like structure on distal anterior sur- 
face. Thu'd 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 uTegulariy 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 



with large, irregular, heavily sclerotized projection of outer portion 
that forms articulation surface for third segment. Third segment 
small, proxhnal 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 thu-d; ratio of 
proximity equals 1 (first to second leg) : 5.5 (second to third) : 5.5 (thu-d 
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 appearmg 
uniramous, both in adult female (fig. 9/) and larval exuvium (fig. 9j), 
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 m alcohol-preserved specimens. For nature and armature of 
thoracic legs, see figures 9e-j and table 5. 

Tablk 5. — Armature of thoracic legs I-IV of the female of Pennella histiophori? 

Thomson, 1889 



Leg 


Surface 


Interpodal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


I 


Outer 
Inner 




P 
P' 


P 


2P 
3P 


P 


3P 
4P 


II 


Outer 
Inner 






P 


3P 
3P 






III 


Outer 
Inner 




P' 










IV 


Outer 
Inner 




P' 











Discussion. — The identification of the Hawaiian 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 mde 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;/, second (endopodite missing), anterior view; g, third (ramus 
or rami missing), posterior view; h, fourth (ramus missing), posterior view; i, distal end 
of protopodite of fourth, posterior view. Pedigerous region of larval exuvium: ;', ventral 
view (A = anterior). 



COPEPOD CRUSTACEANS — LEWIS 



25 




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 structiu-es 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 (pi. 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 

Material. — 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=l) 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=l) 21.00 

Width of trunk (N=l) 1.50 

Approximate length of abdominal pluinosities (N=l) 7.13 

Length of attachment structures (N=2) (2.25, 2.25), (3.15, 4.05) 

Description. — Cephalothorax (figs. 10a, b) presumably consisting 
of cephalon and first thoracic segment although maxOlipeds not dis- 
cernible. Pair of long, dactyliform attachment processes present 
immediately posterior to cephalothorax. Cephalothorax with 2 types 



COPEPOD CRUSTACEANS — LEWIS 



27 




I — 1—1 — I — ^- 

0.5 mm. 



Figure W.—Pennella species, female: a, doisal view (specimen 1); b, anterior trunk region 
and cephalo thorax, dorsal view (spec. 2); c, cephalothorax, ventral view showing large 
peripheral and smaller oral papillae (spec. 2) ; d, three oral papillae (spec. 2) ; e, 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. 
I0e,f), each with extension of alimentary tract running to blind termi- 
nation at distal end. Distal end of abdomen bUobed, anal opening at 
median indentation, each lobe with minute caudal ramus. Caudal 
rami (fig. 11a) 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. lid) 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 knobhke heavily sclerotized region distally. 
Second segment sHghtly 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 lle-h and table 6. Some errors are beheved 
incorporated into both the figures and the table because of the method 



COPEPOD CRUSTACEANS — LEWIS 



29 




O.lmm 



0.1 mm. 



Figure 11. — Pennella species, female: a, caudal ramus, dorsal view (a = anus) ; Z), 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;/, left second (exopodite missing); g, 
right third; h, right fourth. 



30 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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 



Leg 


Surface 


Interpodal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


I 


Outer 
Inner 




s, p' 
P' 


? 


5P? 


? 


6P? 


II 


Outer 
Inner 




s 


? 


6P? 


P 


7P 


III 


Outer 
Inner 




s 




5P, p 


P 


6P, p 


IV 


Outer 
Inner 








4P, p 







Discussion. — 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 bUobed 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 

Diagnosis. — See Lewis, 1964a. 



COPEPOD CRUSTACEANS — LEWIS 



31 



Norion expansus Nordmann 

Norton expansus Nordmann, 1864, p. 489, pi. 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 liluratus 

N. hexacanlhus Lewis, 1964a 

Description. — See Lewis, 1964a. 

Family Pandaridae 
Nesippus Heller, 1865 

Diagnosis. — See Lewis, 1964a. 

Nesippus costatus? Wilson 

Nesippus costatus Wilson, 1924, p. 213, pi. 20. 
Nesippus costatus? — Lewis, 1964a, p. 211, figs. 19-21. 

Hosts and distribution.— 7+ host records: 

location 
Galapagos Islands 
Hawaiian Islands 



reference 
Wilson, 1924 
Lewis, 1964a 



hosts 
"A nine-foot shark" 
Cysts on: 
Acanihurus triostegus sandvi- 

censis 
A. nigroris 
A. xanthopierus 
Ctenochaetus strigoszis 
Diodon holocanthus 
Scarus species 
labrids 
pomacentrids 
zanclids 

Material. — 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 (Lmnaeus), Chae- 
todon miliaris Quoy and Gaimard, and Myripristis pralinius Cuvier 
and Valenciennes. 

Measurements.— Length of body, excluding caudal setae, 1.96mm. 
Length of cephalothorax, including frontal region, 1.39mm; width 
1.55mm. Length of genital segment 0.34mm; width 0.47mm. 
Length of caudal rami 0.22mm. 

Description. — See Lewis, 1964a. 

Family Euryphoridae 
Euryphorus Milne-Edwards, 1840 

Diagnosis. — Cephalothorax suborbicular, consisting of cephalon 



32 PROCEEDINGS OF THE NATIONAL MUSEUM 



and first 4 thoracic segments, frontal region well defined. Free fourth 
pedigerous segment with alae (larger m female than in male) . Female 
genital segment large, siiborbicular, 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. Antennide 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 consistuig of 2 nodides 
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, pi. 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, pi. 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. 

Euryphorus coryphaenae Kr0yer, 1863, p. 161, pi. 10, fig. 4a-h. — Wilson, 1913, p. 
225.— Yamaguti, 1936b, p. 1, pi. 1, figs. 1-17.— Bonnet, 1948, p. 7.— Causey, 
1953a, 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 Coryphacna hippurus Yamaguti, 1936b 

Neothynnus macropterus Shiino, 1954b 

Hawaii Coryphaena hippurus Bonnet, 1948 

Indian Ocean "Seefischen" Kner, 1859 

Coryphaena hippurus 

C. h. equisetis Kirtisinghe, 1937 



"°- ^"* COPEPOD CRUSTACEANS — LEWIS 33 

"'^"'^^ f^ost reference 

Tropical, sub- 
tropical 

Atlantic Lampugus punciulatus Steenstrup and Lutken, 1861 

Coryphaena hippurus Kr0yer, 1863 

Gulf of Mexico Coryphaena hippurus Causey, 1953a 

_ Material.— 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 trollmg 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 mUes south of Niihau, Hawaii 
(USFWS). 

Measurements.— (In mm) 5 females and 5 males: 

female male 

Length of body, excluding caudal setae 

Length of cephalothorax, including frontal 
region 

Width of cephalothorax 

Length of genital segment, excluding gen- 
ital flap 

Width of genital segment 

Length of abdomen 

Length of alae 

Length of genital flap 

Length of abdominal projection 

Length of caudal rami 

Length of egg string (1 string) 

Description.— Female cephalothorax (fig. 12a) ovoid, consisting 
of cephalon, maxiUiped-bearing, and first 3 pedigerous segments. 
Frontal region broad, approxunately one-tenth the length of cepha- 
lothorax, distmctly separable from rest of cephalothorax; anterior 
edge with fine membrane. Lateral cephalo thoracic margms flatly 
convex except for sharp median mdentation, margm bearmg narrow 
membranous flange. Posterior lateral cephalo thoracic regions sharply 
rounded, extending posteriorly shghtly past median cephalothoracic 
region. Posterior sinuses distmct (fig. 12c), U-shaped, with narrow 
but distmct flange extending medially from outer margin and bear- 
ing fine membrane. Dorsal cephalothoracic grooves forming uTeg- 
ular H, posterior longitudinal grooves of H formed by junction of 
posterior lateral cephalothoracic region and median cephalothoracic 
region; crossbar uTegular, in posterior thhd of body; anterior longi- 
tudinal grooves extending anteriorly to just lateral to 3-parted ocufar 
region. Ocular region consisting of 2 ovoid contiguous pigmented 
bodies, each with lens, and smaller pigmented area between posterior 

226-321—67 3 



mean 
9. 65 


range 

7. 80-10. 50 


mean 
5.84 


range 
5. 33-6. 45 


2.77 
2.75 


2. 63- 2. 96 
2. 63- 2. 81 


2.46 
2.35 


2. 37-2. 52 
2. 26-2. 52 


L98 
2.55 
4.31 
0.78 
0.67 


1. 52- 2. 22 

1. 85- 2. 85 

2. 70- 5. 10 
0. 74- 0. 83 
0. 61- 0. 72 


L27 
L 13 
L50 
0.55 


1. 15-1. 44 
1. 04-1. 18 
1. 43-1. 58 
0. 45-0. 61 


4.85 
0.46 


2. 70- 6. 08 
0. 43- 0. 49 
9.23 


1. 10 
0.34 


0. 98-1. 28 
0. 31-0. 36 



34 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 12— Euryphorus nordmanni Milne-Edwards, 1840, dorsal view: a, female;^ b, male; 
c, female posterior cephalothoracic sinus and free fourth pedigerous segment, including 
ala. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 35 

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 u-regular, 
lateral siu-face projecting laterally past ventrally swollen medial 
region. Anterior end of segment narrow, bearing pair of heavily 
sclerotized, leglike plates (fig. 13e) 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 structiu-e on male or on members of 
genus Elytrophora, which appears closely related to Euryphorus. 
Posterior sm-face 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. 12a) 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 paii' of large, curled, or uncm'led flaps projecting posteriorly 
past caudal rami. Division between fii'st 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 thu'd "segments" indicated 
by cuticidar constriction, third "segment" short, approximately 9 
percent of abdomen length, median posterior sm'face projecting 
slightly in biconvex anal region. Caudal rami (fig. 13/) ovate, mar- 
gins irregular, distal inner surface plumose; distal sm-face 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. 125) 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 

136) 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. 13c?) 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 
setvdes. 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. 13A) 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. 13i) 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. 13^, j) 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; e, anterior 
end of female genital segment showing projecting plates;/, 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; i, male left antenna;;, mandible. 



COPEPOD CRUSTACEANS — LEWIS 



37 




0.4mm. 9 



Clmm-Jd* 



38 



PROCEEDINGS OF THE NATIONAL MUSEUM 



less than twice the length of fu'st, distal region ciu'ved inward 
slightly, with 12 denticulations on inner surface. Female and male 
postantennal process (fig. 13^^) evidenced only by pah" of nodules 
lateral to antenna base, each with several hauiike processes. Female 
and male postoral process (fig. 13A) spikelike, with broad base, tapered 
to sharp distal end, outer margm sharply mdented medially. Female 
and male maxillule (fig. 13^) nodular, with 3 setule-like processes 
distally. V-shaped heavily sclerotized ridge present posterior to 




0.3 mm 



J-" d 



0.3 mm.cf 



Figure 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; e, male sternal furca. 



mouth cone (fig. 13^), in same region as adhesion pads of pandarids, 
some euryphorids and some caligids. Female and male maxilla (fig. 
14a) 2-segmented, situated lateral and slightly posterior to postoral 
process, arising from padlike projection. First segment approximately 
thi'ee-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, mnermost 
slightly less than twice the length of outermost, with fine membrane 



COPEPOD CRUSTACEANS — LEWIS 



39 



along inner margin; outermost process with frilled membrane along 
outer margin. 

Female maxilliped (fig. 146) 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 sub triangular projections from inner sm-face. Second segment 
and terminal process clawlike, segment short, distinct from terminal 
process, with setule-like accessory process from distal inner siu-face. 
Male maxilliped (fig. 14c) similar to that of female except with addi- 
tional, loioblike projection on proximal inner sm^face of first segment. 
Female sternal furca (fig.l4(/) V-shaped, tines bluntly pointed. Male 
sternal fiu^ca (fig. 14e) with V-shaped sinus but lateral margins almost 
parallel, tines not diverging as in female. 

For nature of thoracic legs and armatm'e, see figiu-e 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-FV of the female and male of 
Euryphorus nordmanni Milne-Edwards, 1840 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




P 
P 




h 


d,dH,2dH, 

P 
3P 






C,P 
2P 




II 


Outer 
Inner 


m 


s,P 


m,p 
m,s 


m.d.fmH 
c,P 


fmH 
c,P 


2fmn,Q,2P 
3P 


c 
P 


c 
c,2P 


3P 
3P 


III* 


Outer 
Inner 


m 


m,P 
2s,rt,P,ni,3s 




d,II 
c,P 


d,h 
c,P 


3h,2P 
3P 


c 

p 


c 
C,2P 


3P 
P 


IV* 


Outer 
Inner 




P 

fm 




d,fmH 


d.froH 
P 


d,fmH,d, 
2fniH 
4P 


c 

p 


C,2P 
C,2P 





♦Numerous hairlike projections (s) in middle of segment, not Included in table, 
t Roughened area (in this case by denticulations) . 

Discussion. — The synonymyzing of Euryjphorus coryphaenae Kr0yer 
with E. 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 Kr0yer's original description of E. coryphaenae so did not give 
an analysis of the two species. 

In a comparison of E. nympha and E. coryphaenae, the major differ- 
ences appear, from the publications, to be in the sternal furca (that of 
E. nympha having more widely spaced tines) and in the armature and 
its position on the first thoracic leg (E. coryphaenae figured as having 



40 



PROCEEDINGS OF THE NATIONAL MUSEUM 




0.3mm.? d* 



Figure 15. — EuTyphorus nordmanni Milne- Edwards, 1840, right thoracic legs, anterior view: 
a, first; h, distal end of second segment of exopodite of first; c, second; J, third; ^.exopodite 
of third (posterior view);/, fourth. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 41 

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 f ureal 
tines, the rather confusing picture given by the terminal elements on 
the second segment of the exopodite of the first thoracic leg (fig. 156), 
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 E. nordmanni Milne-Edwards are : 

1. E. nordmanni has a better developed pair of antennules than 
E. nympha. 

2. The specimens in the Paris museum {E. nordmanni) were 
from the "Asiatic" ocean while E. nympha was described from material 
collected in the subtropical Atlantic. 

Kner (1859) described E. 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 E. nympha. 

Specimens identified as either E. 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 E. nympha (recognizing E. coryp)haenae 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 E. nordmanni has been 
reported only once, from Coryphaenae hippurus, since Kner's publi- 
cation (Ku-tisinghe, 1937). Even this report was later believed 
erroneous, Kirtisinghe (1964 and personal communication) feeling 
that his specimens should have been identified as E. nympha. 

The validity of E. nympha appears to be dependent upon the 
absence of the host name in the original description of E. nordmanni, 
the different locality from which E. nordmanni was collected, and 
the belief that E. nordmanni has a better developed pan- of anten- 
nules. There even appears to have been some question, by Steen- 
strup and Liitken (1861), as to whether their species, E. 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. nymjpha-E. coryphaenae complex indicates, 
to this author, that these 3 species are synonymous. 

Elytrophora Gerstaecker, 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 immatm'e females, 
with pair of posteriorly projecting, lobe-shaped plates, plates straight 
or angled inward slightly. Male genital segment without plates. 
Abdomen 2-segmented, fh'st 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 cm'ved 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 bhamous although endopodite of first and fourth legs re- 
duced; ramal count 2-2, 3-3, 3-3, 3-2. Fifth legs present, setiform. 

Remarks. — The similarity of Euryphorus 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 Euryphorus), and number of seg- 
ments in the rami of the first 4 pairs of thoracic legs (identical in 
members of both genera). 

Elytrophora brachyptera Gerstaecker 

Figures 16-21 

Elytrophora brachyptera Gerstaecker, 1853, p. 60, pi. 3, fig. 12. — Nordmann, 1864, 
p. 468.— Heller, 1866, p. 753.— Van Beneden, lS70a, p. 57.— Richiardi, 
1880, p. 3.— Stossich, 1880, p. 257.— Valle, 1880, p. 60.— Carus, 1885, p. 360.— 



COPEPOD CRUSTACEANS — LEWIS 



43 



Bassett-Smith, 1896a, p. 158; 1896b, p. 12, pi. 4, fig. 3.— Brian, 1899a, p. 4.— 
Bassett-Smith, 1899, p. 462.— Brian, 1906, p. 51, pi. 1, figs. 2-3; 1908, p. 3; 
1912, p. 10.— Scott and Scott, 1913, p. 83, pi. 19, fig. 10; pi. 23, figs. 1-2; 
pi. 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, pi. 123, fig. 1. 

Dinematura Thynni Kr0yer, 1863, p. 157. — Wilson, 1907a, p. 376 (as thynni). 
[Nomen nudum, label name attributed to KoUar.] 

Arnaeus thynni Kr0yer, 1863, p. 157, pi. 8, fig. 5a-g. 



Material.- — 15+ tiost records: 



locality 
Mediterranean 



Northeastern Atlantic 



Subtropical Atlantic 
Pacific 

Western Pacific 



Hawaii 



hosts 
Unknown 
Thynnus vulgaris 
Thynnvs thynnus 
"Germon" 
"Thon rouge" 

Thynnus vulgaris 
Thynnus thynnus 
Orcynus thynnus 
Plankton? 
Thynnus thynnus 
T. alahinga 
Thynnus orientalis 
Parathynnus sibi 
Neothunnus albacora 
Parathunnus obcsus 
Thynnus species 



reference 
Gerstaecker, 1853 
Heller, 1866 
Brian, 1899a 
Brian, 1908 

Delamare-Deboutteville 
and Nunes-Ruivo, 1953 
Van Beneden, 1870a 
Bassett-Smith, 1896a 
Scott and Scott, 1913 
Heegaard, 1955 

Yamaguti, 1936a 

Shiino, 1954a 

Shiino, 1957a 
Bonnet. 1948 



Material.— 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, longhne 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 



Measurements. — (In mm) 10 females and 10 males: 

female 



Total length, excluding caudal setae 

Length of cephalothorax, including frontal 
region 

Width of cephalothorax 

Length of genital segment, excluding gen- 
ital flap 

Width of genital segment 

Length of abdomen 

Length of alae (N = 9 in female) 

Length of genital flap 

Length of caudal rami 

Length of egg strings (4 strings) 



male 



mean 


range 


mean 




range 




8.37 


C. 23- 9. 68 


6.33 


5. 


10-7. 


80 


4.43 


2. 85- 4. 88 


3.45 


3. 


23-4. 


05 


4.51 


3. 48- 4. 95 


3. 10 


2. 


48-3. 


60 


1.93 


1. 13- 2. 37 


1.45 


1. 


18-1. 


70 


1. 70 


0. 72- 1. 95 


L27 


0. 


58-1. 


43 


1.27 


0. 99- 1. 52 


1.04 


0. 


70-1. 


44 


L21 


1. 00- 1. 33 


0.73 


0. 


56-0. 


85 


0.87 


0. 70- 1. 15 










0.57 


0. 49- 0. 63 


0.53 


0. 


44-0. 


63 



9. 13 7. 50-10. 50 



Remarks. — Two sets of descriptions are given below, the first for 
the matui-e 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 behef 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. 16(7) 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- 



COPEPOD CRUSTACEANS — LEWIS 



45 



tral region. Fifth legs (fig. 16/) arising from posterior lateral surface 
of dorsal overlap, consisting of 2 nodules, anteriormost with single 
setule, posterior with 3 setules. 




Figure 16. — Elytrophora brachyptera Gerstieckev, 1853, matuie adult, dorsal view: a, fe- 
male; b, male; c, posterior cephalothoracic sinus. Ventral view: d, female genital segment 
showing spermatophore (sp), abdomen, and caudal ramus; e, male genital segment, abdo- 
men and caudal ramus;/, 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. 166) 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. 16e) 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 setides on distal half of anterior ventral surface and distal ven- 
tral surface. Second segment club shaped, with 14 naked setules 
distally. Female antenna (fig. 176) 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. 176) 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 ciu-ved 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. 17 d) 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. 



COPEPOD CRUSTACEANS — ^LEWIS 



47 




0.5mm. d 



0.5mm. Jd* 



Figure 17. — Elytrophora hrachyptera Gerstaecker, 1853, mature adult, ventral view: a, right 
antennule; h, female oral region, right side showing antenna, postantennal process, mouth 
cone, mandible, maxillule, and postoral process; c, male right antenna; </, left maxilla;^, 
female maxilliped;/, 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. 17e) 2-segmented, situated posterior and 
slightly medial to maxilla base. First segment strongly developed, 
narrow proximally and distally, broad medially, ^^dth 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. 17/) 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. 17 g) V-shaped, tines slender, 
extending posteriorly over interpodal plate of first thoracic legs. 

For nature of legs and armature, see figures 18a-/ 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 Euryphorus nordmanni Milne-Edwards. 

Description. — Immature adult. Female cephalo thorax (fig. 19a) 
ovoid, consisting of cephalon and first 4 thoracic segments. Frontal 
region approximately one-twelfth the total length of cephalothorax, 
'SAdth median cleft and narrow membrane on anterior margin. Lateral 
cephalothoracic margins convex, Avith membrane; mth 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 Avith 
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 Avith niunerous, elongate pliunosities arranged in 2 
gTOups, 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. 



COPEPOD CRUSTACEANS — LEWIS 



49 




Figure 18. — Elytrophora brachyptera Gerstaecker, 1853, mature adult, right thoracic legs, 
anterior view: a, first; b, second segment of exopodite of first; c, second; i, third; f,exopo- 
dite of third (posterior view); /", 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 wddest anteriorly, con- 
stricted medially, broadly rounded posteriorly, covering anterior third 
of genital segment. 

Female genital segment (fig. 19c^) 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, w4th 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, \\ath row of fine denticulations 
around base of outermost seta. 

Male cephalothorax (fig. 196) similar to that of female although 
somewhat narrower, mthout dorsal plumosities or distinct projections 
indicating previous presence. Free fourth pedigerous segment more 
drawn out medially than in female, alae not as large, wdthout medial 
constriction present in female. Genital segment (fig. 19e) basically 
ovoid, without platelike dorsal part and posterior extensions of female; 
fifth leg (fig. 19^) similar to that of female, genital segment \\dthout 
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 smafl, 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, ^\ith 1 naked setule from 
median dorsal surface, second from distal posterior surface, and 8 or 9 
naked setules from distal surface. Female antenna (fig. 20b) 3-seg- 
mented, situated medial and posterior to antennule base. First 



COPEPOD CRUSTACEANS — LEWIS 



51 




iios'm'nn.'? ' 



Figure 19. — Elytrophora brachyptera Gerstaecker, 1853, immature adult, dorsal view: 
a, female; b, male; c, 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; e, male free fourth 
pedigerous segment, genital segment, abdomen, and caudal ramus; /, right fifth leg and 
genital fold of female; g, male right fifth leg. 



52 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. i2i 




0.2 mm.? 



0.3 mm. o- 



Figure 20. — Elylrophora hrachyptera 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;/, 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; i, 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, mth small, pointed 
secondary projection proximally. 

Female and male mandible (fig. 206) 2-parted, first part short, 
tapered distally, \vith indistinct break at beginning of taper. Second 
part elongate, flattened distally, mth indistinct break just proximal 
to flattened part, distal inner surface with 12 denticulations. Female 
and male postantennal process (fig. 206) 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, ^vith 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 margm. 

Female maxilliped (fig. 20/) 2-segmented, situated posterior and 
slightly medial to maxilla base. First segment strongly developed, 
with pair of small, shelflike projections from posterior medial siu-face; 
second segment small, distinct from long, clawlike terminal process, 
bearing setalike accessory process from inner distal sm^face. Male 
maxilliped (fig. 20g) sunilar to that of female although shelflike pro- 
jections of first segment smaller and proximal end of segment more 
complex, "svith heavily sclerotized, clawlike projection from proximal 
inner surface. 

Female sternal fiu-ca (fig. 20h) V-shaped, overlapping interpodal 
plate of first thoracic legs. Male sternal furca (fig. 20i) simflar to 
that of female although tines not as regular and not appearing as 
heavily sclerotized. 



54 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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-/ 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 viale of 
Elytrophora brachyptera Gerstdecker, 1853 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




s,p 
P 




rh§ 
c 


dm,3dmH 
4P 






C 
3p 




II 


Outer 
Inner 


m 


S,P 


m,p 

m,s 


m,d,dmH 
c,P 


dmH 
c,P 


2dmH,Q 
c§,5P 


C 
P 


C 
c,2P 


2P 
4P 


III* 


Outer 
Inner 


m 


fm,dm,P 

dt,2d,5s, 
P,m,4s 




dm.dmll 
c,P 


dm,fmH 
c,P 


3fniH,2P 
c§,3P 


c 
P 


c 
c,2P 


2P 
2P 


IV 


Outer 
Inner 




s§,p 
fm,s§ 




dm.frnH 


fm,dm,fmH 
c§,4pt 


fm.dm, 
fmH,fm, 
2fmH 


c 

Pi 


C,3pt 
Pt 





♦Numerous, hairlike processes on protopodite not tabulated, 
f Denticulations very small. 
JLarger in male than in female. 
§Not present In immature adults. 

Discussion. — 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 furca 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, 

Figure 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); /, fourth. 



COPEPOD CRUSTACEANS — LEWIS 



55 




56 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

can also be found in many other characteristics. The presence of 
pliimosities 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 adidt 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 E. hrachyptera 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 Ltitken, 1861 

Diagnosis. — 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 pak 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. Antenniile 
2-segmented; antenna 3-segmented, with clav/like 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 hau'lilce proc- 
esses; postoral process spinelike, bifurcate, or simple. Maxilliped 
2-segmented, prehensile. Thoracic legs I-III biramous although 
endopodite of first leg reduced ; fom-th 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 wdth 
spinelike accessory process. 



COPEPOD CRUSTACEANS — LEWIS 



57 



Gloiopotes huttoni (Thomson) 

Figures 22-24 

Lepeophtheirns huttoni Thomson, 18S9, p. 354, pi. 28, fig. lOa-c; pi. 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, pi. 21; 1937, p. 429.— Yamaguti, 1963, 

p. 103. 
Caligus longicaudatus Marukawa, 1925, p. 1243, fig. 2396; 1947, p. 927, fig. 2654. 
Gloiopotes watsoni Elirtisinghe, 1934, p. 167, figs. 1-17. — Yamaguti, 1963, p. 104, 

pi. 126, fig. 1. 
Gloiopotes species Yamaguti, 1936b, p. 4, pi. 2, fig. 20; pi. 3, figs. 21-35. 
Gloiopotes zeugopteri Rao, 1951, p. 248, figs. 1-15. 
Gloiopotes longicaudatus (Marukawa) Shiino, 1954b, p. 273, 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, 

pi. 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 mitsukurii 
Martina zelandica 
Istiompax australis 
Histiophonis brevirostris 
H. gladius 
Xiphias zeugopteri 
Makaira indica 
M. mazara 

Tetrapturus mitsukurii 
T. marlina 
"Swordfish" 

Istiophorus greyi "Marlin" 
"Black Marlin" 
Makaira audax 
Makaira species 
Xiphias gladius 
"Marlin" 

Tetrapturus mitsukurii 
Parathynnus sibi 
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 



Material. — 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 audax (Philippi) captured off Waianae, Hawaii. 



58 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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 
12. 02 



range 
10. 20-12. 90 



mean 
10.25 



range 
9. 53-10. 95 



5. 81 5. 03- 6. 08 

5. 06 4. 35- 5. 63 

3. 21 2. 40- 3. 60 

3. 03 2. 70- 3. 30 

2. 29 1. 67- 2. 55 
1. 76 1. 33- 2. 11 

1. 61 1. 41- 1. 78 

1. 06 0. 89- 1. 30 

3. 34 2. 40- 4. 65 



2. 16 1. 95- 2. 25 

1. 94 1. 88- 2. 03 

1. 83 1. 52- 2. 04 

1. 08 0. 93- 1. 15 



Total length, excluding caudal setae 
Length of cephalothorax, including 

frontal region 5. 81 5. 03- 6. 08 4. 63 4. 35- 4. 88 

Width of cephalothorax 5. 06 4. 35- 5. 63 4. 00 3. 60- 4. 28 

Length of genital segment, excluding 

fifth legs 
Width of genital segment 
Length of abdomen 
Length of alae (male N = 5) 
Length of fifth leg, measured along 

outer margin (male N = 5) 1. 61 1. 41- 1. 78 1. 56 1. 30- 1. 63 

Length of caudal rami 1. 06 0. 89- 1. 30 1. 23 1. 04- 1. 37 

Length of egg strings (12 strings) 

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 smiace. 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 projecting mem- 
brane, from dorsal lateral surface. Median cephalothoracic region 
extending to or slightly past posterior end of lateral cephalothoracic 
regions, margin irregular. Dorsal siuface of cephalothorax Avith 
numerous haulike projections laterally, gi\^ng fuzzy appearance; 3 
rows of long, slender, bifid plumosities present on lateral regions, 
lateral to ocular region. Dorsal surface also \\\ih 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. 



COPEPOD CRUSTACEANS — LEWIS 



59 




Figure 22.— Glotopotes 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: e, female genital segment, fifth leg, abdomen, 
and caudal ramus;/, male genital segment, fifth leg abdomen, and caudal ramus. 



60 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Female free fourth pedigeroiis 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 spmule; 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 mth club-shaped 
swelling on dorsal suiface. Both segments mth 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 mth single 
spine, distal ventral surface mth 1 and distal sm'face 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. 22/) distinct 
from fourth pedigerous segment, ovoid in outline, ^^dthout lobate 
posterior projections. Heavily sclerotized fifth legs projecting from 
posterior ventral surface, mthout 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 altliough inner terminal process longer. 

Female and male antennule (fig. 23a) 2-segmented, first segment 

Figure 23. — Gloiopoies huitoni (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; /, male, same, right side; g, right maxilla (pop= 
postoral process; a = adhesion pad, not present in male); h, left maxilliped. 



COPEPOD CRUSTACEANS — LEWIS 



61 



|[lIE2S 




0,5mm. gd* 0.5 mm.? d* 



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. 236) 3-segmented, situated posterior and medial to anten- 
nule base. First segment sub triangular, small, with roughened, 
triangular projection from proximal posterior surface. Second seg- 
ment almost twice the size of first, with small, lappet-Hke 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 e) indistinctly 3-partied, rodlike; thu'd 
part flattened, distal inner surface mth 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, \\^th 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. 23c) similar to 
that of figured female although lacking smaller of 2 single spinelike 
projections. Female postoral process (fig. 2Se) bifurcate, each ramus 
spinelike; additional, minute, sharply pointed projection present 
just lateral to bifurcation. Male postoral process (fig. 23/) spinelike, 
not bifurcate 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. 23/) nodular, with 2 or 3 setules. Female and 
male maxilla (fig. 23g) 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; 



COPEPOD CRUSTACEANS — LEWIS 



63 



distal surface bearing 2 saber-shaped processes, inner longer than 
outer, both with fine membranes along margins. 

Female maxilliped (fig. 23/0 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; fu'st 
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 maxilhped 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 dis tally, 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 246-(/ and table 9. 

Table 9. — Armature of thoracic legs I-IV of the female and male of 
Gloiopotes huttoni {Thomson, 1889) 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




sss.p 

p 




h 
c 


3P,p,3dH 




d* 


2p 
P' 




II 


Outer 
Inner 


m 


S,P 


m,p' 
m,s 


m.fmHf 
c,P 


dH 
c,P 


dH,H,Q,P 
4P 


C 
P 


c 
c,2P 


c,4P 
2P 


III 


Outer 
Inner 


m 


s,m,p 

2ss,P, m, 
2s,C§ 




H 
P 


H 
c,P 


3H,2P 
C,3P 


C 
0,2 P 


C,2P 
2P 




IV 


Outert 




li,pll 




rh,3h, 


D,H 


D,3H,D 









*Denticulations in clump in female, in row in male, 
t Spine denticulated In male, not in female. 

X Numerous spinules on protopodite and first segment of exopodite not tabulated. Spinule arrangement 
slightly different in two sexes. 
§ Elements between rami. 
II 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 mth 
the postantennal process. In the figured female specimen (fig. 2Zd), 



64 



PROCEEDINGS OF THE NATIONAL MUSEUM 




0.5mnn.?cf 



05 mm.?d' 



Figure U.—Gloiopotes huttoni (Thomson, 1889): a, sternal furca, ventral view (male with- 
out pads lateral to furca). Thoracic legs, anterior view: b, right first; c, second segment 
of exopodite of right first; d, endopodite of right first; e, right second;/, right third; g, left 
fourth. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 65 

the process has a single, posteriorly du-ected 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 figm-ed by Shiino (1954b, fig. 
le) and by Hewitt (1964a, fig. 13), having only the posteriorly di- 
rected spine and the bifid inwardly directed spine. In the same female 
specimen, the postantennal process on the right side has a posteriorly 
du-ected 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 2Sd 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 (Hemtt, 
1964a). Hewitt, however (p. 93), states that "the shape of the 
plates [alae] on the fom-th 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 vnih 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 (HcAvitt, 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 specunens and, from an examination of 
material in the U.S. National Musemn, 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 figiu-ed 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 specunens of G. huttoni (the variation in the number 
and position of spinules and the shape and mclination 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 
thii'd 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 

226-321—67 5 



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. huttoni 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. huttoni is the species of Gloiojwtes that is found 
on biUfishes 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. huttoni). 

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, pi. 5, fig. 9. — Bassett- 
Smith, 1899, p. 458.— Stabbing, 1900, p. 670, pi. 74, fig. A.— Shiino, 1960b, 
p. 533, figs. 4-6.— Yamaguti, 1963, p. 103, pi. 124, fig. 3. 

Gloiopotes species Bonnet, 1948, p. 7. 

Distribution and hosts. — 4 hosts: 



locality 


hosts 


reference 


itlantic 


Unknown 


Steenstrup and Liitken, 1861 


acific 


"Albacore" 


Stebbing, 1900 




Acanthocybium solandri 


Shiino, 1960b 


Hawaii 


Acanthocybium solandri 


Bonnet, 1948 



Material. — 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 1 12883) from A. solandri 
(Cuvier) captm-ed 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 



Length of cephalothorax, including 

frontal region 
Width of cephalothorax 
Length of genital segment, including 

posterior projections but not fifth 

legs 
Width of genital segment 
Length of abdomen 
Length of alae 
Length of fifth leg, measured along 

outer margin 
Length of caudal rami 
Length of egg strings (4 strings) 

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 smaU, posteriorly projecting membrane from posterior haK and 
fine membrane along aU of margin, membrane projecting across 
most of sinus. Anterior end of sinus sharply rounded, with smaU, 
lappet-like, posterodorsaUy 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. 



mean 


range 


mean 


range 


14.33 


13. 65-15. 15 


9.92 


9. 38-10. 35 


6.85 


6. 68- 7. 13 


4.82 


4. 50- 5. 25 


5.44 


5. 25- 5. 78 


3.32 


3. 08- 3. 53 


5.37 


4. 73- 6. 15 


1.86 


1. 73- 2. 03 


2.99 


2. 63- 3. 30 


1.63 


1.58- 1.65 


3. 15 


2. 89- 3. 37 


1.69 


1. 52- 1. 78 


3.68 


3. 44- 4. 00 


L56 


1. 52- 1. 59 


1. 51 


1. 41- 1. 67 


L43 


1. 30- 1. 52 


1.47 


1. 26- 1. 59 


1.57 


1. 41- 1. 78 


12.32 


10. 95-13. 88 







68 



PROCEEDINGS OF THE NATIONAL MUSEUM 




Figure 25. — Gloiopotes hygomianus Steenstrup and Lutken, 1861, dorsal view: a, female; 
b, male. Ventral view: c, female genital segment, abdomen, and caudal ramus; d, male, 
same; e, tip of female left fifth leg; /, 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 pak 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. 256) generally similar to that of female 
although more elongate. Apex of posterior sinus Avithout lappet- 
like membrane although remaining membranes present; cross groove 
of major dorsal grooves with indistinct apex. Ala of free fourth 
pedigerous segment Avith 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. 25/) with 3 spinules, proximal 
inner surface with minute, pkmiose 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 



PROCEEDINGS OF THE NATIONAL MUSEUM 




0.5mm? d^ I' 



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, Avith 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 flaphke 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 setide-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 distaUy, 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 siu"- 
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. 26c) large, spinelike, female with small spine arising from 
inner proximal surface, process absent on male. Female and male 
maxillule (fig. 26c) nodular, with setule distally. Female with pair 

Figure 26. — Gloiopotes hygomianus Steenstrup and Lutken, 1861, ventral view: a, left 
antennule; h, female left antenna; c, male left antenna and postantennal process; d, fe- 
male right postantennal process; e, oral region, left side showing mouth cone, mandible 
(mdbl), maxillule (ma-1), postoral process, maxilla base (ma-2), and postoral adhesion 
process (pa);/, left maxilla; g, female right maxilliped; h, male left maxilliped; t, sternal 
furca. 



72 



PROCEEDINGS OF THE NATIONAL MUSEUM 



of small, knob-shaped projections posterior to mouth cone base, 
similar in position to postoral adhesion pads. Female and male 
maxilla (fig. 26/) 2-segmented, situated lateral and posterior to post- 
oral process. Fhst segment approximately two-thu-ds 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, Avith fine membrane along inner 
margin, outer with membranes along both margins. 

Female maxilliped (fig. 2Qg) 2-segmented, situated posterior and 
medial to maxilla base. First segm.ent 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. 26A) 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 distaUy, 
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. 26'i) situated between and 
posterior to maxiUiped bases, consisting of plate bearing 2 widely 
separated, bluntly pointed tines and 2 accessory spines, at base of 
tines. Pah- of lappet-like projections present just posterior and 
lateral to furca. 

For natiu-e of armatm-e 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 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodlte 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 

c,2P 
P 


3 


I 


Outer 
Inner 




ss,p 
P 




h 

c 


3P,p,3dH 




c 




II 


Outer 
Inner 


m 


S,P 


m,p' 
m,s 


2m,dH 
c,P 


dH 
c,P 


dH.mH, 

Q,P 
4P 


c 
P 


c 
c,2P 


c,4P 
2P 


III 


Outer 
Inner 


m 


m,p 

2s,P,m,2s, 
c* 




3s,H 
P 


h 
P 


3h,2P 
3P 


c 
c,2P 


3P 
P 




IV 


Outert 




Pt 




d,D,dH 


D,dH 


D,3£in,3dH 









•Elements between rami. 

fElement in middle of segment. 

tSpinules along iimer surface not tabulated. 




0.5mm.?d'' g 



Figure 27. — Gloiopotes hygomianus Steenstrup and Liitken, 1861, right thoracic legs, ante- 
rior view: a, first; b, endopodite of first; c, distal end of second segment of exopodite of first; 
d, second; e, third;/, 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 tilua Lewis, 1964b, p. 351, figures 2-4; 12a, e, i; 13a. 
Hosts and distribution. — 1 host record: 

location hosts reference 

Hawaiian Islands Caranx rnelampygus? Lewis, 1964b 

Material. — Two females and 1 male (retained by author) from 
external sm"face of Caranx 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: 

Total length, excluding fifth legs and caudal setae 
Length of cephalothorax, including frontal region 
Width of cephalothorax 
Length of genital segment, including lobes but excluding 

fifth legs 
Width of genital segment 
Length of abdomen 
Length of caudal rami 

Length of fifth legs, measured along outer margin 
Length of fifth leg terminal process 
Length of sixth leg, measured along outer margin 
Length of sixth leg terminal process 

Description. — See Lewis, 1964b, 

Dentigryps bifurcatiis Lewis 

Dentigryps bifurcatiis Lewis, 1964a, p. 203, figs. 17a-c, e-k, m-q; 18a-c, e-f; 1964b, 
p. 356, figs. 5; 12b, f, j; 13b. 

Distribution and hosts. — 5 host records: 

locality hosts reference 

Hawaiian Islands Acanthurus olivaceus 

A. triostegus 

sandvicensis Lewis, 1964a 

Naso hexacanthus 
Fistularia petimba Lewis, 1964b 

Material. — Three males (retained by author) from external 
surface of Bodianus bilunulatus (Lacepede) from the Honolulu 
Aquarium. One male (retained by author) from external surface 
of Chaetodon jremhlii Bennett captured in a fishtrap by vSamuel 
Kaolulo between Diamond Head and Koko Head, Oahu, Hawaii. 



female 


male 


6. 60, 6. 83 


5. 18 


4. 43, 4. 43 


3.45 


3. 40, 3. 66 


2.92 


1. 67, 1. 67 


0.93 


1. 70, 1. 70 


1.04 


0. 61, 0. 76 


0.49 


0. 31, 0. 29 


0.29 


1. 19, 1. 28 


0.99 


0. 05 


0.06 




0. 15 




0. 14 



NO. 3574 COPEPOD CRUSTACEANS — ^LEWIS 75 

Two immature males (retained by author) from external surface of 
Aulostomus chinensis (Linnaeus) captiu-ed 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 1.58, 1.55, 1.48, 1.53 

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 0.12, 0.11, 0.11, 0.12 

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, 0.13, 0.14 

Description. — See Lewis, 1964a. 

Anuretes Heller, 1865 

Diagnosis. — See Lewis, 1964a. 

Anuretes serratus Shiino 

Anuretes serratusShiino, 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 

Description. — 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 

Diagnosis. — Cephalothorax consisting of cephalon and first 4 
thoracic segments; frontal region distinct, without lunules. Free 
fom-th 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 
sm-face. 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. MaxUlule nodular, with setule-like processes; 
maxilla 2-segmented, with 2 saber-like terminal processes. Sternal 
fiu-ca present. Thoracic legs I-III biramous although endopodite of 
first legs rudimentary; fourth legs uniramous, typically with 4 
segments. 

Lepeophtheirus crassiis (Wilson and Bere) 

Figures 28-30 

Gloiopotcs crassus Wilson and Bere, in Bere, 1936, p. 590, pi. 5, figs. 109-111; pi. 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 V/ilson and Bere, 1936 

Eastern Pacific "Remora" 

Remilegia australis Shiino, 1963a 

Indian Ocean Echeneis albescens Shiino, 1960c 

Material. — Five females (USNM 112884) from external siu-face 
of Rhombochirus osteochir (Cuvier) taken from Tetrapturus angusti- 
rostris Tanaka captm-ed by longline at 21°04.5'N, 173°47.5'E (USF 
WS). Two females and 1 male (USNM 112885) from external siu-- 
isice oi Rhombochirus osteochir (Cuvier) taken at20°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. SO 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 



COPEPOD CRUSTACEANS — LEWIS 



77 



Description. — Female cephalothorax (fig. 28a) ovoid, consisting of 
cephalon and first 4 thoracic segments. Frontal region narrow, 




Figure 28. — Lepeoph/heirus crassus (WihonandBere, 1936), dorsal view: a, female; i, male. 
Ventral view: c, female genital segment, fifth leg, abdomen, and caudal ramus; J, male, 
same; e, male tip of fifth leg;/, 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 
groove 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. 2Sc) 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 dactyhform although sHghtly u-reg- 
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 plimiose 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 ventraUy, V-shaped dorsaUy, 
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; knobUke projection present on inner distal 
surface, bearing minute, spikehke terminal process. 

Cephalo thorax 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 
flrst abdominal segment slightly. Fifth legs dactyUform, originating 
on anterior-medial ventral surface, extending posteriorly and shghtly 
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. 28/) small, somewhat irregular, projecting shghtly 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 distaUy 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 
smaU, 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 weU 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 




C *~o.2mm. ? / o.lmm.'d ^ 0.2mm. ? /^ 0.1mm. c^ 




0.2mm. 9 




] 0.2mm. 6 



Figure 29. — Lepeopluheirus crassus (Wilson and Bere, 1936), ventral view: a,leftantennule; 
b, female right antenna ; c, male, same; ^, right mandible; <?, female left postantennal process; 
/, male, same; g, female right maxillule and postoral process; h, male, same; i, female post- 
oral adhesion pads, postoral processes, posterior end of mouth cone and maxilliped bases 
(mb);/, 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 swelhng 
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 denticidations 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, cm'ving medially sharply in male, 
slightly in female, sharply rounded, or pointed, distally. Female 
postoral process (fig. 29g) bifurcate, both tines pointed; male postoral 
process (fig. 29h) spinelike, with small spine projecting from middle of 
outer surface and small, heavily sclerotized swelhng on middle of 
inner surface. Female and male maxillule (figs. 29g, 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 VA 
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. 29/) extend- 
ing posteriorly and medially from just posterior to postoral processes 
to just anterior to maxiUiped bases, surfaces distinctly rugose. Female 
also with surfaces but not rugose (fig. 29i). 

Female maxilliped (fig. 306) 2-segmented, situated posterior and 
shghtly 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 shghtly greater than half 
that of first segment, terminal process clawhke, sharply pointed; 
single small, setalike accessory process present on inner surface at 
irregular division between segment and terminal process. Male max- 
ilhped (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— 67— !— 6 



82 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




0.3mm. d? 



0.05mm. d? 



COPEPOD CRUSTACEANS — LEWIS 



S3 



segment flexed. Terminal process of second segment not as sharply 
curved as in female, accessory process slightly longer. 

Female and male sternal furca (fig. 30c^) 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) 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodlte 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss,p 
P 




rh 
c 


3H,P 
3P 










II 


Outer 
Inner 


m 


S,P 


m,p 
m,s 


m,fmH 
c,P 


fniH 
c,P 


H,qH,Q, 
c,3P 


c 
P 


c 
c,2P 


c,3P 
c,3P 


III 


Outer 
Inner 


m 


m,P 

P,s,m,lls* 




3s 
p,mH 


c,p' 
c,P 


c,3p',P 
c,3P 


c 

p 


c,2p 
c,4P 




IV 


Outer 




s 




dm,h 


dm.fmH 


3dm,3fmH 









*0n both anterior and posterior surfaces of protopodite. 

Lepeophtheirus dissimulatus Wilson 

Lepeophtheirus dissimulatus Wilson, 1905b, p. 631, pi. 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 
Bermuda 



Dry Tortugas 
Galapagos Islands 

Eastern Pacific 



hosts 
Epinephalus morio 
E. striatus 
Mycteroperca apua 
Thynnus pelamys 
Lactophrys triqueter 
Epinephelus labriformis 
Mycteroperca species 
Bodianus diplotaenis 
Epinephelus labrifrons 
Merluccius productus 
Paralichthys californicus 
Hypsopsetta gattulata 
Sphyraena argentea 



reference 
Wilson, 1905b 

Linton, 1907 
Heegaard, 1943b 
Wilson, 1935b 
Wilson, 1905b 
Heegaard, 1943b 

Shiino, 1959d 



Figure 30. — Lepeophtheirus crassus (Wilson and Bere, 1936), lateral view: a, maxilla; b, 
female maxilliped; c, male maxilliped; d, sternal furca (ventral view). Right thoracic 
legs, anterior view: e, first; /, second segment of exopodite of first; g, second; h, third; i, 
exopodite and endopodite of third; ;", fourth; k, 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 



locality 



Hawaii 



Mauritania 



hosts 
Paralahrax nehulijer 
Sphaeroides annulatus 
Galeichthys guatemalensis 
Gadus macrocephalus 
Acanthurus olivaceus 
A. dussu7nieri 
A. triostegus sandivcensis 
Zebrasoma flavescens 
N'aso hexacanthus 
Chaetodon quadrimaculatus 
Labrus species sensu lato 



reference 



Causey, 1960 
Lewis, 1964a 



Lewis, 1964a 
Brian, 1924 



Material. — 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. 53-2. 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. lO-O. 12 


Length of caudal rami 






0.08 


0. 07-0. 09 


0. 10 


0.08-0. 11 


Length of egg strings (2 st 


■ings) 


1.00 


1.48 








Description. — See Lewis, 


1964a 











Lepeophtheirus? falloliinulits, new species 

Figures 31-33 

Material.— One female (holotype, USNM 112886) from the gill 
cavity of Naso unicornis (Forskal) speared off Lehua Rock, Niihau, 
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: 

Total length, excluding caudal setae 

Length of cephalothorax, including frontal region 

Width of cephalothorax 

Length of genital segment 

Width of genital segment 

Length of abdomen 

Length of caudal rami 

Length of egg strings 



female 


male 


3.05 


2.25 


1.43 


L30 


1.30 


L23 


LOS 


0.54 


0.85 


0.45 


0. 15 


0. 14 


0.07 


0.08 


1.20 





COPEPOD CRUSTACEANS — LEWIS 



85 




0.3 mm. d* 



O.OSmm.cT 



Figure 31. — Lepeophtheirus} fallolunulus, new species, dorsal view: a, female; b, immature 
adult male; c, posterior cephalothoracic sinus. Ventral view: d, female genital segment, 
abdomen, and caudal ramus; e, male, same;/, female right fifth leg; g, male, same; h 
male right sizth leg. 



86 



PROCEEDINGS OF THE NATIONAL MUSEUM 




NO. 3574 COPEPOD CRUSTACEANS — LEWIS 87 

Description. — Female cephalo thorax (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 sm-face median to modi- 
fication. Lunular modifications (fig. 326) with denticulated, dorsally 
facing anterior margin, beaiing 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 fiange 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 cephalothoracic grooves 
running longitudinally, cross groove absent. Free fourth pedigerous 
segment slender, without dorsal plates. 

Female genital segment (fig. 31f/) ovoid, indistinctly separable from 
free fourth pedigerous segment. Posterior end of segment irregular, 
irregularities primarily in region of oviducal openings. Fifth leg 
(fig. 31/) 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. 316) 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 fovirth 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 

Figure 32. — Lepeophtheirust 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-1), postoral process (pop), postoral adhesion process (pa), 
and maxilla base (ma-2); e, male oral region, right side showing antenna (stippled), 
postantennal process, mouth cone (hatched), mandible (hatched), maxillule (ma-1), 
postoral process (pop), postoral adhesion process (pa) and maxilla base (ma-2); /, 
right maxilla; g, female left maxilliped; h, male right maxilliped. 



88 



PROCEEDINGS OF THE NATIONAL MUSEUM 




0.2 mm.? d" 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 89 

segment without plates. Segment-like projection present between 
fom'tli pedigerous and genital segTnents, separable from fourth pedig- 
erous segment (at least superficially) both dorsaUy and ventrally, 
distinct from genital segment; projection without processes. Genital 
segment (fig. 31g) barrel shaped, fifth and sLxth legs (figs. 31^, 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 
segment 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. 32c?) 
3-segmented, situated posterior to subtriangular projection associ- 
ated with antennule and frontal region. Fu-st 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 sub terminal process and setule-like accessory 
process. 

Female and male mandible (figs. ?,2d, e) 4-parted, rodlike, distal 
part curving inward, inner surface with 12 denticulations. Female 
and male postantennal process (figs. ?>2d, e) consisting of sharply 
curved, spinelilce process, with outer margm denticidated distally, 
in addition to 3 nodules, 2 on base of spinelike process, third posterior 
to process, each with several hau'lil^e processes. Female postoral 
process (fig. ?>2d) formed of heavily sclerotized plate bearing small 
bifurcate projection, tines rounded distally. Male postoral process 
(fig. 32e) small, lappet-lil<;e projection of heavily sclerotized ridge. 
Female maxiUule (fig. 32(/) 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 fii'st segments of antennae; postoral process, 
antenna base, mouth cone, maxillule, and maxilla in close proximity. 

Figure 33. — Lepeophtheirust 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 



Small, heavily sclerotized, knoblike postoral adhesion process present 
in both female and male (figs. 32c?, e), arising from Y-shaped region 
of heavy sclerotization, just posterior to postoral process. Female 
and male maxilla (fig. 32/) 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. 32^, h) 2-segmented, situated 
medial and posterior to maxilla base. Fu'st 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, h) situated well posterior to maxilliped 
bases. Bifurcate portion of female furca at end of necklike process, 
bifurcate 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-y 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, neiv species 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




P 
P 




rh 
C 


H,2dH,P' 
3rh 




P' 
P' 






II 


Outer 
Inner 


ra 


s,P 


m,p 
m,p' 


m,Ht 
c,P 


h 
c,P 


h,mh,Q,2P 
3P 


C 
P 


c 
c,2P 


C,3P 
3P 


III 


Outer 
Inner 


m 


Dt,s,a*,m,p 
r§,2s,P,m 




H 
P 


P,p,2P 
4P,p 




c 


c,3P 
3P 




IV 


Outer 




pll 




s,H 


3mH,H 











*Five-lobed adhesion pad. 

tSingle large denticulation. 

JRamitied spine; female with 6 rami (excluding tip), male with 7. 

sThrec roughened areas. 

liSeveral hairUke processes not tabulated. 

Discussion. — The species is tentatively placed m 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 kmule-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 foiu-th 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 leg of members of 
this genus is reduced or rudimentary but it is present (e.g., L. dissimu- 
latus Wilson, 1905b; L. gonistii Yamaguti, 1936a) and the leg is 
biramous. 

5. The postoral process is reduced, especially in the male. 

The lunule-like structure of L. jallolunulus 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 sunple (e.g., C. enormis Wilson, 1913) 
or the membrane can be well developed and the sinus remain open 
(e.g., C. zei 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 L. jallolunulus there is no sinus, the structure is 
directed dorsally and not ventrally, and it is found onl}^ 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, aniennule, 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. jallolunulus 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 Shiino, 1954a. 

The name "falloluniiliis" is derived from the Latin terms "fallo" 
(deceive) and "lunula" (small moon) and refers to the natm-e of the 
lunule-lilce structm'e on the female. 

Midias Wilson, 1911b 

Diagnosis. — Cephalothorax ovoid, consisting of cephalon and first 
4 thoracic segments. Frontal region distinct, with lunules. Free 
fom-th pedigerous segment wider than long, tergal region heavily 
sclerotized (as is most of the dorsal body surface), with flapiike 
lateral extensions, segment without alae or separate plates in either 
sex. Genital segment with short, heavily sclerotized, spinelike fifth 
leg projections on posterior lateral siu-face. 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 hau'like processes; postoral process spine- 
like, either bifid or simple, male with adhesion siu-face 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, foiu-th 
thoracic leg uniramous. 

Remarks. — Two species have been described in this genus and 
are compared in the discussion section following the description of 
M. lohodes. 

Wilson (1911b, p. 628) placed M. lohodes 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-segmented 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 Tiixophorinae, which includes both 
Midias and Tuxophorus, the latter of which has distinct alae on the 
fourth pedigerous segment. 

The division between the Cahgidae and the Euryphoridae is tenuous, 
both famihes sharing common characteristics. Dentigryps, a caligid 
genus, for example, possesses euryphorid-like fifth legs; Pupulina, a 
caligid, 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. 

Midias, 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). 
Midias 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 DarteveUia, and the posterior lobes on the abdomen are pres- 
ent in Dentigryps curtus Wilson, 1913, both of the last two being 
caligids. 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. 



94 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Midias lobodes Wilson 

Figures 34-36 

Midias lohodes Wilson, 1911b, p. 626, pi. 65; 1913, p. 225.— Causey, 1953b, p. 
11.— Shiino, 1958, p. 98, figs. 1-3; 1963a, p. 343.— Yamaguti, 1963, p. 107, 
pi. 127, fig. 1. 

Distribution 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 

Material.— One female and 1 male (USNM 112888) from the ex- 
ternal surface of Sphyraena barracuda (Walbaiim) 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). 

Measurements. — (In mm) 1 female and 3 

Total length, excluding caudal setae 

Length of cephalothorax, including frontal region 

Width of cephalothorax 

Length of genital segment, excluding fifth legs 

Width of genital segment 

Length of abdomen 

Width of first segment of abdomen 

Length of fifth leg 

Length of caudal rami 

Description. — Female cephalothorax (fig. 34o) 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. 



lales : 








female 

12.00 

4.73 

4.35 


7.13, 
3.68, 
3.38, 


male 
6.90, 
3.68, 
3.38, 


6.30 
3.38 
3.08 


3.29 
3.33 
3.29 


1.55, 
1.81, 
1.70, 


1.44, 
1.92, 

1.78, 


1.33 
1.85 
1.67 


3.11 
0.27 


0.48, 


0.54, 


0.56 


0.35 


0.29, 


0.31, 


0.28 



COPEPOD CRUSTACEANS — LEWIS 



95 




0.2mm.? ■^' 



0.2mnn.d' 



Figure 34. — Midias lohodes Wilson, 191 1, dorsal view: a, female; h, male; c, posterior cepha- 
lothoracic sinus. Ventral view: d, female genital segment; e, male genital segment. 
Dorsal view:/, female right fifth leg; g, male, same. Ventral view: h, male right sixth 
leg; 1, female caudal ramus and abdominal projection; /, 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. Md) 
broader posteriorly, tapered regularly from slightly narrower anterior 
end, lateral posterior surface slightly lobate. Fifth legs (fig. 34/) 
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 plimiose, 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 fomth pedigerous seg- 
ment and abdomen, lateral surfaces broadly convex, fifth legs (fig. 34/) 
spikelike, projecting from posterior lateral sm-faces, 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. 34^) present medial to fifth, consisting of 
node bearing 2 plumose setules distaUy. (The condition of the fiftli 
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. 356) 2-segmented, attached to 
ventral-lateral anterior surface of cephalothorax, just posterior to 
division between frontal region and remaining cephalothorax. First 
segment approximately IK times the length of second, distal half of 
anterior and anterior ventral surface bearing approximately 26 hghtly 
plumose setae and setules. Second segment elongate, slightly longer 
in male than in female, with single naked setule from posterior medial 
surface, 1 1 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 subcorneal pro- 
jection from posterior surface. Second segment short, subrectangular ; 



COPEPOD CRUSTACEANS — LEWIS 



97 




Smnn.?^ 



Figure 3S.-Midias lobodes Wilson, 1911, ventral view: a, right side of frontal region show- 
ing lunule; b, 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; e, male right postantennal process;/, male right postoral process and max- 
illule; g, left maxilla; h, right maxiJliped; i, sternal furca. 



226-321—67-^—7 



98 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

third segment fused with clawUke 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. S5d) basically similar to that of female although segments 
and processes relatively longer and first 2 segments bearing adhesion 
surfaces. 

Female and male mandible (fig. 35c) 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. 35c) 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. 35/) 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. S5g) 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 IK times the length 
of outermost, both with very fightly frilled membranes along both 
inner and outer margins. 

Female maxilliped (fig. 35A) 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. 35i) situated between and posterior to maxilliped 
bases, associated with heavily sclerotized transverse band on cepha- 
lo thorax. 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 3Qa-h and table 13. 



Figure 36. — Mz(^taj-/oioJifj Wilson, 1911, right thoracic legs, anterior view: fl, first; b, distal 
end of second segment of exopodite of first; c, second; d, third; e, exopodite of third (pos- 
terior view);/, first segment of exopodite of female third; g, male fourth; h, protopodite of 
female fourth. 



COPEPOD CRUSTACEANS — ^LEWIS 



99 




100 



PROCEEDINGS OF THE NATIONAL MUSEUM 





Table 13 — 


-Armature of thoracic legs I-IV of the female and male of 
Midias lobodcs Wilson, 1911b 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodlte 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss,p 
P 




fm.rh 
C 


3fm,3dH, 

H 
3P 






ss 




II 


Outer 
Inner 


m 


s,P 


m,p 
m,s 


m,fm, 

dmll 
c,P 


dmH 
c,P 


2dmH,Q, 

P 
c,4P 


c,C 
P 


c 
c,2P 


C,3P 
C,3P 


III 


Outer 
Inner 


m 


a,d,ni,p 

d,3s,P,s, 
m,s 




s.fmH 


3s,c,p' 
c.P 


c,3p',P 
c,3P 


c 
P 


C,2P 
C,4P 




IV 


Outer 




s,ss,s*(pt) 




2s,fm,fmH 


fm.fmH 


3fm,3fmH 









•Female only. 
fMale only. 



Discussion. — 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 hairhke processes, as 
well as a spinule-)ike 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-seginented 
although in all of the specimens examined, including the type mate- 
rial, the endopodite is 2-segmented. There is a cuticular break in 
the second segment in all of the specimens (fig. 36^) 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 thu'd thoracic leg in M. carangis; this 
claw is bifid in M. lobodes. 

Caligus MuUer, 1785 

Diagnosis. — Cephalothorax consisting of cephalon and first 4 
thoracic segments; frontal region distinct, with Imiules. Free fom-th 
pedigerous segment without alae, sometimes with small, lappet-like 
lateral projections of tergum extendmg over proximal sm-face of fom'th 
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 
fom-th 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 hau'like 
processes, usually with spinelike projection. Postoral process spine- 
like, simple or with small accessory spine. MaxUlule 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 

Figures 37-39 

ICaligus scutaius Milne-Edwards, 1840, p. 453. 

Caligus coryphaenae Steenstrup and Liitken, 1861, p. 360, pi. 4, fig. 7. — Richiardi, 
1880, p. 148.— Valle, 1880, p. 58.— Cams, 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, pi. 59, fig. 3. 

Caligus bengoensis Scott, 1894, p. 130, pi. 14, fig. 19. 

Caligus aliuncus Wilson, 1905b, p. 576, pi. 9; 1935b, p. 330.— Bonnet, 1948, 
p. 7. — Causey, 1953b, p. 8. 

Caligus elongatus Heegaard, 1943b, p. 11, figs. 21-31. 

Caligus iesserifer Shiino, 1952, p. 89, fig. 5. 

Not Caligus coryphaenae. — Brian, 1935, p. 51, figs. 19, 20. — Yamaguti, 1936a, 
p. 5, pi. 4, figs. 40-54.— Barnard, 1955, p. 246, figs. 8a-d. 



102 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Distribution and hosts. — 19 + host records: 



locality 
Subtropical Atlantic 



Gulf of Mexico 
Eastern Pacific 

Hawaii 

Western Pacific 
Indian Ocean 

Mediterranean 



hosts 
Coryphaena 
"Bonito" 
C. hippums 
Plankton 
Parathunus obesus 
Euthynnus alleteratus 
Coryphaena hippurus 
Sarda species 
Coryphaena hippurus 
Euthynnus linealus 
Katsuwonus pelaniis 
Euthynnus alleteratus 
Coryphaena hippurus 
Katsuwonus pelaniis 
Coryphaena hippurus 
Unknown 
Squalus acanthias 
Neothunnus albacora 
Coryphaena hippurus 



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 



Material.— 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 surf ace of Ti. ^eZamis (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 at20°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, Hawaii 
(USFWS). One female and 2 males (USNM 112895) from the exter- 
nal surface of Euthynnus 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 Euthynnus 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 Katsuwonus pelamis 
(Lmnaeus) collected at 2°S, 132°W (USFWS, JRM cruise 34, station 
5). One male (USNM 112899) from the external surface of Euthyn^ 
nus yaito (J. and E.) collected in imknown 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 


L77 


1. 30-2. 11 


0.99 


0. 81-1. 11 


Width of genital segment 


L58 


1. 33-1. 81 


1.25 


1. 11-1.33 


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 cephalo thoracic 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 sm-face 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 slightly, 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 




Figure 37. — Caltgus coryphaenae Steenstrup and Liitken, 1861, dorsal view: a, female; b, 
male; c, posterior cephalothoracic sinus. Ventral view: d, female free fourth pedigerous 
segment, genital segment, and anterior end of abdomen; e, male, same;/, female fifth leg; 
g, male fifth (V) and sixth (VI) legs. Dorsal view: h, left caudal ramus; i, 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 siu-face, 2 additional plumose setae from outer posterior 
surface; sixth plumose seta present on inner posterior surface. 

Male cephalothorax and free fourth pedigerous segment (fig. 376) 
similar to female. Genital segment (fig. 37^) suborbicidar, lateral 
posterior siu-face indented, indentations bearing fifth legs. Fifth 
leg (fig. 37 g) similar to that of female; sixth leg (fig. 37 g) 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 siu-face 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. 386) 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. 386) 4-parted, fourth part flattened, 
with 12 denticulations along inner margin. Female and male post- 
antennal process (fig. 386) consisting of 3 nodules, each with several 
fine, hairlike processes. Female and male postoral process (fig. 386) 
consisting of sub triangular, spinelike 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. izi 




0.2mm.2(cr) 6 



J 0.2 r 



mnn(?)d' 



Figure 38. — Caligus coryphaenae Steenstrup and Liitken, 1861, ventral view: a, 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;/, ster- 
nal furca, maxilliped bases (mxpd), and interpodal plate of first thoracic leg (ip). 



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. 38e) 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 simUar to that of female although first seg- 
ment with small, ledge-shaped projection on proximal inner surface. 
Female and male sternal furca (fig. 38/) 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 



T,ef 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss,s,p 

p 




rh 
c 


3dniH,p' 
3P 










II 


Outer 
Inner 


m 


p',p 


ni,p 
m,p 


m,in,mH 
c,P 


mH 
c,P 


2H,Q 
c,5P 


C 
P 


C 
C.2P 


C,3P 
c,3P 


III 


Outer 
Inner 


m 


d.m.p 
d,P,s,m,s 




mil 


p'*,c,p' 
c,P 


C,3p',p 
3P 


c 
P 


C,2P 
4P 




IV 


Outer 




P 




fra.dHf 


fm,dHt 


fm,dH,fm, 
dn,fm,dHf 









•Element in middle of segment. 

fFemale 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. Ill 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 




Q2'mm'?d' 



Figure 39. — Caligus coryphaenae Steenstrup and Liitken, 1861, right thoracic legs, anterior 
view: a, first; b, distal region of second segnaent of exopodite of first; c, second; d, third; 
e, exopodite of third (posterior view);/, 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 m 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 Caranx pelagica, Seriola dorsalis, Rachycentron 
canadus, and Caranx hippos as reported hosts. 

Caligiis quadratus Shiino 

Figures 40-42 

Caligus productus. — Rathbun, 1884, p. 487. — Wilson, 1905b, p. 597, pi. 14, figs. 

162-170. 
Caligus coryphaenae. — Yamaguti, 1936a, p. 5, pi. 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, pi. 75, fig. 4. 

Distribution and hosts. — 8 host records: 

locality host reference 

Japan Neothynnus macropterus 

Histiophorus orientalis 

Coryphaena hippurus 

Neothunnus albacora 

Rhinobatus schlegelii Shiino, 1954c, 1959b, 1960a 

"Pacific" Coryphaena hippurus Yamaguti, 1936a 

Atlantic Coryphaena hippurus Wilson, 1905b 

Western North At- Coryphaena species Rathbun, 1884 

lantic 

Material.— 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 mUes south of Niihau, 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. 



mean 


range 




5.02 


3. 90-6. 15 


3. 50, 3. 25, 2. 85 


1. 79 


1. 28-2. 08 


1. 93, 1. 95, 1. 53 


1. 65 


1. 30-1. 88 


1. 65, 1. 58, 1. 33 


1. 55 


1. 33-1. 80 


0. 73, 0. 60, 0. 55 


1. 24 


1. 08-1. 43 


0. 58, 0. 55, 0. 43 


1. 64 


1. 20-2. 18 


0. 73, 0. 60, 0. 53 


0. 15 


0. 10-0. 18 


0. 18, 0. 19, 0. 13 


2.33 


1. 25-3. 05 





110 PROCEEDINGS OF THE NATIONAL MUSEUM v( 

Measurements. — (In mm) 11 females and 3 males: 

female male 

mea'i 
Total length, excluding caudal setae 
Length of cephalothorax, including frontal 

region 
Width of cephalothorax 
Length of genital segment 
Width of genital segment 
Length of abdomen 
Length of caudal rami 
Length of egg strings (4 strings) 

Description. — 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 j miction of frontal region and rest of cepha- 
lothorax. Lateral margin of cephalothorax slightly uTegular, 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) UTegu- 
larly V-shaped, with fine membrane along outer surface. Posterior 
margin of median cephalothoracic area irregularly convex, area 
extendmg 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 thkd 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. 40c^) variable in 
shape, subovoid to rectangular, anterior lateral and lateral posterior 
surfaces frequently slightly lobate. Fifth leg (fig. 40/) situated on 
lateral ventral surface in posterior fourth of segment, consisting of 
2 nodules, outermost with single lightly plumose setule, innermost 
with 2 lightlj^ 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- 



COPEPOD CRUSTACEANS — LEWIS 



111 




Figure 40. — Caligus quadratus Shiino, 1954, dorsal view: a, female; b, male;c, posterior 
cephalothoracic sinus. Ventral view: d, female genital segment; e, male, same; /, female 
fifth leg; g, male, same; h, male sixth leg; i, caudal ramus. 



112 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. 40i) 
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. 406) 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 foiu-th leg attachment surface. Genital segment (fig. 40e) 
barrel shaped, constricted slightly anteriorly. Fifth legs (fig. 40^) 
situated in posterior half of ventral lateral surface, consisting of 3 
lightly plumose setules. Sixth legs (fig. 40A) situated on posterior- 
lateral ventral siu*face, 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. 416) 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. 41c^) 3-segmented, situated posterior and slightly medial to anten- 
nule base. First segment stubby, irregular, base subtriangular, 
bearing small, heavUy 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 setide-like accessory process from 
distal lateral surface and small, lappet-like process from proximal 
posterior siuface. Male antenna (fig. 41e) 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. Tines of trifid terminal process short, sit- 
uated at distal end of process. 



COPEPOD CRUSTACEANS — LEWIS 



113 




0.1mm fd" 



Figure 41. — Caliguj quadratus Shiino, 1954, ventral view: a, frontal region, right side show- 
ing lunula 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;*?, male right antenna, post- 
antennal process and base of antennule;/, claw and accessory process of male left antenna; 
g, left maxilla; h, female left raaxilliped; i, male, same;/, sternal furca. 



22&-321— 67- 



114 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

Female and male mandible (fig. 41o?) 4-parted, distalmost part 
curved medially, with 12 denticulations along inner surface. Female 
postantennal process (fig. 41o?) 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. 41g) also consisting of spinelike projection and 3 
nodules although projection much longer and strongly curved. Fe- 
male and male postoral process (fig. 41c^) spinelike, elongate. Female 
and male maxillule (fig. 41d) nodular, with 3 setules distally. Female 
and male maxilla (fig. 41^) 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 IK 
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. 41A) 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. Alale 
maxilliped (fig. 4H) more strongly developed, proximal projection 
of first segment shorter, inner medial surface of segment with small 
nodes proximally, larger node distally (node spinehke on 1 maxilliped 
of 1 of the 3 specimens); second segment and processes as in female. 
Female and male sternal furca (fig. 4:lj) situated between and shghtly 
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. 
guadratus Shiino 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. guadratus and the Hawaiian specimens is not indicated. The 



COPEPOD CRUSTACEANS — LEWIS 



115 




0.1mm ?d; 



Figure 42. — Caligus quadratus Shiino, 1954, right thoracic legs, anterior view: a, first; 
b, distal end of second segment of exopodite of first; c, second; d, third; e, exopodite of 
third; /, fourth. 



116 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Table 15. — Armature of thoracic legs I-IV of the female and male of 
Caligus quadratus Shiino, 1964c 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss,p 
d,p 




rh 
c,c 


H,2dH,P 
3P 










II 


Outer 
Inner 


m 


s.P 


ni,p 
m,s 


m,fm,fmH 
c,P 


fmH 
c,P 


h,mH,Q 
c,5P 


D 
P 


D 
c,2P 


D,3P 
C,3P 


III 


Outer 
Inner 


m 


d,m,p 
2s,P,s,m,s 




H 


c,p' 
c,P 


c,3p',P 
c,3P 


c 
P 


c,2P 
c,4P 




IV 


Outer 




s,p 




s,fm,fmH 


4fni,4niH 











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 

Figures 43-45 

Caligxis productus Dana, 1853, p. 1354, pi. 94, fig. 4. — Steenstrup and Liitken, 
1861, p. 357, pi. 3, fig. 6.— Kr0yer, 1863, p. 138. pi. 3, fig. 4a-i.— Brian, 
1898, p. 10.— Basse tt-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, pi. 73, fig. 
3.— Kirtisinghe, 1964, p. 53, figs. 25-26. 

Caligus monacanthi Kr0yer, 1863, p. 59, pi. 3, fig. 2a-e. — Wilson 1905b, p. 
607; 1937, p. 424, figs. 6-19. 

Caligus lobatus Wilson, 1935a, p. I, pi. 1, figs. 1-10. 

Caligus katuwo Yamaguti, 1936a, p. 6, pi. 4, fig. 55; pi. 5, figs. 56-68. — Shiino, 
1954f, 1 fig.— Nunes-Ruivo, 1956, p. 11, pi. 1, fig. b; pi. 2, fig. b. 



Distribution and hosts. — 37+host records: 



locality 
Subtropical Atlantic 



hosts 
Thymnus pelamys [sic] 
Thynnus 

Coryphaena 
Monacanthus species 
Bonito 

Naucrates ductor 
Neothunnus albacora 



references 
Dana, 1853 

Steenstrup and Liitken, 
1861 

Kroyer, 1863 
Fowler, 1912 
WUson, 1935a 
Nunes-Ruivo, 1956 



COPEPOD CRUSTACEANS — LEWIS 



117 



locality 
Gulf of Mexico 



Eastern Pacific 



Japan 

"Pacific" 
Indian Ocean 



Mediterranean 



hosts 
Scomberomorus cavalla 
Sarda sarda 
Flops saurus 
Coryphaena hippurus 
Scomberomorus maculatus 
Pogonias cromis 
Bonito 

Auxis thazard 
Coryphaena hippurus 
N eothunnus albacora 
Seriola dorsalis 
Paralabrax clathratus 
P. maculatofasciatus 
Verrunculus polylepos 
Mackerel 

Katsuwonus vagans 
Sphyraena argentea 
Scomberomorrus sierra 
Coryphaena species 
Seabass 

Calamus brachysomus 
Lutianus species 
Centropomus species 
Katsuwonus pelamis 
Coryphaena hippurus 
Euthynnus pelamys 
Coryphaena hippurus 
Katsuwonnus pelamis 
Euthynnus affinis 
Chrysophrys aurata 



references 



Pearse, 1952a 
Causey, 1953b 
Causey, 1955 
Wilson, 1937 



Shiino, 1959b 



Causey, 1960 

Shiino, 1959b 
Yamaguti, 1936a 



Kirtisinghe, 1964 
Brian, 1898 



Material.— Twenty-one females and 15 males (USNM 112904) 
from the extenal surface of Katsuwonus 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 Auxis thazard 
(Lacepede) collected south of Nawiliwili, Kauai, Hawaii (USFWS, HMS 
cruise 39, station 37). One female and 1 male (USNM 112907) from 
the gill cavity of Euthynnus yaito (Jordan and Evermann) from an 
unknown locaHty (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 
locaHty (USFWS). Three females (retained by author) from the 
gill cavity of 3 specimens of K. pelamis (Linnaeus) captured southeast 



mean 


range 


mean 


range 


4.80 


4. 10-5. 45 


4.58 


4.05-5. 15 


2. 16 


1. 83-2. 33 


2.39 


2. 10-2. 78 


1.92 


1. 50-2. 10 


2.04 


1. 88-2. 28 


1.58 


1. 05-1. 93 


1.07 


0. 90-1. 20 


1.09 


0. 80-1. 28 


0.75 


0. 65-0. S3 


1. 14 


0. 96-1. 39 


0.87 


0. 47-1. 00 


0.24 


0. 19-0. 27 


0.26 


0. 22-0. 29 


2.86 


1. 50-4. 15 







118 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

of Kauai, Hawaii (USFWS, HIMS 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 
Total length, excluding caudal setae 
Length of cephalothorax including frontal 

region 
Width of cephalothorax 
Length of genital segment 
Width of genital segment 
Length of abdomen 
Length of caudal rami 
Length of egg strings (17 strings) 

Description. — 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. Aledial 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. 43/) 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- 



COPEPOD CRUSTACEANS — LEWIS 



119 




0.05 mm. ? 



0.05 mm. d 



005mm. cf 



Figure 43. — Caligus productus Dana, 1853, dorsal view: a, female; b, male; c, posterior 
cephalothoracic sinus. Ventral view: d, female genital segment; e, male, same. Legs: 
/, female fifth; g, male fifth; h, 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. 45A) 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. 43e) distinct from fourth 
pedigerous segment, barrel shaped, posterior surface slightly irregular, 
ventral surface with padlike swelling posteriorly. Fifth leg (fig. 43^) 
situated on lateral ventral surface in posterior half of segment, simi- 
lar to that of female ; sixth leg (fig. 43A) 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 IK 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. 
44(Z) 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 accessoiy process proximally. 

Female and male mandible (fig. 44c) 4-parted, distal part curved 



COPEPOD CRUSTACEANS — LEWIS 



121 




0.2mm. d* 



Figure 44.—~Caligus produdus Dana, 1853, ventral view: a, frontal region, right side show- 
ing lunule; h, 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; e, left 
maxilla;/, right maxiUiped; g, sternal furca. 



122 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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. 44(Z) much longer than that of female, recurved; nodules 
and hairlike processes as in female. Female postoral process (fig. 
44c) long, spinelike; male (fig. 44c?) 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 Ij^ times the length of outermost, with fine 
membrane along inner margin; outermost process with finely frilled 
membrane along both margins. 

Female and male maxUliped (fig. 44/) 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. 44^) situated on median longitudinal axis posterior to 
maxUliped 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 o 
Calig 


f thoracic legs I-IV of the female and male of 
us productus Dana, 1852 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




2s*,p 
d,p 




rh 
C,C 


n,2dH,dP' 
3s 










II 


Outer 
Inner 


m 


2s, P 


m,p 
m,s 


m,mH 
c,P 


mH 
c,P 


mH,2Q 
c,5P 


C 
P 


D 

C,2P 


3P 
c,3P 


III 


Outer 
Inner 


m 


d.m.p 
d,3s,P,s,m,s 




s,H 


c,p' 
c,P 


c,3p',P 
c,3P 


c 
P 


c,2P 
C,4P 




IV 


Outer 




4s, p 




4s,dmnt 


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 



COPEPOD CRUSTACEANS — LEWIS 



123 




02mm9cr 



Figure 45. — Caligus produdus Dana, 1853, right thoracic legs: a, first, anterior view; h, 
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; /, 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 Cali(}us productus 
(USNM 6109) for his 1884 publication, all of the specimens were 
fomid to belong to C. quadratus Shiino. 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, figiu-ed, 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, pi. 13, figs. 150-153, 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, 
pi. 5, figs. 69-71; pi. 6, figs. 72-85.— Bere, 1936, p. 582.— Causey, 1953a, 
p. 4; 1953b, p. 8.— Nunes-Ruivo, 1956, p. 6, pi. 2, fig. a.— Causey, 1960, 
p. 328.— Shiino, 1960b, p. 527, fig. 1; 1963a, p. 336.— Yamaguti, 1963, p. 50, 
pi. 56, fig. 3. 

ICaligiis kuroshio Shiino, 1959c, p. 51, figs. 1-2.— Pillai, 1963, p. 80, fig. 9.— 
Yamaguti, 1963, p. 55, pi. 71, fig. 3. 



COPEPOD CRUSTACEANS — LEWIS 



125 



Distribution and hosts. — 20 host records: 



locality 
Western North Atlantic 

Subtropical Atlantic 

Gulf of Mexico 



Eastern Pacific 



Japan 



Indian Ocean 
Mediterranean 



hosts 
Bonito 

Gymnosarda pelarnis 
Cyhiuin 

Katsxiwonus pelarnis 
Pomatomas saltatrix 
Scomberomorus maculaius 
Lutianus grisens 
Mugil cephalus 
Oligoplites saurus 
Scomberomorus cavalla 
Gymnosarda alleterato 
Sarda sarda 
Sarda chilensis 
ILutjanus species 
Katsuwonus pelarnis 
Euthynnus pelarnis 
Katsuwonus pelarnis 
Thunnus thynnus 
Euthynnus affinis 
Pelamys sarda 



references 
Wilson, 1905b 
Rathbun, 1905 
Brian, 1924 
Nunes-Ruivo, 1956 



Bere, 1936 
Causey, 1953a 
Causey, 1953b 

Causey, 1960 
Shiino, 1963a 
Yamaguti, 1936a 

Shiino, 1959c 
Pillai, 1963 
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 sm-face 
with small indentation. Posterior sinus (fig. 466) 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 



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. 46c?) 




Figure 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; e, 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- 



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 siurface slightly swollen, swelling terminating 
posteriorly in slight knob bearing plumose setule. Distal inner 
margin plumose, distal surface with 3 plumose setae originating ventral 




Figure 47. — Caligus bonilo Wilson, 1905, female, ventral view: a, right side of frontal 
region showing lunule and antennule base; b, 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; e, 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. ili 

surface with approximately 23 plumose setae and setules, anterior 
dorsal 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 sm-face. 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. 
47(/) 2-segmented, situated lateral and slightly posterior to postoral 
process. First segment approximately foiu'-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. 47c) 2-segmented, situated posterior and medial to 
maxilla base. Fu-st 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. 47/) situated on median 
longitudinal axis of body approximately halfway between maxil- 
liped bases and first thoracic legs. Tines chisel shaped, diverging 
slightly; furca attached to small, platelike projection. 

For nature of legs and armatiu-e, see figm'e 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 specunens. 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 



COPEPOD CRUSTACEANS — LEWIS 



129 




01 mm 



Figure 48. — Caligus bonito Wilson, 1905, right thoracic legs, anterior view: a, first; h. second 
segment of exopodite of first; c, second; d, third; e, exopodite of third (posterior view); 
/, fourth. 



220-S21— G7- 



130 



PROCEEDINGS OF THE NATIONAL MUSEUM 



Tab 


LE 17.— 


Armature of thoracic h 


gs I-IV 
1905b 


f the female 


of Caligus bouito Wilson, 




Surface 


Intcr- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




s,p 
P 




rh 
c 


3dH,P' 
3P 






II 


Outer 
Inner 


m 


s.P 


m,p 
m,s 


m.fm.mH 
c,P 


mH 
c,P 


fmH.mH, 

Q,P 

4P 


C 
P 


D 

C,2P 


D,4P 
C,2P 


III 


Outer 
Inner 


m 


d,m,p 

d,2s,P,s, 

m,s 




s,H 


c,s,p' 
c,P 


c,3p',P 
c,3P 


c 
P 


c 
c,6P 




IV 


Outer 




5s,p 




s.fm.mll 


fm,mH,3fm, 
3H 











genital segment), upon the change that occurs in the length of the 
abdomen in other species (e.g., Caligus conjphaenae) , and upon 
Wilson's statement (1905b, p. 589) that "the joints [arel 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. honito 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 setide 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 C. 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. honito. 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. 48e) while the plumose seta and 
naked setule (Ih in Shiino's table I) are on the second segment. 
Shiino also uses the smaller size of his specimens to dift'erentiate C. 
kuroshio from C. honito. 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. honito) fists 4.5-5.6 mm, 
Brian (1935, C. honito) fists 8mm, Brian (1924, C. honito) lists 4,5,7 
and 8 mm females, and a male at 5 mm, Shiino (1960b, C. honito) 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. — Callgus quadratus, C. productus, and C. 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 e\4dence 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 nmnber 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 plimiose setae normally present on the inner 
margin of the second segment of the exopodite of the first thoracic 
leg. Both Caligus qiiadratus 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 pliunosities 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. IB, D, E. 

Caligus asymmetricus Pillai. — Kabata, 1965b, pp. 109, 110. 

Distribution and host.^ — ^Trivandrimi, South India, Euthynnus 
affinis {in Pillai, 1963) ; Queensland, Austraha, Euthynnus alleteratus 
{in Kabata, 1956b). 

Remarks. — Dana (1853, p. 1353) described the caligid Caligus 
thymni "from the external surface of a Bonito {Thymnus pelamys) 
. . . ." Presiunably the species was named after the host although 
the spelling 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 Thynnus. 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 Euthynnus affinis, thereby 
becomes a junior homonym. 

After the pubhcation 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 sjoecies. 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. 

Material.— One adult female and 3 males (USNM 112912) from 
the external sm-face of Euthynnus yaito (Jordan and Evermann) 
captiu-ed off Moku Manu Island, near Oahu, Hawaii. One adult 
female (USNM 112913) from the gill cavity of Euthynnus yaito (Jor- 
dan and Evermann) captm-ed 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. SO, 2. 55 

Length of cephalothorax, including frontal region 1. 95, 2. 05 2. 03, 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 

Length of caudal rami 0. 11, 0. 11 0. 13, 0. 11, 0. 10 
Length of egg strings (neither female ovigerous) 

Description. — Female cephalothorax (fig. 49a) ovoid, consisting 
of cephalon and fii"st 4 thoracic segments. Frontal region approxi- 
mately one-thirteenth the greatest length of cephalothorax, Avith 
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, A\dth fine 
membrane projecting laterally and second membrane projecting med- 
ially on ventral surface. Posterior lateral surface with small but 



COPEPOD CRUSTACEANS — LEWIS 



133 




0.1 mm.?c? 



0.3 mm.? 



0.3 mm.o' 



Figure 49. — Caligus asymmelricus Kab.nta, 1965b, dorsal view: a, female; i, 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; i, caudal ramus. 



134 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

distinct concavity. Posterior sinus (fig. 49c) narrow, bearing fine 
membrane along outer siu-face. 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. 49(/) narrower anteriorly than posteriorly, lateral margins 
flatly convex, posterior flatly concave. Fifth legs (fig. 49/) situated 
on ventral-posterior lateral surface, consisting of pair of knobs, ante- 
riormost with single pkimose 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. 49i) 
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. 49b) 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. 49^) 
on sHght 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 Iji 
times the length of second, broad medially, narrower proximafly and 
distally, distal two-thirds of anterior and anterior ventral surface 
bearing approximately 26 lightly plumose setae and setules, including 
1 long, 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- 



COPEPOD CRUSTACEANS — LEWIS 



135 




Figure 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;/, 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 distaUy 
than proximally. Third segment fused ^\dth 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, clawhke 
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. 50c) 
2-segmented, situated lateral and slightly posterior to postoral process. 
Fu-st segment slightly shorter than second, more strongly developed; 
second segment elongate, tapered proximally, wdth fine membrane 
along inner medial surface and 2 saber-shaped terminal processes. 
Innermost terminal process slighty less then IK times the length of 
outermost, with fine membrane along inner margin; outermost process 
mth finely frilled membrane along both margins. 

Female and male maxilliped (fig. 50/) 2-segmented, 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, subcorneal projection pres- 
ent on inner medial surface. Second segment distinct from clawHke 
terminal process, with setule-like accessory process on distal inner 
surface. Female and male sternal furca (fig. 50g) situated on median 
longitudinal axis of body, posterior to maxilliped bases. Base of furca 

Figure 51. — Caligus asymmetricus Kabata, 1965b, right thoracic legs, anterior view: a, first; 
b, second segment of exopodite of first; c, second; d, third; e, exopodite of third (posterior 
view);/, fourth. 



COPEPOD CRUSTACEANS — LEWIS 




0.05 mm.^' 



0.2 nnnn.9cr 



138 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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-/ and table 18. 

Table 18. — Armature of thoracic legs I-IV of the female and male of Caligus asym- 
metricus Kabata, 1965b 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss,p 
P 




rh 
c,c 


3fmH,P' 
3P 










II 


Outer 
Inner 


m 


s,P 


m,p 
m,s 


m.uiH 
c,P 


niH 
c,P 


2inH,Q 
5P 


C 
P 


D,c 
C,2P 


c,3P 
3P 


III 


Outer 
Inner 


m 


d,m,P 

d,2s,P,s, 
m,s 




mH 


c,p' 
c,P 


c,3p',P 
c,3P 


c 
P 


c,2P 
4P 




IV 


Outer 




P 




s,fm,mH 


fiu.mH, 
fm.mH, 
fm.H 











Discussion. — 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 nmnber (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 Euthynnus affinis along with several specunens of C. kuroshio 
(=?C. bonito Wilson, 19056), while the Hawaiian material came 
from a specimen of Euthynnus yaito. 

Caligus pelamydis Kr0yer 

Figures 52, 53 

Caligus pelamydis Kr0yer, 1863, p. 124, pi. 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.— WUson, 1905b, p. 594, pi. 13, figs. 154-161; pi. 14, fig. 
161a.— Brian, 1906, p. 43.— Brady, 1910, p. 589, fig. 69.— Stebbing, 1910, 



COPEPOD CRUSTACEANS — LEWIS 



139 



p. 558.— Guiart, 1913, p. 7.— Scott and Scott, 1913, p. 57, pi. 7, tigs. 2,3; 
pi. 9, figs. 1-5; pi. 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, pi. 1, fig. a. — Barnard, 1955, p. 245. — 
Causey, 1960, p. 329.— Hewitt, 1963, p. 78, fig. 6. 

Caligus scomberi Bassett-Smith, 1896b, p. 11, pi. 3, fig. 2; 1899, p. 450. 

Caligus scombri Scott T., 1901, p. 148, pi. 5, fig. 9-10.— Scott A., 1906, p. 196, 
pi. 6.— Guiart, 1913, p. 6. 

Distribution and hosts. — -12 host records: 



locality 
North Atlantic 

East coast of North 

Atlantic 
South Atlantic 



Gulf of Mexico 



New Zealand 
Mediterranean 



hosts 
Pelamys sarda 
Scomber scombrus 
Unknown 

Unknown 

Sarda sarda 
Pogonias cromis 
Scomberomorus cavalla 
Sarda sarda 
Thyrsites atun 
Scomber scomber 
Pelamys sarda 
Unknown 



references 
Kr0yer, 1863 
Scott and Scott, 1913 
Wilson, 1905b 

Brady, 1910 
Stebbing, 1910 
Nunes-Ruivo, 1956 

Causey, 1953a 
Causey, 1953b 
Hewitt, 1963 

Richiardi, 1880 
Guiart, 1913 



Material.— 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 1- 10 

Length of abdomen 1- 45 

Length of caudal rami 0. 17 
Length of egg strings (female nonovigerous) 

Description of female. — Cephalothorax (fig. 52a) ovoid, con- 
sisting of cephalon and first 4 thoracic segments. Frontal region 
narrow, middle of anterior sm-face distinctly indented, margin with 
fine membrane. Lunules (fig. 52/) large but filmy, extending 
posteriorly, on ventral surface, past junction of frontal region and 
remaining cephalothorax. Lateral margin of cephalothorax slightly 
kregular, 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 vnth 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. 52e) approxi- 
mately iy2 times as long as mde, distal half of inner surface plumose; 
three plumose setae present on distal surface, single plumose seta on 
outer distal siu"face and 1 plumose setule on either side of setae. 

Antennule (fig. 52g) 2-segmented, attached to lateral-anterior 
ventral surface of cephalothorax. Firet segment slightly less than 
tAnce the length of second, broader proximaily than distall}^; ventrally 
curved distal half of anterior surface mth approximately 18 plumose 
setules. Second segment rodlike, distal end M'ith approximately 12 
naked setules. Antenna (fig. 52h) 3-segmented, attached medial and 
posterior to antennule base. First segment squat, UTegular, heavily 
sclerotized; second segment tapered slightly from broad proximal to 
slightly narrower distal end, proximal outer surface -with small, sub- 
triangular projection. Thhd segment and clawlike terminal process 
indistinctly separable, segment elongate, wdth small, lappet-like pro- 
jection from posterior medial surface. 

Mandible (fig. 52/;,) 4-parted, rodlike, distalmost part ciu-ved 
inward, with 12 denticulations on inner surface. Postantennal process 

Figure 52. — Caligus pelaviydis Kr0yer, 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;/, 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; i, left maxilla;;', left maxilliped; k, sternal furca. 



COPEPOD CRUSTACEANS — LEWIS 



141 




142 



PROCEEDINGS OF THE NATIONAL MUSEUM 



(fig, b2h) situated lateral and slightly posterior to antenna base, 
consisting of short, heavUy sclerotized, dagger-shaped projection and 
3 nodules, 2 at base of projection, third slightly posterior, each with 
several hairlike processes. Postoral process (fig. 52A,) large, heavily 
sclerotized, spinelike. Maxillule (fig. 52/^) nodular, situated adjacent 
to inner proximal surface of postoral process, with 3 setules distally. 
Maxilla (fig. 32 i) 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 siu"face, 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. 

MaxHliped (fig. b2j) 2-segmented, situated medial and posterior 
to maxilla base. First segment strongly developed, with narrow, 
heavily sclerotized, strongly curved proximal articulation and muscle 
attachment sm'face. Second segment indistinctly separable from 
clawlike terminal process, with single, small, setule-like accessory 
process from distal inner sm^face. Sternal furca (fig. b2k) situated 
between and slightly posterior to maxilliped bases, tines broad, 
parallel, terminating in rounded surface. Furca 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 

Kr^yer, 1863 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




sss.p 

p 




h 
c 


4H 
3P 




s 
s 






n 


Outer 
Inner 


m 


S,P 


m 
m,s 


m,fm,H 
c,P 


II 
C,P 


H.mH.Q 
5P 


P 


C 
c,2P 


C,3P 
3P 


III 


Outer 
Inner 


m 


m,d,p 
2s,P,s,m,s 




H 


c,p' 
P 


c,3p',P 
3P 




p 


C,2P 
4P 




IV* 


Outer 




2s,p 




s,fm,pll 


fm,2pH,p', 
3pH,fm 











♦Segmentation of this leg in doubt: protopodite and first segment of exopodite recognizable, others ques- 
tionable. 



Figure S3. — Caligus pelamydis Kr0yer, 1863, female, right thoracic legs, anterior view: 
a, first; b, distal end of second segmentof exopodite of first; f, second; d, third; e, exopodite 
of third;/, fourth; g, exopodite of fourth. 



COPEPOD CRUSTACEANS — LEWIS 



143 




0,1 mm. 



0.05 mm. 



144 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Discussion. — Wilson (1905b) suggests that Caligus pelamydis 
Kj0yer, and C. scomberi 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 espece [C. pelamy- 
dis] est beaucoup plus volumineuse que la precedente [C. scombri, 
T. Scott's (1901) modification of Bassett-Smith's C. scomberi], le 
segTiient genital beancoup (sic) plus long et Fabdomen nettement 
forme de deux segments. Beaucoup d'autres caracteres s'opposent 
du reste a la fusion de cette espece avec la precedente, conmae 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 Kr0yer 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 figiu-es 52a,c, while 
in others the lobes are reduced or absent. 

The fourth thoracic leg is complex, the armatiu-e elements are 
poorly defined, and, in most cases, lightly sclerotized. The figure 
given by Wilson (1905b, pi. 13: fig. 161) does not show the frilled 
membranes adjacent to the bases of at least the fu'st 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 
ui 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-Sinith 

Figures 54-56 

Caligus longipedis Bassett-Smith, 1898b, p. 359, pi. 10, figs. 2-3. — Yamaguti, 

1963, p. 56. 
Caligus longipes Bassett-Smith, 1899, p. 452. — Wilson, 1905b, p. 556 (in key). 



I 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 145 

Caligus ampUfurcus 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 hosts. — 4 host records: 

locality hosts references 

Gulf of Aden Caranx melamphigus Bassett-Smith, 1898b 

Australia Cantherhines axjraud Heegaard, 1962 

Eastern Pacific Caranx lugubrius Shiino, 1959 

Gulf of Mexico Caranx crysos Pearse, 1952b 

Material.— 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 




1. 12 


0. 94-1. 35 


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 







Description. — Female cephalothorax (fig. 546) 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 kregular, 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 

226-321—67 10 



146 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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, 




1 — I — I — 1 
0.2mm.cr 



Figure 54. — Caligus longipedis Bassett-Smith, 1898, dorsal view: a, male; b, female; c; 
posterior cephalothoracic sinus. Ventral view: d, female genital segment; e, male, same, 
/, female fifth leg; g, male fifth and sixth legs; h, female caudal ramus; i, 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. 54c?) 
obcordate. Fifth legs (fig. 54/) situated on posterior lateral surface, 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 147 

consisting of 2 nodules, anieriormost 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. 54A) 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. 54^) consisting of nodule 
bearing 3 plumose setules; sixth legs (fig. 54^) 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. 541) 
larger than those of female, inner surface convex, plumose in distal 
half, remaining armature as in female. 

Female and male antennule (fig. 556) 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. 55c) 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 swelhng 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 sm-face. 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 nodides, 2 on base 
of projection, thh*d 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 
setides distally. Female and male maxilla (figs. 55^, /) 2-segmented, 
situated slightly lateral and posterior to postoral process. Second 
segment approximately IK 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 maxUliped (fig. 55^) 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 natm'e, tapered proximally to narrow proximal end. Second 
segment slender, approximately fom'-tenths the length of first, dis- 
tinct from short, clawlike terminal process, bearing setule-like acces- 
sory process from distal inner sm-face (not figiu-ed) and small, hahlike 
process from nodule on distal posterior sui'face. 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 fm'ca (fig. 55i) situated between and posterior to maxilliped bases. 
Fmxal 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 
natiu-e of thoracic legs, see figiu-e 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 
subtermmal 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 amplijurcus Pearse (USNM 93710) 



COPEPOD CRUSTACEANS — LEWIS 



149 




0.2mm.cr 



Figure 55. — Caligus longipedis Bassett-Smith, 1898, ventral view: a, frontal region, left 
side showing lunule and antennule base; b, 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; e, left maxilla; f, distal half of second segment 
of left maxilla; g, female right maxilliped; h, male, same; i, sternal furca. 



150 



PROCEEDINGS OF THE NATIONAL MUSEUM 





Table 20.— 


Armature 
Caligus 


of thoracic legs I-IV of the female and male of 
longipedis Bassett-Smith, 1898b 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




s,p 
P 




rh 
c,c 


3H,P' 
3P 










II 


Outer 
Inner 


m 


S,P 


m,p* 
m,s 


m,fm,mH 
c,P 


mH 
c,P 


mh.mH, 
5P^ 


c 
P 




c,2P 


C,2P 
4P 


III 


Outer 
Inner 


m 


2s,m,p 
2s,P,s,m,s 




m,s,H 


c,2s,p' 
P 


c,3p' 
4P 


c 

p 


c 
6P 




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 coidd 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 Pearee'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 maxUla (Heegaard's 
"first maxilliped," no figure number given but identified as MxPi 
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. amplijurcus 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 



COPEPOD CRUSTACEANS — LEWIS 



151 




Figure SS.—Caligus longipedis Bassett-Smith, 1898, right thoracic legs, anterior view: a, 
first; h, distal region of second segment of exopodite of first; c, second; d, third; <•, exopodite 
of third (posterior view);/, 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. 56e) 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 Cejdon. 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 oft' the sur- 
face of the host when it starts to dry. 

C. rugosus Shiino, 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 

Material.— 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 giU cavity of an unknown chaetodontid 
captured in the Oahu area, Hawaii. One female (paratype, USNM 
112922) from the gill cavity of Chaetodon jremhlii 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., captiu-ed 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., captiu-ed 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; e, male, same;/, male fifth and sixth legs; g, female 
fifth leg; h, caudal ramus 



COPEPOD CRUSTACEANS — LEWIS 



153 




154 



PROCEEDINGS OF THE NATIONAL MUSEUM 



USNM 112926) from the gill cavity of several specimens of C.fremblii 
Bennett, captm-ed 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 giU cavity of C. 
jremhlii Bennett, speared by Carolyn Lewis off Rabbit Island, Oahu, 
Hawaii. Two females (1 damaged) (paratypes, retained by author) 
from the giU 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- 










eluding 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. 14-0. 15 


Length of caudal rami 




0. 06 


0. 05-0. 07 


0.05 


(aU) 



Length of egg strings (3 strings) 0. 74, 0. 56, 0. 72 

Description. — Female cephalothorax (fig. 576) 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 foiu'th 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. 57g) 



COPEPOD CRUSTACEANS — LEWIS 



155 




0.1 mm. 2 ^ 



Figure 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); c, male left antenna; d, right maxilla; e, left maxilliped; /, sternal furca and 
maxilliped bases (mxpd). 



156 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

situated on posterior lateral siu'face 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. blh) short, rounded distal half of inner margin 
plumose, distal sm-face with 4 plumose setae and 2 plumose setules, 
1 on either side of setae. 

Male cephalothorax (fig. 57a) similar to that of female. Free 
fom'th 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 siu-face. 
Fifth legs (fig. 57/) consisting of 2 plumose setules; sLxth legs (fig. 57/) 
arising from 1 of lobate projections on posterior ventral sm-face 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-segmented, 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 sm-face with approximately 21 lightly plumose 
setules. Second segment rodlike, with 12 naked setules distally. Fe- 
male antenna (fig. 586) 3-segmented, situated posterior and medial to 
antennule base. First segment socket-like, in'egular ; second segment 
broader proximally than distally. Thkd segment fused with clawlike 
terminal process, bearing setule-like accessory process proximally. 
Male antenna (fig. 58c) with similar fu-st 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 jimction of segment and terminal process. 

Female and male mandible (fig. 586) 4-parted, distalmost part flat- 
tened, with 12 denticulations along inner surface. Female and male 
postantennal process (fig. 586) 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. 586) elongate, spinelike. Female and male maxillule (fig. 586) 
nodular, with 3 setules distally. Female and male maxilla (fig. 58c?) 
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 



COPEPOD CRUSTACEANS — LEWIS 



157 




0.C5 mm. Q cr 



0:1 mm. 9 cT 



Figure 59. — Caligus kapuhili, new species, right thoracic legs. Posterior view: a, first. 
Anterior view: b, second; c, third; d, exopodite and endopodite of third; e, fourth. 



158 



PROCEEDINGS OF THE NATIONAL MUSEUM 



processes. Innermost terminal process approximately Iji times the 
length of outermost, both with fine membranes along margins. 

Female and male maxilliped (fig. 586) 2-segmented, situated pos- 
terior and slightly medial to base of maxilla. First segment strongly 
developed, with distally concave ledge on inner siu-face 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. 58/) 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, fiat. 

Spine at distal end of &st 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 
Caligus kapuhili, 


I-IV of the female and male of 
new species 




Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




P 
P 




h 
c 


h,mH,2H 
3P 






rh? 
rh? 




II 


Outer 
Inner 


m 


P,s 


m,p 
m,s 


m,H 
c,P 


H 
c,P 


2h,Q 
5P 


P 


C 
c,2P 


C,3P 
3P 


III 


Outer 
Inner 


m 


a,m,P 
3s,P,m,s 




2ss,H 


c,p' 
P 


c,3p' 
4P 


c 
P 


c,3P 
c,3P 




IV* 


Outer 




3s,p 




mH 


mh 


3mH,m 









'Division between second and third segments of exopodite indistinct, incomplete. 

Discussion. — Caligus kapuhili exhibits several chai'acteristics 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. kajnihili, 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 
fom"th thoracic leg of C. laticaudus has a friUed membrane at the 
base of each spine; with the exception of the innermost terminal 
spine, these membranes are absent in C. kapuhili. 

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 

Figures 60, 61 
Caligus kala Lewis, 1964a, p. 142, figs. 2-3 

Distribution and hosts. — Hawaiian Islands, Naso hexacanthus. 

Material.— 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 fish trap 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 WiUiam Gosline. One 
female (USNM 112931) from the gill cavity of D. albisella Gill, from 
the Honokdu 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 bj' 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 




L27 


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 str 


ings) 


1. 30, 1. 48 









160 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 521 

Description. — Female 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 sm"f ace, 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 fom'th pedigerous segment. For 
description of rest of cephalothorax and fourth pedigerous segment, 
see original description. 

Female genital segment (fig. 60c?) of general obcordate shape al- 
though anterior end broadly rounded. Posterior lobes projecting to 
middle of abdomen, tipped by fifth legs. Fifth legs (fig. 60/) 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. QOh) as origionally described 
except for additional small setule present on outer distal ventral 
surface and hairlike process from middorsal surface. 

Male genital segment (fig. QOe) of varible shape, variation appear- 
ing due to longitudinal contraction of free body segments. Dis- 
tinct separation of fom'th pedigerous and genital segments in holotype 
specimen, separation not visible in presently figm'ed specimen but 
present in specimens exhibiting contracted condition. "Posterior 
margin" of original description (p. 144) thus formed by cuticular 
fold. Sixth leg (fig. QOg) 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. Qlb) 
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. 616) situated lateral to antenna base, 



COPEPOD CRUSTACEANS — LEWIS 



161 




0.1 mm.2cr 



Figure 60.—Caligus kala Lewis, 1964, dorsal view: a, female; b, male; c, posterior cepha- 
lothoracic sinus. Ventral view: d, female genital segment; e, male, same; /, 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 



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 




Figure 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; c, 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, 



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 tuberciilate 
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 sm-face 
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 
pubHcation, figures 3a-/ in Lewis, 1964a, 





Table 22.— 


-Armature of thoracic legs I-IV of the female and male of 
Caligus kala Lewis, 1964a 




Leg 


Surface 


Inter- 
podal 
plate 


Protopodite 


Exopodite 


Endopodite 




1 


2 


1 


2 


3 


1 


2 3 


I 


Outer 
Inner 




SSS.P 

d,p 




rh 
c,c 


3H,P 
3P 










II 


Outer 
Inner 


m 


s,P 


m,p 
m,s 


m,m,mH 
c,P 


mH 
c,P 


h.H.Q.P 
c,4P 


c,C 
P 


D 
P 


3P 
c,3P 


III 


Outer 
Inner 


m 


d,m,p 

d,2s,P,s, 
m,s 




H 


c,p' 
c,P 


c,3p',P 
c,3P 


c 
P 


c,3P 
C,3P 




IV 


Outer 




2s,p 




s,m,mH 


4m,4mH 











Caligus ligatus Lewis 

Figures 62-64 
Caligus ligatus Lewis, 1964a, p. 164, figs. 8-9. 

Distribution and hosts. — Hawaiian Islands, Acanthurus dussu- 
mieri, Naso hexacanthus. 

Material. — 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 



mean 


range 




3.29 


3. 20-3. 35 


2. 25, 2. 55, 2. 65 


1.62 


1. 55-1. 68 


1. 40, 1. 58, 1. 60 


1.49 


1. 43-1. 53 


1. 18, 1. 30, 1. 33 


0.95 


0. 93-0. 98 


0. 43, 0. 50, 0. 53 


0.93 


0. 88-1. 03 


0. 38, 0. 40, 0. 43 


0.59 


0. 58-0. 61 


0. 29, 0. 43, 0. 34 


0. 13 


(all) 


0. 12, 0. 15, 0. 14 


1. 55, 0. 73 







164 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Holocentrus xantherythrus 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). 

Measueements. — (In mm) 5 females and 3 males: 

female male 

mean 
Total length, excluding caudal setae 
Length of cephalothorax, including 

frontal region 
Width of cephalothorax 
Length of genital segment 
Width of genital segment 
Length of abdomen 
Length of caudal rami 
Length of egg strings (2 strings) 

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. 62c^) apple shaped, broader posteriorly, with fifth legs from 
ventral-posterior lateral surface. Fifth legs (fig. 62/) consisting of 2 
knobs, anteriormost with 1 plumose setule, posteriormost with 2, 

Female abdomen indistinctly separable from genital segment, con- 
sisting of 1 or 2 segments. 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. 62A) subrectangular, with smaU indenta- 



COPEPOD CRUSTACEANS — LEWIS 



165 



tion on medial outer surface, indentation bearing single plumose seta. 
Three plumose setae present on distal surface; single plumose setule 
on inner distal surface, second on outer surface, just medial to indenta- 




0.05 nnmoc 



0.05mm.9cr^ 



Figure 62. — Caligus ligatus Lewis, 1964, dorsal view: a, female; h, male; c, posterior ceph- 
alothoracic simus. Ventral view: d, female genital segment; e, male, same; /, female left 
fifth leg; g, male right fifth (V) and sixth (VI) 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. Q2e) subovoid, fifth legs (fig. 62^) 
arising from ventral lateral surface, consisting of 2 knobs, each with 
single, plumose setule; sixth legs (fig. 62^) arising from slight swell- 
ing on lateral-posterior ventral surface, consisting of minute nodiJe 
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. 636) 2-segmented, attached to 
anterior ventral surface of cephalothorax, not attached to adjacent 
frontal region. First segment slightly less than IK times the length 
of second, narrow proxunally and distally, broad medially; distal 
half of anterior and anterior ventral smiace with approximately 
23 plumose setae and setules. Second segment club shaped, with 
single naked setule from distal half of posterior sm-f ace, approximately 
11 naked setules from distal surface. Female antenna (fig. 63c) 
3-segmented, situated posterior and medial to antennide base. Fu'st 
segment dactyliform, ventrally facing sm'face 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 smiace. Male antenna (fig. 
Q3d) 3-segmented, first segment elongate, irregular; second segment 
broader proximally than distaUy, with adhesion surface on proximal 
half of anterior surface and small, shelf like adhesion surface project- 
ing from distal half. Third segment short, incompletely fused with 
short, bifurcate 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 sHghtly 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. Q3d) 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. 63c) nodular, with 2 small and 1 large setule. Female 
and male maxilla (fig. 636) 2-segmented, situated lateral and shghtly 
posterior to postoral process. First segment slightly more than 
foiu--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 



COPEPOD CRUSTACEANS — LEWIS 



167 




0.1nnm.?d' 



Figure 63. — Caligus ligatus Lewis, 1964, ventral view: a, frontal region, left side showing 
lunule and antennule base; b, 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; e, left maxilla;/, 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 v<u.. ili 




0.1mm ?^ 



0.1mm. ?d| 



Figure 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; e, exopodite of third; 
/, fourth. 



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. 63^) 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-hke accessory process present on distal inner 
surface of segment. Fu'st segment of female maxiUiped (fig. 63/) 
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. 

Remarks. — 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 



Leg 


Surface 


Inter- 
podal 
Plate 


Protopodite 


Exopodite 


Endopodite 


1 


2 


1 


2 


3 


1 


2 


3 


I 


Outer 
Inner 




ss.p 
P 




rh 
c 


3H,P 
3P 










II 


Outer 

Inner 


m 


S,P 


ni,p 
m,s 


m.fm.mH 
c,P 


dmH 
c.P 


h,mH,Q,P 
4P 


c 
P 


C* 
c,2P 


c,3P 
3P 


III 


Outer 

Inner 


m 


m,d,p 
s,P,s,m,s 




s,H 


c,s,p' 

C,P 


c,3p' 
c,4P 


c 
P 


c,3P 
C,3P 




IV 


Outer 




P 




s,fm,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. 

Material.— 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. 



rM 


VOL. 121 


female 


male 


2.37 


2. 29 


1.53 


1.67 


1.21 


1.26 


0.59 


0.38 


0.68 


0.38 


0.20 


0.20 




0.20 



170 PROCEEDINGS OF THE NATIONAL MUSEUM 

Measurements. — (In mm) 1 female and 1 male: 

Total length, excluding caudal setae 

Length of cephalothorax, including frontal region 

Width of cephalothorax 

Length of genital segment 

Width of genital segment 

Length of abdomen 

Length of caudal rami 

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-1, n-q; fig. 7. 

Distribution and hosts. — Hawaiian Islands, Acanthurus trio- 
stegus sandvicensis. 

Description. — See Lewis, 1964a. 

Caligus kaliimai Lewis 

Caligus kalumai Lewis, 1964a, p. 171, fig. 10. 
Distribution and hosts.— Hawaiian Islands, Acanthurus guttatus. 
Description. — See Lewis, 1964a. 

Order Lernaeopodoida 

Family Lernaeopodidae 

Charopinopsis Yamaguti, 1963 

Diagnosis. — 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 openmgs. Abdomen and caudal rami not visible. Anten- 
nule 3-segmented, subconical; antenna bu-amous, not chelate, ex- 
opodite broadly rounded, without exteiTially 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?). MaxiUae posterior 
to maxillipeds, joined at buUa, distal region convoluted, giving 
knobhke appearance. Maxillipeds 2-segmented, prehensile, second 
segment with short, clawlike terminal process. 

Male: See Yamaguti, 1963, p. 251. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 171 

Charopinopsis quaternia (Wilson) 

Figure 65 

Charopinus quaternius Wilson, 1935b, p. 343, pi. 4, figs. 42-49. — Causey, 1953a, 

p. 11, fig. 15. 
BrachieUa coryphaenae Pearse, 1952a, p. 35, figs. 129-135. — ^Pillai, 1962a, p. 85, 

fig. 18.— Yamaguti, 1963, p. 248, pi. 270, fig. 7. 
Charopinopsis quaternia (Wilson) Yamaguti, 1963, p. 251, pi. 271, fig. 3. 

Distribution and hosts. — 4 host records: 

locality hosts references 

Gulf of Mexico Peristedion gracilis 

Coryphaena hippurus Wilson, 1935b 

Scomheromorus cavalla Causey, 1953a 

Indian Ocean Coryphaena hippurxis Filial, 1962 

Material. — Nine females (USNM 112934) from the gill filaments 
of Coryphaena hippurus Linnaeus, captured 120 mUes south of Oahu, 
Hawaii (USFWS, HMS cruise 34). 

Measurements. — (In mm) 9 females: 

Total length, excluding posterior processes 

Length of cephalothorax 

Width of anterior region of cephalothorax 

Width of posterior region of cephalothorax (at level of 

maxillae) 
Length of "neck" between maxillae and trunk 
Length of trunk, excluding posterior processes 
Width of trunk 

Length of lateral-dorsal posterior processes 
Length of median-ventral posterior processes 
Length of egg strings (8 strings) 

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. 
656), 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 outlme to surface. 



mean 


range 


6.75 


6. 23-7. 35 


1.54 


1. 37-1. 70 


0.78 


0. 74-0. 85 


0.54 


0. 47-0. 63 


0.95 


0. 81-1. 11 


4.38 


4. 13-4. 65 


L75 


1. 30-2. 00 


0.66 


0. 44-0. 85 


2. 91 


2. 48-3. 55 


7.54 


6-53-8. 18 



172 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Antennule (fig. 65c) 3-segmented, attached to concave frontal 
projection, immediately anterior and slightly lateral to labriim. 
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. Q5d) 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 
siu-face 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 
"segment" approximately two-thirds the length of second, concave 
distally, with minute, setule-like projection arising from concavity. 

Mandible (fig. Q5d) rodlike, distal end flattened, inner edge rounded, 
without apparent denticulations. Maxillule (fig. 65e) situated 
adjacent to posterior lateral edge of mouth cone base, consisting 
of stalk with palp on distal half of inner siu"f ace, 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. 65/) tapered to bulbous swelling formed by convolution 
of distal end of each maxUla; swellings enclosing tip of capstan- 
shaped bulla. 

Maxilliped (fig. 65^) 2-segmented, situated posterior and medial 
to maxillule base, arising from small, apron-shaped projection of 
ventral surface of cephalothorax. Fii'st segment well developed, 
approximately twice the length of second, medial inner surface with 
knob-shaped swelling bearing small, conical projection. Second 
segment Avith 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 specunens 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 f om'-fif ths 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- 



COPEPOD CRUSTACEANS — LEWIS 



173 




Figure 6S.—Cliaropinopsis 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 maxiUiped; e, left maxillule; /, 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, bvit 
the author attributes this to natural variation, PUlai (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 

Diagnosis. — 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. MaxLllule with 
palp; maxillae removed well behind maxillipeds, distaUy 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 

Figures 66, 67 

B. thynni Cuvier, 1829, p. 257, pi. 15, fig. 5.— Gu^rin-M^neville, 1829-1844, pi. 
9, fig. 6a-c.— Nordmann, 1832, p. 90.— Milne-Edwards, 1840, p. 512.— 
Steenstrup and Lutken, 1861, p. 420, pi. 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, pi. 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, pi. 9, fig. 1.— Scott 
and Scott, 1913, p. 204, pi. 64, figs. 4-6.— Wilson, 1915, p. 703, pi. 25, fig. C; 
pi. 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, 
1953b, p. 15. — Delamare-Deboutteville and Nunes-Ruivo, 1953, p. 217. — 
Shiino, 1956b, p. 283, 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.l.— 
Kirtisinghe, 1964, p. 119, figs. 171, 172. 

Thynnicola ziegleri Miculicich, 1904, p. 48, figs. 1-3. 



NO. 3574 


COPEPOD CRUSTACEANS — LE 


WIS 175 


Distribution 


AND HOSTS. — 28 host records : 




locality 


hosts 


references 


Unknown 


Scomber thynnus 


Nordmann, 1832 




"tuna" 


Milne-Edwards, 1840 


North Atlantic 


Thynnus thynmis 


Bassett-Smith, 1896a 




T. vulgaris 


Rathbun, 1905 




"tuna" 


Wilson, 1915 




Orcynnus thynnus 


Scott and Scott, 1913 


Equatorial Atlantic 


"Albacore" 
Sciaena aquila 






"Baracottaer" 


Steenstrup and Lutken, 
1861 


Baltic 


Thynnus vulgaris 


Van Beneden, 1870a 


Gulf of Mexico 


Pomatomus saltatrix 


Bere, 1936 




Scomberomorus cavalla 


Causey, 1953b 


Hawaii 


Thynnus species 
Thunnus albacares 
Parathunnus sibi 


Bonnet, 1948 




Thunnus obesus 


Shiino, 1963a 


Japan 


Acanthocybixim solandri 


Shiino, 1956b 


Indian Ocean 


Chirocentrus dorab 


Thompson and Scott, 
1903 




Thunnus {Germo) macroptcrus 


Kirtisinghe, 1935 




Parathunnus obesus 






Neothunnus albacora 


Shiino, 1958 




Acanthocybium solandri 


Shiino, 1960b 




Indocybium lineolatum 


Pillai, 1962a 




Neothynnus macroptcrus 


Kirtisinghe, 1964 


Mediterranean 


Thynnus vulgaris 


Carus, 1885 




Thynnus thynnus 


Brian, 1899a 




"tuna" 


Brian, 1901 




Thunnus thynnus 


Delamare-Deboutteville 
and Nunes-Ruivo, 
1953 



Material.— Tliree females and 2 males (USNM 112935) from the 
external surface of Acanthocybium solandri (Cuvier) caught in the 
Hawaiian region (USFWS, HMS cruise 38). 

Measurements. — (In mm) 3 females and 1 male : 

female 
Total length, excluding posterior processes 
Length of cephalo thorax, 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 6.23, 13.05, 15.90 



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 



176 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

male 

Total length 1. 85 

Length of prosome 0. 7G 

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. 666) 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. 66c) 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 IK times the length of second, 
rounded distally, with single, minute spinule from medial anterior 
surface and 2 hairlike processes distally. Antennae (fig. 66e) 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. 66/) rodlike, wavy, flattened distally, distal inner 
surface with 6 denticidations (including distal end). Maxillule (fig. 
QQg) 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. 



COPEPOD CRUSTACEANS — LEWIS 



177 




0.05 mm. 



0.0 5mm. 



0.2mm. 



Figure 66. — Brachiella ihynni 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-1), and maxilliped; e, left antenna;/, 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 
maxilliile (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. 666) subconical, with numer- 
ous superficial annulations, distal end concave on inner half and 
enveloping cylindrical bulla (not figured). 

Maxilhped (fig. 66A) 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. 

Description of male. — Cephalothorax or prosome (figs. 67cr,6) 
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 proximaUy, 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. 67(/) 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 siu-face (spines not noted by Shiino, 1956b). 
Endopodite 2-segmented, first segment approximately twice the length 
of second, second with sharp indentation on distal inner siu-face, 
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 



COPEPOD CRUSTACEANS — LEWIS 



179 



flat, with membranous lateral sm-face; labium rounded, with heavily 

sclerotized band distally, band bearing fine membrane projecting 

, medially and membrane-like projection directed distally, latter topped 




00 5 mm. 



0.1mm 



Figure 67. — Brachiella thynni Cuvler, 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);/, right maxilla; g, right maxilliped; h, projections 
on right maxilliped. 



by row of small, bifid pads, pads topped by finely frilled membrane. 
Mandible (fig. Q7e) 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 sm-face 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 bifiu-cate band of heavy 
sclerotization. First segment well developed, with heavily sclerotized 
ring at proximal end, duct from maxillary gland visible inside proximal 
inner sui'face of appendage. Second segment fused to clawlike 
terminal process. 

Maxilliped (fig. Q7g) 2-segmented, situated on posterior half of 
apron-like projection. First segment of maxilliped elongate, with 
3 adhesion surfaces on inner siu'face. Adhesion sm'faces formed by 
numerous minute, padlike or hook-shaped projections (fig. 67A). 
Proximal adhesion sm-face associated with heavily sclerotized ridge 
and pad, medial surface associated with swelling, swelling with 
spinule; distal adhesion sm'face associated with swelling on distal 
inner sm-face. Second segment elongate, with adhesion process 
on distal half of posterior siu*face; 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 
maxilliped, 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 

Figures 68, 69 

Material. — Two 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 

Measurements. — (In mm) 1 female and 1 male: 

Total length, excluding lateral 

posterior processes but female 

including median projection 9.08 
Length of ceiihalothorax, 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 l.GO 

Lengtli of median posterior process 1.11 

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. 68e) 3-segmented, situated on ventral anterior sm'- 
face of cephalothorax. First segment large, irregular, approximately 
1 Yi 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 setiiles. Antenna 
(fig. 68^) biramoiis, 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 siu-face, at origin of endop- 
odite. Exopodite appearing 1 -segmented, approximately three- 
fourths the length of protopodite. Distal end of exopodite curved 
inward slightly, outer margin broadly convex, siu-face 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. Q8h) appearing 3-parted, proximal part broad proxi- 
mally, tapered abruptly, remaining parts tapered gradually to bluntly 
rounded distal end. Inner siu-face 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. 68i) situated just posterior to mandible base, 
consisting of stalk bearing palp on median posterior surface, palp with 
2 spinides distally; distal surface of maxillide with 2 nodides and spi- 
nide, both nodules with spinide distally. Maxillae (fig. 68a) elongate, 
approximately lYi times the length of cephalothorax, situated at 
posterior end of cephalothorax, well behind maxillipeds. Maxillae 
separate to distal end, attached to small, top-shaped bidla. 

Maxilliped (fig. Q8j) 2-segmented, extending anteriorly from pad- 
like ledge slightly posterior to mouth cone. First segment strongly 
developed, with smoothly irregidar outline, bearing small, spinule- 
like projection from medial inner siu-face. 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 siu'face, bearing 2 small, spinelike accessory 
processes from inner siu-face and single, large, clawlike terminal 
process. 

Description of male. — Body (figs. 69a, b) tapered at both ends, 
lateral margins parallel throughout most of length although constric- 

FiGURE 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;/, 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]; i, right maxillule, 
ventral view;/, right maxilliped, ventral view. 



COPEPOD CRUSTACEANS — LEWIS 



183 




Q025mm.? 




184 



PROCEEDINGS OF THE NATIONAL MUSEUM 



tion present immediately posterior to maxillipeds in 1 specimen (not 
shown in figures). Posterior region with pah- 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 IK times the combined lengths of second and third segments, 




0.1 mm.o" 



Figure 69. — Brachiella regia, new species, male: a, lateral view; i, dorsal view. Lateral 
view: c, right antennule; d, right antenna; e, denticulated portion of mandible [drawn 
by Z. Kabata];/, right maxillule; g, right maxilla; h, 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. 69c?) 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. 69e) appearing 3-parted, rodlike; distalmost part 
flattened, inner surface with 9 denticidations, the third slightly smaller 
and considered a secondary denticulation by Kabata (from correspond- 
ence) . Maxillule (fig. 69/) ad j acent 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. MaxUla 
(fig. 69^) 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. Q9h) 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 
smaU, 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 pau's of posterior processes, an unpau'ed 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 maxUlae. 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, whUe 
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 

Naohranchia Hesse, 1863 

Diagnosis. — 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 rodhke. 
Maxillule bipartite, with or without palp; maxilla fofiaceous, usually 
with muscle bands extending length of appendage, forming organ 
of attachment that envelops gill filaments of host. Alaxilliped 
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. 

Naohranchia species 

Figure 70 

Material.— One female (USNM 112937) from gill cavity of 
spotted moray eel from Honolulu Aquarium. 



NO. 3574 COPEPOD CRUSTACEANS — LEWIS 187 

Measurements. — (In mm) 1 female: 

female 

Total length, including egg naasses 3. 75 

Length of cephalo thorax, 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 romided trmik. 
Anterior end of cephalothorax slightly expanded, from dorsal view- 
point; tapered to narrow, rounded end, in lateral view, ventral sm^- 
face bearing antennules, antennae, mandibles, maxillules, and 
maxillipeds. Margin of anterior end of cephalothorax with heavily 
sclerotized band broken by several mmute swellings associated with 
minute channels through band, similar to nodules bearing hau'like 
processes on caligoids. Remaining part of cephalothorax iiTegular 
(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 sm'faces 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 imless 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. lOd) 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 imiramous; 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 



inner margin.^: Maxillule (fig. 70e) nodular, situated adjacent to pos- 
terior lateral surface of labium base, with 2 setule-like projections 
distally. Maxillae (fig. 70/) fused, foliaceous, situated at posterior 




0.5 mm 



(^ 004 mm. 



Figure 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; e, oral region showing antenna (a-2), 
mouth cone, mandibles, and maxillule (ma-1);/, maxilla; g, right maxilliped. 

end of vermiform cephalothorax and included in anterior portion of 
trimk. 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. 70^) 2-segmented, situated in small concavity 
slightly posterior and lateral to labium base. First segment with 
shelflike indentation on distal half of inner siu"face, 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 siuface 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 TV. aulojn Yamaguti (1939c), although in this species they 
are oriented in a longitudinal dhection whUe 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 immatiu-e condition as Pearse (1952a, fig. 126) indicates for N. 
spinosa Pearse. The possible immature condition of the specimen 
may also explain the distinctive cuplike structiu"es. Yamaguti (1939c) 
figures a distinct cuplike structm-e for a 3.2 mm specimen of N. aulopi 
while Shiino (1958) states "striated pads just behind 2nd max- 
illipeds [= maxiUipeds] 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 cm^ved, 
heavily sclerotized rodlike structures are visible in the cuticle. 

The Hawaiian specimen appears to most closely resemble A^. varia- 
bilis Brian. The similarity is not only in the general shape of the 
body in the 3-segmented antennule although Pfllai (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 maxfllides and maxiUipeds also are similar to those described 
for A^. 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 
it in Hawaiian waters. The list includes the copepods described 
in the present paper and in Lewis, 1964a,b, 1966. With few excep- 
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 cuvicr (tiger shark) 

Pterolamiops longimanus (whitetip shark) 
Carcharhinus melanopterusl (blacktip shark) 
Sphyrna lewini (hammerhead shark) 



Hexanchus griseus? 
Unidentified sharks 



7Aetobaius narinari (eagle ray) 
Saurida gracilis (lizard fish) 
Muraenidae (moray eels) 

Fistularia petimha (cornet fish) 
Aulostomus chinensis (trumpet fish) 



Holocentrus xantherythrus 
Myripristis pralinius 
Lampris regixis 
Sphyraena barracuda 
Pranesus insularuml 
Seriola dumerilii (yellowtail) 
Caranx melampygusi 

Coryphaena hippurus (dolphin) 



190 



copepod parasites 
Anthosoma crassum 
Dinematura latifolia 
Pandarus satyrus 
Phyllolhyreus cornutus 
Nesippus crypturus 
Alehion echinatus 
Pandarus cranchii 
Pandarus smithii 
Kroyeria praelongacicula 
Pandarus cranchii 
Alehion echinatus 
Paeon vaissierei 
Demoleus heptapus 
Alehion gracilis 
Dineynatura latifolia 
Pandarus cranchii 
Pandarus smithii 
Trehius caudatus 
Caligus kola 

Pseudotaeniacanthus puhi 
Naohranchia species 
Dentigryps bifurcatus 
Nesippus costatus? 
Dentigryps bifurcatus 
Caligus ligatus 
Caligus flexispina 
Caligus ligatus 
Nesippus costatus? 
Brachiella regia 
Midias lohodes 
Caligus ligatus 
Nesippus costatus? 
Dentigryps ulua 
Caligus longipedis 
Euryphorus nordmanni 
Caligus coryphaenae 
Caligus quadralus 
Charopinopsis quaternia 



COPEPOD CRUSTACEANS — LEWIS 



191 



host 
Mulloidichthys aurifiarnma 
Parupeneus pleurostignia 
Chaetodon fremblii 

Chaetodon quadrimaculatus 
Chaetodon miliaris 

Dascyllus albisella 

Pomacentrus jenkinsi 
Bodianus bilunnlatus 
Unidentified labrid 
Scarus species 
Zanclus canescens 
Acanthurus guttatus 
Acanthurus nigroris 
Acanthurus olivaceus 



Acanthurus dussumieri 



Acanthurus xanthopterus 

Acanthurus mata 

Acanthurus triostegus sandvicensis 



Ctenochaetus strigosus 

Zebrasoma flavescens 
Naso lituratus 
Naso hexacanthus 



Naso unicornis 



Acanthocybium solandri (wahoo, ono) 

Auxis thazard (frigate mackerel) 
Katsuwonus pelamis (skipjack) 



copepod parasites 
Hatschekia breviramus 
Lepeophthelrus dissimulatus 
Dentigryps bifurcatus 
Caligus kapuhili 
Lepeophtheirus dissimulatus 
Nesippus costatus? 
Caligus kapuhili 
Caligus kala 
Caligus ligat^is 
Caligus kala 
Dentigryps bifurcatus 
Caligus flexispina 
Nesippus costatus"! 
Nesippus costatus"? 
Caligus kalumai 
Nesippus costatus"? 
Dentigryps bifurcatus 
Lepeophtheirus dissimulatus 
Peniculus calarmisl 
Caligus ligatus 
Lepeophtheirus dissimulatus 
Peniculus calamus? 
Nesippus costatus? 
Peniculus calamus? 
Nesippus costatus? 
Dentigryps bifurcatus 
Lepeophtheirus dissinmlatus 
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 sibi 

Thunnus alhacares (in Sliiino, 1963a) 
Thunnus ohesus (in Shiino, 1963a) 
Istiompax 7uarlinal (black marlin) 
Makaira ampla? (Pacific blue marlin) 
Makaira audax (striped marlin) 

Remoropsis brachypterus 
Rhombochirus osteochir 
Pervagor spilosoma 
Ostracion lentiginosus 
Diodon holocanthus 
Plankton 



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 histiophori? 
Gloiopotes huttoni 
Pennella species 
Lepeophtheirus crassus 
Peniculus calamus? 
Anchistrotos moa 
Nesippus costatusi 
Caligus coryphaenae (immature 
male) 



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KiRTISINGHE, P. 

1932. Pennella zeylanica n.sp., a parasitic copepod of Histiophorus gladius 
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1955. Parasitic copepods of Travancore-Cochin. Bull. Cent. Res. Inst. 

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1926. A list of parasitic Copepoda found at Plymouth. Parasitology, vol. 

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1933. A list of British fishes with their characteristic parasitic Copepoda. 

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1956. Caliqus oligoplilisi sp. n. copepodo parasito del "Zapatero" Oligo- 
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1948. A second report on parasitic copepods collected at Beaufort, N,C. 

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1953. A propos des pennelles (crustaces copepodes) parasites des germons, 
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1952. Copepods parasitic on Japanese fishes, 1: On the species of Caligus 

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Rep. Fac. Fish. Pref. Univ. Mie, vol. 1, no. 3, pp. 273-290, figs. 1-6. 
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the fish Myxocephalus raninus J. & S. Bull. Japanese Soc. Sci. 

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Rep. Fac. Fish. Pref. Univ. Mie, vol. 2, no. 2, pp. 269-311, figs. 

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1959d. Ostpazifische parasitierende Copepoden. Rep. Fac. Fish. Pref. Univ. 

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*1880. Prospetto della fauna del mare Adriatico. Boll. Soc. Adriatica Sci. 
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1915. North American parasitic copepods belonging to the Lernaeopodidae, 

with a revision of the entire family. Proc. U.S. Nat. Mus., vol. 47, 

no. 2063, pp. 565-729, pis. 25-56, figs. 1-15. 
1917. North American parasitic copepods belonging to the Lernaeidae, with 

a revision of the entire family. Proc. U.S. Nat. Mus., vol. 53, no. 

2194, pp. 1-150, pis. 1-21, figs. a-c. 
1919. A new species of parasitic copepod, with notes on species already de- 
scribed. Proc. U.S. Nat. Mus., vol. 55, no. 2274, pp. 313-316, 

pl. 21. 



204 PROCEEDINGS OF THE NATIONAL MUSEUM vol. m 

Wilson, C. B. 

1921. New species and a new genus of parasitic copepods. Proc. U.S. Nat. 

Mas., vol. 59, no. 2354, pp. 1-17, pis. 1-7. 

1922. North American parasitic copepods belonging to the family Dicheles- 

thiidae. Proc. U.S. Nat. Mus., vol. 60, no. 2400, pp. 1-100, pis. 
1-13. 

1923. Parasitic copepods in the collection of the Riksmuseum at Stockholm. 

Ark. Zool., vol. 15, no. 3, pp. 1-15, pis. 1-2. 

1924. Parasitic copepods from the Williams Galapagos Expedition. Zoolo- 

gica. New York Zool. Soc, vol. 5, no. 19, pp. 212-217, pi. 20. 

1932. The copepods of the Woods Hole region, Mas.sachusetts. U.S. Nat. 
Mus. Bull., vol. 158, xix + 635 pp., 41 pis., 316 figs. 

1935a. Pveports on the collections obtained by the first Johnson-Smithsonian 
Deep-Sea Expedition to the Puerto Rican Deep. Smithsonian 
Misc. Coll., vol. 91, no. 19, pp. 1-9, pis. 1-3. 

1935b. Parasitic copepods from the Dry Tortugas. Carnegie Inst. Wash- 
ington, publ. 452, Pap. Tortugas Lab., vol. 29, no. 12, pp. 327-347, 
pis. 1-6. 

1936. Note on a copepod from Cuba. Mem. Soc. Cubana Hist. Nat., vol. 

10, no. 1, p. 32. 

1937. Some parasitic copepods from Panama Bay. Journ. Washington 

Acad. Sci., vol. 27, no. 10, pp. 423-431, figs. 1-34. 
Yamaguti, S. 

1936a. Caligoida, I. Pt. 2 in Parasitic copepods from fishes of Japan, 22 pp. 

12 pis. [Publ. by author.] 
1936b. Caligoida, II. Pt. 3 in Parasitic copepods from fishes of Japan, 21 

pp., 9 pis. [Publ. by author.] 
1939a. Cyclopoida, II. Pt. 4 in Parasitic copepods from fishes of Japan, 

pp. 391-415, pis. 1-13. [Vol. Jub. pro Prof. Sadao Yoshida, vol. 2.] 
1939b. Caligoida, III. Pt. 5 in Parasitic copepods from fishes of Japan, 

pp. 44.3-487, pis. 14-33. [Vol. Jub. pro Prof. Sadao Yoshida, vol. 

2.] 
1939c. Lernaeopoida, I. Pt. in Parasitic copepods from fishes of Japan, 

pp. 529-578, pis. 34-58. [Vol. Jub. pro Prof. Sadao Yoshida, vol. 

2.] 

1953. Cyclopoida, III, and Caligoida, IV. Pt. 7 in Parasitic copepods 

from fishes of Japan. Publ. Seto Mar. Biol. Lab., vol. 3, no. 2, I 
pp. 221-231, pis. 1-5. 

1954. Parasitic copepods from fishes of Celebes and Borneo. Publ. Seto ^ 

Mar. Biol. Lab., vol. 3, no. 3, pp. 375-398, pis. 1-6. | 

1963. Parasitic Copepoda and Branchiura of fishes, vii + 1104 pp., 333 pis. j 

Yamaguti, S., and Yamasu, T. j 

1959. Parasitic copepods from fishes of Japan with descriptions of 26 new |l 

species and remarks on two known species. Biol. Journ. Okayama 

Univ., vol. 5, nos. 3, 4, pp. 89-165, pis. 1-14. 



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. Pettibone 

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 poljaioids 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 mdicates, it shows certain affinities to Admetella 
Mcintosh. 

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- 

1 



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; segments 
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 (i.e., lepidonotoid) ; Avith paired antennal scales. 
Tentacular segment (I) achaetous, with 2 pairs tentacular cirri, 
Avith 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 
distaUy, with faint spinous rows and tapered tips. 

Admetella longipedata (Mcintosh) 

Figures 1, 2 

Polynoe (Admetella) longipedata Mcintosh, 1885, p. 124, pi. 14, fig. 5; pi. 20, 
fig. 6; pi. 12A, fig. 17. 

Admetella longipedata Darboux, 1900, p. 103. — Augener, 1906, p. 123. — Ehlers, 
1908, p. 40, pi. 2, figs. 10, 11; pi. 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, 1S85.] 



NO. 3575 BATHYAL POLYNOIDS — PETTIBONE 3 

Admetella hastigerens Chamberlin, 1919, p. 64, pi. 9, figs. 6-8. — Treadwell, 1923, 

p. 3. 
Admetella dolichopus Chamberlin, 1919, p. 67, pi. 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 Hawaii, Philippine Islands, and Lower California to off 
Central America. The specimens were identified by Treadwell as 
Polynoe mirahilis Mcintosh (TreadweU, 1906, 1920) and Admetella 
hastigerens Chamberlin (TreadweU, 1923) and by Chamberlin (1919) 
as 2 new species of Admetella, A. hastigerens, and A. dolichopus. AU 
of the above are herein referred to Admetella longipedata (Mcintosh) . 
Fauvel (1914, p. 39) questioned the identification by Treadwell (1906, 
p. 1149) of Polynoe mirahilis from off Hawaii. Uschakov (1950, p. 
157) also questioned the identifications in both records of TreadweU 
(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- 
899 fms, coral, sand, foraminifera, rock (USNM 5458, identified by 
Treadwell as Polynoe mirahilis, 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, 
PhUippine Islands, 13°36' N, 120°45' E, 340 fms, fine sand (USNM 
17507, identified by Treadwell as Polynoe mirahilis, 5 spec); Sta. 
5122 (Feb. 2, 1908), east coast of Mindoro, PhUippine Islands, 13°2r 
N, 120°30' E, 220 fms, green mud (USNM 17606, identified by Tread- 
well as Polynoe mirahilis, 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 TreadweU 
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 spincUe shaped, mdest in middle, tapering anteriorly and 
posteriorly, arched dorsally, flattened ventraUy. Parapodia very long, 
as long as width of body, inflated and flattened transversely (fig. Ic). 
Elytra (usually missing) on large inflated elytrophores, 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 




Figure 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 (III); b, 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-III= 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, la, 6) 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 prostomiimi 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, la, 6) 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 bidbous facial tubercle between bases of palps, con- 
tinuous with a longitudinal ridge on upper lip. Second or buccal 
segment with first pair ely trophores ; parapodia rudimentary, achae- 
tous; with dorsal, low, slightly bilobed, transverse nuchal fold between 
ely trophores ; 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,b). 

Parapodia (figs. 2a,b) elongated, inflated, thin walled, flattened 
transversely, subbiramous. Notopodia small lobes on anterodorsal 
faces of elongated neuropodia, inflated basaUy 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 distaUy, postsetal lobes subtriangular, presetal lobes extending 
as long digitiform acicular processes. Acicida 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, ii'idescent, 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 pi. 
12A, fig. 17 in Mcintosh, 18S5; pi. 3, figs. 1-5 in Ehlers, 1908; pi. 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. lc,26) on ventral bases 
of parapodia, begin on segment 6 and continue posteriorly, inflated 
basally, tapered distally to short tubes directed between parapodia. 



i 



6 



PROCEEDINGS OF THE NATIONAL MUSEUM 



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. la,b). 
Pharynx reddish to blackish; dis tally with soft papillae and 2 pairs 
light-brown interlocking jaws. Color (according to Tread well, 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- 




FiGURE 2. — Admetella longipedata: a, parapodium of elytragerous segment, anterior view, 
elytron missing, position of acicula shown in dotted lines; h, parapodium of cirrigerous \ 
segment, posterior view, lateral part of dorsal tubercle indicated, style of dorsal cirrus 
missing; c, 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. | 

Bathyadmetella, new genus 

Type species: 5. commanc^o, new species. Gender: feminine. j 

Diagnosis: Polynoids mth body large, oval, flattened, segments j 

more than 50 (58). Prostomium ^^ith 2 long palps and 3 antennae; \ 



NO. 3575 BATHYAL POLYNOIDS — PETTIBONE 7 

median antenna with ceratophore inserted on anterior part of pro- 
stomiuni; lateral antennae inserted on anterolateral extensions of pro- 
stomiimi (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 elytrophores. 
With lamellar papillae on ventral hp 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 

Figures 3, 4 

Material examined: The species is represented by a single damaged 
specimen, collected in a trawl haul, southwest of the mouth of the 
Cohmibia River, Oregon, 45°45' N, 125°09' W, 900 fms, 2.6° C 
bottom temperature, 34.3727oo 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, 6) bilobed, wider than long, with paired very 
large bidging 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 
prostomimn; 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 



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. 




Figure 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 
(II). (anSh = antennal sheath, bC = buccal cirrus, elp = elytrophore, fT==facIal tubercle, 
lP = lamellar papilla, tSh=^ tentacular sheath.) 

Fhst or tentacular segment (figs. 3a-c) forms short ring dorsally, 
projecting laterally to prostomium, achaetous, with 2 pairs long 
slender tentacular chri similar to lateral antennae, cirrophores short, 
cylindrical, with a projectmg acicular process. Pahed long tentacular 



BATHYAL POLYNOIDS — PETTIBONE 



9 



sheaths attached basally on tentacular segment, wider basally, tapermg 
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. 36, d) with first pau- elytrophores; parapodia rudimentary, not 
elongated, with no to- and neuroacicular achaetous lobes; ventral or 
buccal cbri 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; 




'"'.0.3' 



'V,, 



Figure 4. — Bathyadmetella commando, new species: a, cirrigerous parapodium, posterior 
view, style of dorsal cirrus missing; h, 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 cu-ri on these 2 segments extra long, extending beyond the 
distal tips of the neuropodia (fig. 36). Rest of ventral cirri short, 
subulate, in middle of elongated neuropodia (figs. 4a, 6) . 

Parapodia (figs. 4a,6) elongated, subbu-amous. Notopodia small 
lobes on anterodorsal faces of neuropodia, inflated basaUy, 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 nem'oacicula. Neurosetae 
munerous, forming brushlike transparent bundles. Neurosetae with 
long, bare, basal regions, distaUy wider, flattened, with somewhat 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

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 this area) . Anal cirri not observed. Nephrid- 
ial papillae on ventral bases of parapodia, beginning on segment 4 
and continuing posteriorly, small at first, becoming longer (shorter 
than ventral cim), 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 bulgmg 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(?) renotv.hulata Moore, 1910. Gender: 
feminine. 

Diagnosis: Polynoids with relatively few segments (33). Pro- 
stomium with 2 long palps and 3 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 pau's tentacular cuTi, 
with bulbous facial tubercle between bases of palps. Buccal segment 
(II) with first pair elytra, setigerous parapodia, and long ventral 
buccal cu-ri. Elytra 14 pau's, 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 famt spinous rows and 
tapered tips. Nephi-idial papillae remarkably elongated, some ex- 
tending to tips of neuropodia. 

Bathymoorea renotubulata (^loore), new combination 

Figure 5 

Polynoeil) renoluhulata Moore, 1910, p. 368, pi. 31, figs. 59-64. 
Admetella renoluhulata Hartman, 1938, p. 123. 

Material examined: Holotype Polynoe{l) renotuhulata (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 H 

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 sm.all. 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 cu'rigerous segments inflated; cuto- 
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 pau-ed, thick, smooth, tapering. 

First or tentacular segment projecting lateral to prostomimn, 
achaetous, with 2 pau's tentacular cirri; styles long, slender, tapered. 
A bulbous facial tubercle between palps, continuous with longitudmal 
ridge on upper lip. Second or buccal segment with fu-st pair elytro- 
phores, notopodial acicular lobes, and neuropodial acicular lobes 
bearing bundles of neurosetae; cuTophores of ventral cutI large, 
stvles missing. Ventral cirri on remaining segments short, subulate 
(fig. 5c). 

Parapodia (figs. 5b, 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 subcorneal postsetal lobes; 
presetal lobes prolonged into dehcate acicular processes, vnth long, 
dense, brushlike bundles of neurosetae. Neurosetae transparent, dis- 
taUy expanded and flattened, ^vith faint spinous rows, abruptly 
tapering to entire tips (may be split, appearing bifid, fig. Be). 

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, Bathy admetella, and Bathymoorea, 
have a number of characters in common, e.g., prostomia with paired 



12 



PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 




Figure 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), h, elytragerous parapodium, anterior view, elytron missing, 
position of acicula shown in dotted lines; c, cirrigerous parapodium, posterior view, style 
of dorsal cirrus missing; d, tip of notoseta; e, tips of neurosetae. (1-11 = 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 elj'trophores) 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. renotuhulata (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. Nephridial papillae not especially 
elongated, shorter than ventral cirri 2 

2. Antennal scales short, triangular, attached between ceratophores 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 pan- elytrophores. Lower lip region bare, without 
papillae. Notopodia with long acicular processes and small bundles noto- 
setae Admetella Mcintosh 

A. longipedata (Mcintosh) 
Antennal sheaths long, digitiform, attached basally to and enclosing lateral 

antennae. Ceratophores of median antemaa 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 pis. 
Chamberlin, Ralph V. 

1919. The Annelida Polychaeta. Mem. Mus. Comp. Zool. Harvard, vol. 
48, pp. 1-514, 80 pis. 
Darboux, J. Gaston 

1900. Recherches sur les Aphroditiens. Bull. Sci. France et Belgique, 
vol. 33, pp. 1-274, 83 figs. 
Ditlevsen, Hjalmar 

1917. Annelides I. Pt. 4 oj vol. 4 in The Danish Ingolf expedition, 71 pp., 
6 pis. 
Ehlers, Ernst 

1908. Die bodensassigen Anneliden aus den Sammlungen der Deutschen 
Tiefsee-Expedition. In Wissenschaftliche Ergebnisse der Deut- 
schen Tiefsee-Expedition, 1898-1899, vol. 16, pp. 1-168, 23 pis. 
Eliason, Anders 

1951. Polychaeta. In Report of the Swedish Deep-Sea Expedition, 1947- 
1948, Zool., vol. 2, no. 11, pp. 131-148, 2 pis., 5 figs. 
Fauvel, Pierre 

1914. Annelides Polychetes non pelagiques provenant des campagnes de 
VHirondelle et de la Princesse- Alice (1885-1910). Res. Camp. 
Sci. Monaco, vol. 46, pp. 1-432, 31 pis. 
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 (Sfbo^a-Expedition, vol. 24b, pp. 1-140, 19 
pis., 5 figs. 
McIntosh, William C. 

1885. Annelida Polychaeta. In Challenger reports, vol. 12, pt. 3, pp. 1-554, 
94 pis. 
Moore, 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, pis. 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 



NO. 3575 BATETTAL 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. 

USCHAKOV, P. 

1950. [Polychaeta from the Sea of Okhotsk.] Issled. dalnevost morei 
SSSR, no. 2, pp. 140-234, 2 pis., 39 figs. [In Russian.] 



U.S. GOVERNMENT PRINTINS 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 Barnard 

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 Byhlis 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, 
yeUow 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, o£f 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 mUes, 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 III 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. Bai*nard 

Ampelisca brevisimulata J. L. Barnard, 1954a, pp. 33-35, pis. 23-24; 1954b, p. 7; 
1964a, p. 212. 

Material: Albatross 2S37 (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, pis. 14-16; 1954b, p. 3, pi. 1, 
figs. k-1. 

Material: Albatross 2834 (3), 2835 (5), 2837 (6). 

Kecords: 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, pis. 17-18 (includes forma microdentata) ; 1954b, pp. 3-4, pi. 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. 9^2 m. 

Ampelisca cristoides J. L. Barnard 

Ampelisca cristoides J. L. Barnard, 1954a, pp. 29-31, pis. 19-20; 1954b, p. 4, 
pi. 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 
Cahfornia, 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, pis. 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. 3576 AMPELISCIDAE — BARNARD 5 

Ampelisca eschrichtii Kr0yer 

Figure lo 

Ampelisca Eschrichtii Kr0yer, 1842, p. 155. — Sars, 1895, pp. 174-176, pi. 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 eschrichti. — 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 obUquely truncated 
ventral margin of pereopod 5 is a contrast to the transversely trun- 
cated margin in A. macrocephala (see Stephensen, 1935). In A. 
eschrichtii the ventrolateral corneal lens occurs posterior to the anter- 
oventral cephaUc 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 
ui'opod 2 is an invahd 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. eschrichtii 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. eschrichtii eschrichtii 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 cephaHc 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. eschrichtii 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 stroogly 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, pi. 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, pi. 30; 1964a, pp. 213-214. 

Material: Albatross 2837 (1). 
Record: Cedros Island, Baja Cahfornia, 23 fms. 
Distribution: Eastern Pacific Ocean from about Pt. Conception, 
Calif., into the Gulf of California, 33-98 m. 



AMPELISCIDAE — BARNARD 



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, pis. 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, pis. 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, pis. 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. m 

Anipelisca pugetica Stimpson 

Ampelisca pugetica Stimpson, 1864, pp. 158-159. — J. L. Barnard, 1954a, pp. 

49-51, pis. 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, pis. 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 Piiget 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, pis. 10-11; 1954b, p. 3; 1960, 

p. 34; 1964a, pp. 215-216. 
Ampelisca isocomea J. L. Barnard, 1954a, pp. 20-21, pi. 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, Calif., 
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, 
1964a, pp. 14-16, pis. 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 




Figure 1. — Ampelisca schellenbergi Shoemaker, Albatross 3809, female, 7.6 mm: a, head; 
b-f, pereopods 1-5; g, telson; h-j, uropods 1-3; k, I, coxae 1, 2; m, continuation of antenna 
2 from fig. a; n, pleonites 2-6, left lateral (5-6 coalesced). Ampelisca eschrichtii Kr0yer, 
female, 20.0 mm. Albatross 3195: o, 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—67 2 



10 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

4; submarginal medial row of setae on postero ventral edge of article 
2 on pereopod 5 more densely packed than in American population; 
notch and distal end of article 5 v/ith 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 
postero ventral 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 aU 3 areas 
have a few serrate spines in the distal groups on article 5 of pereopods 
3-4. 

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, pis. 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 ; 



AMPELISCIDAE — BARNARD 



11 




Figure 2. — Byblis albatrossae, new species, female, holotype, 15.0 mm, Albatross 3738: 
a, head; b, c, coxae 1, 2; d-h, pereopods 1-5; i ,telson;y, outline of maxilla 2, minus setae; 
k~m, uropods 1-3, latter enlarged. 



12 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

article 4 of antenna 2 longer than peduncle of antenna 1 and much 
longer than article 5 of antenna 2, fiagellum 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 sLxth 
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-thkd. 

Remarks: Pleonal epimera are similar to those of Byblis affinis (in 
Sars, 1895, pi. 65). Mouthparts are sunilar to those of B. gaimardi 
(in Sars, 1895, pi. 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 Byhlis. 
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-foiu-ths 
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 ventraUy, 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 3 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 
(Kr0yer) (in Sars, 1895, pi. 64), but the outer plate of maxiUa 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, pi. 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 



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 




Figure 3. — Byblis ampelisciformis, new species, male, holotype, 12.0 mm, Albatross 3708: 
a, head; b-f, pereopods 1-5; g, h, coxae 1, 2; i, telson; ;-/, uropods 1-3, latter enlarged. 

to antenna 2 or greatly exceedmg 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 15 

these uropods which are suggestive of the following species of Am- 
pelisca: lohata 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 orientaliSf 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-thu-d 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-thu'd. 

Remarks: Mouthparts are like those of B. gaimardi (Kr0yer) (in 
Sars, 1895, pi. 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 



PROCEEDENGS OF THE NATIONAL MUSEUM 



VOL. 121 



Type locality: Albatross 3769, Nagane Saki, off Honshu Island, 
Japan, 40-42 fms, June 5, 1900. 




Figure i.—Byblis orientalis, new species, male, holotype, 11.0 mm, Albatross 3769: a, head; 
h-f, pereopods 1-5; g, h, coxae 1, 2; i, telson; j-/, 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 nun). 



NO. 3576 AMPELISCIDAE — BARNARD 17 

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 tuhicola. — 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 Ampelisoidae collected in the eastern Pacific 

Ocean by the Velero III and Velero IV. Allan Hancock Pacific 

Expeditions, vol. 18, pp. 1-137, pis. 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, pis. 1-2. 

1959. Estuarine Amphipoda. In Barnard and Reish, Ecology of Amphi- 

poda and Polj'chaeta of Newport Bay, California. Allan Hancock 
Found. Publ. Occ. Pap. 21, pp. 13-69, pis. 1-14. 

1960. New bathyal and sublittoral ampeliscid amphipods from California, 

with an illustrated key to Ampelisca. Pacific Nat., vol. 1, no. 16, 
pp. 1-36, 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. 
Barnard, K. H. 

1932. Amphipoda. In Discovery Pteports, vol. 5, pp. 1-326, figs. 1-174, 
pi. 1. 

BULYCHEVA, A. 

1936. New species of Amphipoda from the Japan Sea. Ann. Mag. Nat. 
Hist., ser. 10, vol. 18, pp. 242-256, figs. 1-35. 
Dahl, E. 

1944. Amphipoda of the family Ampeliscidae from Professor Sixten Bock's 
expedition to Japan 1914. Arkiv Zool., vol. 35A, pp. 1-18, figs. 1-10. 
Gtjrjanova, 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.] 
Holmes, S. J. 

1905. The Amphipoda of southern New England. Bull. U.S. Bur. Fisheries, 
vol. 24, pp. 459-529, pis. 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. 489-543, figs. 1-46. 

Jones, M. L. 

1961. A quantitative evaluation of the benthic fauna off Point Richmond, 

California. Univ. California Publ. Zool., vol. 67, pp. 219-320, 
figs. 1-30. 



NO. 3576 AMPELISCIDAE — BARNARD 19 

Kr)z(ter, H. 

1842. Une nordiske Slaegter og Arter af Amfipodernes Orden, henh0rende 
til Familien Gammarina (Forel0bigt Uddrag af et St0rre Arbejde). 
Naturh. Tidsskr., vol. 4, pp. 141-166. 

KUNKEL, B. W. 

1918. The Arthrostraca of Connecticut. Connecticut Geol. Nat. Hist. 
Surv., vol. 6, bull. 26 (Amphipoda), pp. 15-181, figs. 1-55. 
Nagata, 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, pis. 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, pis. 13-17. 

1965. Studies on marine gammaridean Amphipoda of the Seto Inland Sea, 1. 
Publ. Seto Mar. Biol. Lab., vol. 13, pp. 131-170, figs. 1-15. 
PiRLOT, J. M. 

1936. Les amphipodes de I'expedition du Siboga, Deuxieme partie: Les 
amphipodes gammarides, II. Les amphipodes de la mer profonde, 3 : 
Addendum et partie gen^rale. III: Les amphipodes littoraux, 1: 
Lysianassidae . . . Gammaridae. Monogr. 33e (vol. 127) in 
Weber, Siboga-Expeditie, pp. 237-328, figs. 102-146. 
Sars, G. 0. 

1895. Amphipoda. In An account of the Crustacea of Norway with short 
descriptions and figures of all the species, vol. 1, pp. i-viii + 1-711, 
pis. 1-240, suppl. pis. 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, pi. 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. Coutr. 

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. 
Stebbing, 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). In Danish Ingolf- 

Expedition, vol. 3, pp. 101-178, figs. 23-53. 
1933. Amphipoda. In The Godthaab Expedition, 1928. Medd. om Gr0n- 

land, vol. 79, no. 7, pp. 1-88, figs. 1-31. 
1935. The Ampliipoda of N. Norway and Spitsbergen with adjacent waters. 
Troms0 Mus. Skr., vol. 3, pp. 1-140, figs. 1-19. 
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. PhUadelphia [vol. 16], 1864, pp. 153-165. 
Stout, V. R. 

1913. Studies in Laguna Amphipoda. Zool. Jahrb., vol. 34 (Syst.), pp. 
633-659, figs. 1-3. 



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." 

1 



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; pi. 1: fig. 1). Small sandy patches of approxi- | 

mately 100 square feet occur within this rock-coral. Usually this | 

ROCKY-CORAL ENVIRONMENT | 



Echinomeiro lucunter (Linnoeusl 
Diodemo ontillorum Philippi 
Tripneusies ventncosus tLomorck) 
EuCidons tribuioides (Lomorckl 



Clypeosler rosoceus (Linnoeusl 
Clypeosier subdepressus (Gro/l 
Meomo v«nlricosa ILomorck) 

in sondy potches — down 

10 85 fi 
Diodemo oniillorum Philippi 

down 10 85 ft. 




Corel, sponges 
encrusli 



Schiiosler (Pomstet) 

floridiensis Kier and 

GronI 



Figure 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 Comadinia 
echinoptera (Muller) lives commonly in crevices (pi. 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 antUlarum Philippi (pi. 2: 
figs. 4, 5) except in areas of strong current, where Echinometra lucunter 
(Linnaeus) is more abundant. I saw a few specimens of Eucidaris 



NO. 3577 ECHINOIDS — KIER 3 

tribuloides (Lamarck) and Tripneustes ventricosus (Lamarck) (pi. 2: 
fig. 4). 

Below 10 feet, Diadema antillarum was the only echinoid seen in 
any number. The spatangoid Meoma ventricosa (Lamarck) lives in 
the sandy patches together wdth 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, Leodia 
sexiesperjorata (Leske), was found in the shallower parts of these 
sandy areas. Between 25 and 65 feet holothurians were encountered 

SAND ENVIRONMENT 



,,__^___^_^Leodio sexiesperforolo (Leske) 
°''<'"' ^^^"^ilX abundant — buried 




Figure 2 



Patches of sponges with 
Meomo ventricosa (tomorck) 



- 10 ft. 
-20 
-30 

- 40 

- 50 

- 60 

- 70 

- 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 
Eucidaris tribuloides (Lamarck) 
Tripneustes ventricosus (Lamarck) 
Echinometra lucunter (Linnaeus) 
Echinometra viridis Agassiz 
Diadema antillarum Philippi 
Clypeaster subdepressus (Gray) 
Clypeaster rosaceus (Linnaeus) 
Leodia sexiesperjorata (Leske) 
Meoma ventricosa (Lamarck) 
Schizaster (Paraster) floridiensis Kier and Grant 



locality 
8 

8, 10, 18 
8, 14-16, 18 
8 

1-3, 5, 6, 8, 10, 13-17 
2, 6, 10, 13 
10 

7, 9, 12 
2-4, 6, 10, 12 
4 



4 



PROCEEDINGS OF THE NATIONAL MUSEUM 



6I'30' 



ens' 




&°iO' 



6\'\5' 



Figure 3 



NO. 3577 ECEONOIDS — KIER 5 

Locality Data 

numerals = location of stations on map (fig. S) 

1 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) Uoridiensis 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 Tripneustes ventricosus at 30 feet. 

11 Sandy area, small amount of grass, many Oreaster reticulatus (Linneaus), 

two basket stars. 

12 Sandy area, Leodia sexiesperforata (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. 

17 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- 



b 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 
(pi. 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 (pi. 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 

Figure 1. — Coral mass in SO-foot depth at locality 8; the crinoid Comactinia echinoptera 
(Muller) can be seen living in crevices in the coral. Figure 2. — Tripneustes ventricosus 
(Lamarck) in 10 feet of water in dense aggregations, presumably to spawn. 



PROC. U.S. NAT. MUS. VOL. 121 



KIER— PLATE I 









A -'.-._ 




PROC. U.S. NAT. MUS. VOL. 121 







7 * 

-m- m 




fym, 



NO. 3577 ECHESrOIDS — 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 En^^i- 
daris tribuloides (Lamarck) and Tripneustes ventricosus (Lamarck) 
live with this species. 

Echinometra viridis Agassia 

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 

Plate 2 (figs. 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 
Uves even deeper. Most specunens 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 
Mernia 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. 



Figure l.—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. Figure 3.—Meoma 
ventricosa (Lamarck) in 3S-foot depth at locality 6, showing lack of detrital material over 
petals and trail made by the echinoid. Figure i.— Tripneustes ventricosus (Lamarck) 
clinging to rock in 2 feet of water at locality 8; Diadema antillarum Philippi can be seen 
to the right. Figure S— 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 hving 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 (figs. 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 (pi. 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 (pi. 2: 
fig. 3), usually two to three feet long, consisting of a furrow with a 
smaU 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 hving in the sandy 
patches with the echinoids. The one exception occurred at a locahty 



NO. 3577 ECHINOIDS — KIER 9 

85 feet deep (pi. 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- 
simiably, 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 Avith 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, mth 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 five 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 

KiER, 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 pis., 
15 figs. 
Lewis, J. B. 

1958. The biology of the tropical sea urchin Tripneustes esculentus Leske 
in Barbados, British West Indies. Canadian Joum. Zool, vol. 36, 
no. 4, p. 607-621, pis. 8-9, 7 text-figs. 

MORTENSEN, Th. 

1928-1951. A monograph of the Echinoidea, 5 vols, and index. Copen- 
hagen: C. A. Reitzel. 



U.S. GOVERNMENT PRINTING OFFICE 1966 




Proceedings of 
the United States 
National Museum 

SMITHSONIAN INSTITUTION « WASHINGTON, D.C. 

Volume 121 1966 Number 3578 

BREDIN-ARCHBGLD-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. Gagne, 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 Olbiogaster 
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 litnanda, 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. 

1 



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 mth sides of mesonotum; hairs and setae of meso- 
notum brown; scutellum yello^\^sh 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 
yeUow bro^vn, the tarsus shghtly 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 nma. 

Male: Coloration as in female. Eyes nearly touching. Genital 
capside 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 Co^ta 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 9 (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 yello\\dsh tip of the abdomen, I use as 
a specific name the generic name of the flounder knoAvn as the YeUow 
TaU. 

The three New World species of Mycetobia may be differentiated 
by the following key: 

1. Subcosta bare (Nearctic) divergens Walker 

Subcosta with numerous long setae ventrally (Neotropical) 2 

2. Thorax, abdomen, and legs strongly patterned with dark markings; wing 

more than 5 mm long stonei Lane and Andretta 

No distinctive pattern except for yellowish at base and tip of abdomen; 
wing not more than 4 mm long limanda, new species 

Olbiogaster danista, new species 

Female: Frons and occiput dark, grayish pruinose, only the ocellar 
triangle shining black; scape and pedicel yellow, flageUum black; face 
yellow, in some slightly darkened above; palpus and mouthparts 
yellow. Thorax mostly yeUow wdth yellowish and some brown hairs, 
with or A\dthout 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 yeUow stem, black knob. 



DIPTERA — STONE 



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 I, 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 ^dth sterna III-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 Mdder black areas. Terminalia indistinguishable from those of 
the type of 0. antillarum. Lane and Andretta (1958, p. 517, fig. 19) 
figured those for antillarum. 

Distribution: Dominica, Lesser Antilles. 

Holotype: 9, Clarke Hall, Jan. 11-20, 1965 (Wirth). 

Paratypes: same data, 1 9, 3 cf cf ; Clarke Hall, Mar. 1-10, 1965, 1 9, 
1 d", and Mar. 21-31, 1965, 1 cf (Wirth) ; 5 miles south of Pont Casse, 
Apr. 11, 1964 (Flint), 1 9; Clarke HaU, Apr. 12, 1964 (FUnt), 1 9; 
Hillsborough Estate, Mar. 15, 1965 (Wirth), 6 cf cf; Cabrits Swamp, 
Feb. 23, 1965 (Wirth), 1 cf^; Layou R. mouth. Mar. 8, 1965 (Wirth), 
1 cf; Antrim, 1000 ft. Mar. 12, 1956 (Clarke), 1 d^ ; Newfoundland, 
on road to Rosalie, 1000 ft. Apr. 13, 1966 (Gagne), 1 cf . (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 
meson otum and pleura mostly dark, and the stigma not crossing 
vein Ri. From antillarum Lane and Andretta, danista differs in 
having the knob of the halter dark, the sternopleuron entirely yellow, 
and terga IV and V mth 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 fkst 
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 yello^^dsh; 
mesopleuron shiny black wdth a few hairs above and a broad trans- 
verse band of gray poUinosity. Halter with yellow stem, black 
knob. Wing hyaline except for narrow stigmal spot crossing cell Ri. 
Legs mostly dark but with the following yellow: A large patch 



PROCEEDINGS OF THE NATIONAL MUSEUM 



anteriorly on forecoxa; fore- and midfemora; fore- and mid tibiae, 
but slightly darkened. Abdomen with terga shiny black except 
for narrow yellow hind margins on I-V, shghtly widened laterally; 
sterna I-V mostly yellow, a dark area on side of sternum II. Wing 
length 5.5 mm. Terminalia as figured (fig. 1). 

Holotype: d", South Chiltern, 1600 feet, Feb. 19, 1965 (Evans), 
(USNM 69076). 

This species would run in the key of Lane and Andretta (1958) 
to 0. 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 



1 ife 





Figure 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, not narrow and rodlike, and in similans the tergal yellow 
bands are much broader and the steina are mostly black. 

This species is dedicated to the coDector, Howard E. Evans. 

Family Bibionidae 

The only Bibionidae collected on Dominica by the Bredin-Ai'chbold 
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 foiu- shiny stripes on 
notum and fumose wings. It 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 wdth 7 cyhndrical 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 thi-ee 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- 





Ventral 



Dorsal 



Lateral 



Figure 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 mth hairs; hairs of mesonotal furrows very 
small. Wing fumose, the stigma and costal cell darker; veins dark 
wdth short fine hairs; vein R3 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 



b 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 broA\Tiish poUinosity; 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), 
cf , (USNM 68228). 

Paratypes: 1800 ft., Pont Casse, Jan. 12, 1965 (Clarke and Clarke), 
1 cf ; mouth of Layou River, Jan. 20, 1965 (Clarke and Clarke), 1 (^ ; 
light trap, Sylvania, Jan. 23, 1965 (Wirth) , 1 cf ;grassy marsh, Sylvania, 
Jan. 25, 1965 (Wirth), 6 d", 1 9; 2.5 miles west of Pont Casse, Jan. 27, 
1965 (Wu-th), 1 cf , 1 9; 2 miles west of Ponte Casse, Jan. 27, 1965 
(Wirth), 1 cf ; Sylvania, Jan. 28, 1965 (Clarke and Clarke), 2 cf ; 0.5 
miles west of Point Lolo, Jan. 28, 1965 (Wirth), 2 cf ; 1 mile east of 
Pont Casse, Jan. 29, 1965 (Wirth), 1 9; Clarke Hall, Apr. 5, 1965, 
(Flint), 1 cf , 1 9. 

Literature Cited 

Hardy, D. Elmo 

1945. Revision of Nearctic Bibionidae including Neotropical Plecia and 
Penthetria (Diptera). Bull. Univ. Kansas, vol. 30, pp. 367-547, 
13 pis. 
Lane, J., and d'Andretta, C, Jr. 

1958. Neotropical Anisopodidae (Diptera, Nematocera) . Stud. Entom., 
vol. 1, pp. 497-528, illustr. 



U.S. GOVERNMENT PRINTING 0FF1CE:I96S 



I 




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 Harald 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 1824-2S 
(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 oric- 
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 anonj^mous 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 

i.e., pubhcation (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 vahdly 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 



^°- 3579 PORTLAND CATALOGUE — REHDER 3 

we cannot, in my estimation, choose to use some of the vahdly pro- 
posed names and not use the others, unless, of course, the latter names 
are truly nomina dubia. 

I, therefore, publish this hst of names in the conviction that they 
have been vahdly proposed and shoidd enter into the ranks of zoolog- 

A 

CATALOGUE 

OF THE 

PORTLAND MUSEUM, 

LATELY THE PROPERTY OF 

■^^e Duchefs Dowager ^/Portland, 
Decease D; 

Which will be SOLD by AUCTION, 

B Y 

Mr. SKINNER a/^d Co; 

On MONDAY the 24th of APRIL, 1786, 

ANDTHE 




THIRTY-SEVEN roLLowiNc DAYS,^ 
At twelve O'CLOCK, 

SCKCAYS, and the 5th of June, (the Day his Majesty's Birth-Day 
■is kept) excepted ; 

At her late DWELLING-HOUSE, 

In PRIVr-GARD EN, WH irEHALL; 

By order of the 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-Street, Price Five Shillincjs, which will admit 

the Bearer during the Time of Exhibition and Sale. 

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-f amihar 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. 



4 PROCEEDINGS OF THE NATIONAL MUSEUM ^"'" '"^ 

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 51 A of the International Code of Zoological Nomenclature, 
which states that the name of the author of a taxon published anon}^- 
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 vahdly 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, HI pertain to moUusks, 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, and 46 synonyms of 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 Humplirey 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 Hterature of the past 140 years and, 
thus, they should be considered nomina obhta. The remaining 10, 
in all cases but one, have been used as senior synonyms in the past 50 
years and, thus, are available. 



portland catalogue — rehder 
Nineteenth Day's Sale.. 

MONDAY THa 15th of M AY» 1781?. 



SHELLS, CORALS, (fc. 

SOT 

J1JJ4 TP^IFTEEN cards, containing various fpecies of Nerlta, among which are 

J^ Canrcna, albumen, MamiUa, pelorontha, &c. 
igjj Four fine Oftrez, viz. Eburnca, .fquamofa.and 2 lurida, A — all rare 

1926 Eight fine Petrifaftions, viz. 2 Strombus iuOator, Srart/frr'j Fofs. HaxHu.fl. J. 

jig. 64. an Anomia Gryphites, L. an Echinitea from Ftrena, &c; 

1927 Fourteen Bivalves of various genera, among which are a Tellina radiata, » 

Oftrca maxima, /,._^ii', 3 Areas, &c. 
IfjiS Nine cards of various fpecies of Englifti Crabs, viz. Cancer plfum, mimitus^. 

plialangium male and female, Dorfettenfis ; Aftscus Bernhatdus, all JefiriM 

ly Pennant, and the eared Crab, nos ilefcrlpt. 
1929 A very large and fine Buccinum Galea, L. or tun (hell from the Meilitrraneait 
19JO Six different curious Ovaries of marine (hell-fi(h, and 4 eggs of Helix ovipara, 

Lijler, 23. 21. feme |>earls, &c.. 
11951 Seven cards containing various fpecies of Cardium,. viz. one valve ofh»ni- 

cardium, 2. odd. valves ofieiufum, medium, i. muricatum, ciliaic & varic- 

gatum, 5. 
193* Fourteen cards of different fpecies of Needle Buccina, viz. Hcifticuni, flrigila- 

tum, bnceatum, S. torolofum, aniculatum, pertufum, caprtolus, cinercum, 

& fufus,.5. 
1933 A ,fmall Echinus rofaceus and 7. varieties of Echinus orbiculus, L.—fam rare 
J934 Six fine f;ccle5 of Venus, viz. Deflorata, literata, mcrctrix, 8ic. 
iQj^ Two pair of very rare fpecies ofVoluta, viz. Sanguifuga, X.. and turrlcula,, 

S.—fne 

1936 Two fine varieties of ChamaXjigas, Z. or furbelovved clamp 

1937 A beautiful Oftrea nodofa, or.Duck's-foot i'ciflen, and Oflrea plica, L. Icth rare 

1938 Three fine varieties of Murex ramofuf, viz. the Stag's horn, and j others 
jrtjo Three kinds of Pbolas, ««f /CTW</r<//« ^wo</, a fmall rare Sulen, a Mya, ic. 

1940 A fine |>air of Voluta calata and another of I'cpo, S. Mariyn, /-'el. III. 

fi- 768—770 

1941 A pair of very fine Trochui Solaris, L. cr^un fhell, from the fK !nil,'e! — raiv 
J94J Six fine fpecimens of Trochus, viz. 2 of Granofus from Nevj-ZealanM, Martyi:, 

Vol. l.fs' 37' '■" ^ maculatus, L. and 2 others 

VdmumJIactous and ethir JNIMylLS^ VEGETABLES, Vc. in SPIRITS. 
J943 A fine clufter of Lepas Cgnita, S. found in the Briiijb Chaniiei, the animal 
Terebella, iviti it's augur, of the Teredo navalis, or fhip-worm, and fundry 
Orwia of BucciDum uodatum, L. i« 3 iotikt 1944 A 

Figure 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, 

Jine, Lister. 854. 12." 
Tibia insulachorab Roding, 1798 

Strombus Fusus L. Persicus [Lightfoot, 1786], p. 3. 
Rosiellaria 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. Tab. 12. C. 1. 2." 
Xenophora conchyliophora (Born, 1780) 

Trochus onustus [Lightfoot, 1786], p. 4. 
Phorus onustus Reeve, 1842, p. 160. 

Not Trochus omistus Nilsson, 1827 ( = Xenophora sp., Creataceoiis, 
Sweden). 

50 "... 2 Strombus tricornis, Martin 843. 45. List. 873." 

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." 
Voiutocorona nobilis [Lightfoot, 1786] 

Valuta nobilis [Lightfoot, 1786], p. 6. 

Voluta scapha Gmelin, 1791, p. 3468. — not Voluta scafa Lightfoot, 

1786. 
Aulica nobilis Soh\nder. — Abbott, 1962, p. 98. 
Voiutocorona 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 Voiutocorona 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 Isognoma (or Isognomon) . 
See lots 968 and 3041, wherein the now generally accepted form 
Isognomon is used. Ostrea isognomon L. ( = 0. 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 Roding, 1798, p. 186. 

Abbott (1954, p. 439), following Dall (1921, p. 99), erroneously used 
this name for T. sanguinolenta Gmelin, in which he has been followed 
by Shikama (1964, pi. 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 *§. bertini Pilsbry and Lowe, as Keen 
(1958, p. 188) thinks possible. 

See lot 1360, wherein Lightfoot spells the trivial name correctly: 
cruenta. 

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 "Echinus 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 oj 
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], pi. 8. 

353 "Two fine species of Placuna, S. viz. placenta, and Ephippium, 
the Chinese Window, and PoHsh 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, Jig. lOe." 
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, pi. 61, 
D. . . . rare." 
Pila urceus (Mueller, 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) 

Limax 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) 

Valuta ponderosa [Lightfoot, 1786], p. 25. 

Xancus pyrutn Roding, 1798, p. 134 (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] 

Valuta virescens [Lightfoot, 1786], p. 26. 

Plejona muta Roding, 1798, p. 60. 

Voluta pohjzonalis 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) 

Valuta arausiaca [Lightfoot, 1786], p. 26. 

Voluta arausiaca Shaw, 1790, pi. 52. 

Voluta vexillum Gmelin, 1791, p. 3464. 

Harpulina arausaica [sic] Solander. — Shikama, 1963, pi. 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, jig. 25. 

?i." 

Placostylus elongatus (Lightfoot, 1786) 

Limax fihratus Martyn, 1785, fig. 24 (nonbinominal) . 
Valuta elongata [Lightfoot, 1786], p. 30. 
Bulimus bovinus Bruguiere, 1792, p. 345. 
Ellobium australe Roding, 1798, p. 106. 

Not Valuta elongata Schroeter, 1804 {=lAurinia sp.); not Voluta 
elongata Swainson, 1821 {=Alcithae 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, earUest names, those of 
Lightfoot, Bruguiere, and Roding. Since the name fihratus 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) amphora (Lightfoot, 1786) 

Valuta amphora [Lightfoot, 1786], p. 30. 

Cymbium flammeum Roding, 1798, p. 151. 

Melo diadema Lamarck, 1811, p. 57. 

Cymbium {Cymbium) ampharus (Solander). — Shikama, 1963, pL 85. 

915 "Hehx Otis an extremely rare species, &.guredhj Favanne, tab. 
QS.fig.n." 
Lampadion otis (Lightfoot, 1786) 

Helix atis [Lightfoot, 1786], p. 38. 

Helix labyrinthus Lamarck, 1792, p. 352. 

Lampadion labyrinthus Roding, 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 Rodmg, 1798. Winckworth (1945, p. 141) lists as 
type Lampadion labyrinthus Roding, citing Gray (1847, p. 173) as 
authority for this type designation. Although Gray gave Helix otis 
as the type of the genus, which was not one of the originally included 
species, Winckworth's action is vahd ("International Code of Zoologi- 
cal Nomenclature;" art. 69(a) (iv)), Wurtz (1955, pp, 106-107), 
apparently overlooldng Winckworth's statement, designated as type 
the same species of Roding, Lampadion labyrinthus, fixing this name 
on the figure in Favanne (1780), cited by Roding, 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 Roding cited the wrong figure in Favanne for his species. 
Roding apparently meant to cite F 11 on plate 63 but referred instead 
to "fig. 2," possibly reading the 11 as a Roman II. That Roding 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 pi. 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, Marty n, vol. III. 

772, 773 .. . ." 

Cymbium (Melo) melo (Lightfoot, 1786) 

Voluta melo [Lightfoot, 1786], p. 41. 

Voluta anguria Lightfoot, 1786, p. 64. 

Voluta indica Gmelin, 1791, p. 3467. 

Cymhium maculatum Roding, 1798, p. 152. 

Cymbium melo Roding, 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) 

Valuta scafa [Lightfoot, 1786], p. 41. 

Adanson's figure is cited by Lightfoot for Solander's "variety b" 
of his Voluta scaja, which is validated in lot 3039 by a reference to a 
figure in Martini. Both of these figures represent Voluta cymhium 
Linnaeus. 

990 "Two fine PateUa Mytihformis, Humphrey's Conch, pi. HI. 
Jig. 9 from Falkland's Islands . . . ." 
Nacella mytilina (Helbling, 1779) 

Patella mytiliformis [Lightfoot, 1786], p. 42. 

Patella mytilina Gmehn, 1791, p. 3698. 

Not Patella mytiliformis Gmehn, 1791 {=Crepidula sp.). 

1001 "Two fine specimens of Area fusca, S. Gualt. 87. G. one of 
them affixed to a pinna." 
Nomen dubium (figure indeterminable) 

Not Area fusca Bruguiere, 1789 { = Barbatia amygdalumtostum 

(Roding, 1798). 



NO- 3579 PORTLAND CATALOGUE — REHDER H 

Solander's name was assigned by earlier authors to various species. 
Donovan (1803, p. 158) gave Solander's name to specimens he beheved 
to be what is now called Area imbricata Bruguiere. In this he was 
followed by Montagu (1808, p. 51), who assigned certain British 
specimens to this species, obviously confusing Area tetragona PoH with 
imbrieata, since he, hke Donovan, cites Lister's figure (1732, pi. 367, 
fig. 207) of imbrieata for Arcajusca. Dillwyn (1817, p. 227) actually 
places A. Jusea Solander (mss.) in the synonjrmy of Area imbrieata 
Bruguiere, in which he is followed by Iredale (1961, p. 92). Lamy 
(1907, p. 41), on the other hand, places the "Portland Catalogue" 
name m the synonymy of A. tetragona PoH, 1795, and Dall (1921, p. 
98) equates the species with jusea 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 styHzed and 
could represent any of several species of Area. 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 
inferior) . . . ." 

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 plebeius 'Solander.' — Dillwyn, 1823, p. 22. 

Tagelus plebeius Solander. — Abbott, 1954, p. 440. 

1046 "A pair of fine Conus Augur, S. or dotted Cone, Knorr. VI. 
tab. 13. fig. 6." 
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 " . . . and hians, S. ibid 18. B. both rare." 
Argonauta hians [Lightfoot, 1786] 

Argonaula hians [Lightfoot, 1786], p. 44. 
Argonauta hians Dillwyn, 1817, p. 334. 
Argonauta hians Solander. — Tryon, 1879, p. 136. 

1206 "Two Cypraea pantherina, S. List. 681. 28 . . . ." 
Cypraea pantherina pl/ightfoot, 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, List. 62. 60 . . . ." 
Sagda epistylium (MuUer, 1774) 

Trochus alveolatus [Lightfoot, 1786], p. 52. 

1242 "Four varieties of Ostrea perna, L. (Isognomon, S.) . . . ." 
Isognomon CLightfoot, 1786] 
See lot 968. 

1266 "Three fine species of Balanus . . . Pyramidalis, S. Ellis, 
Jig. 9." 
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, pi. 34). 

1301 "Two very large specimens of Patella fungoides or Mushroom 
limpet, from the Cape of Good Hope. Humphrey's Conch, pi. 
lY.Jig.l6." 
Patella (Scutellastra) barbara Linnaeus, 1758 

Patella fungoides [Lightfoot, 1786], p. 55. 

Not Patella fungoides Roding, 1798 {—Acmaea jamaicensis Gmelin, 

1791). 

1315 "Four fine specim.ens 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 
scarce . . . Lister, 328. 165." 
Dinocardium robustum [Lightfoot, 1786] 

Cardium robustum [Lightfoot, 1786], p. 58. 
Cardium magnum Born, 1778, p. 34 (not Liun^, 1758). 
Cardium ventricosum Bruguiere, 1789, p. 228. 
Dinocardium robustum Solander. — Abbott, 1954, p. 401. 



NO. 3579 PORTLAND CATALOGUE — REHDER 13 

1359 "Three fine and rare species of Patella, viz. two of Mytili- 
formis, Humphrey's Conchology, 3. 9 . . . ." 
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 III. 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. 1. 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 pi. 79. H." 
Pterynotus (Marchia) elongatus (Lightfoot 1786) 

Murex elongatus [Lightfoot, 1786], p. 65. 
Murex clavus Kiener, 1841, p. Ill (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.-" 
]VlodioIus 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, Humphreifs 
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 unhkely 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. 
KnorrlU. 12. 5" . . . ." 
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) 

Vohda filosa Born, 1778, p. 212. 
Voluta filosa [Lightfoot, 1786], p. 76. 

1709 "Patella Calyptra, vol. I. Alartyn, fig. 18, k . . . ." 
Nomen dubium 
This figiu"e 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 Solancler. — Abbott, 1950, p. 204. 
Not Voluta angulata Swainson, 1821 = Zidona dufresnei (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 
Xancus Roding, 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. 
Comis arachnoideiis Gmelin, 1791, p. 3388. 
Conus araneosus Solander. — Tomlin, 1937, p. 213. 

1718 "A large and fine, and 2 small specimens oj Patella Unguis, L. 
(Mytilus Lingua, or green Duck's-Bill Limpet, S.) Humphrey's 
Conchology,Pl.2.fig.2.fro7nAmboyna . . . extremely scarce." 
Lingula lingua (Lightfoot, 1786) 

Mytilus lingua [Lightfoot, 1786], p. 77. 

Pharetra monoculoides Roding, 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 
Roding, 1798. 

Later (see lot 3717) Lightfoot lists the same shell as Patella unguis 
L., a reminder of the long-standing but erroneous appHcation of 
Linnaeus' name to this species of Lingula. 

1873 "A neat small pair of Voluta Ancilla, S. from the Straits oj 
Magellan, very rare, D'Avila's Catalogue, vol. I. j^l- 8. fig. 5." 
Adelomelon ancilla (Lightfoot, 1786) 

Valuta ancilla [Lightfoot, 1786], p. 84. 

Voluta spedabilis 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. 1. fig. 30. 9." 
Astraea imperialis (Gmehn, 1791) 

Trochus Solaris [Lightfoot, 1786], p. 86. 
Not Trochus Solaris Linnaeus, 1766. 

1930 "— 4 eggs of Hehx ovipara. Lister, 23. 21." 
Strophocheilus oblongus (MuUer, 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. Jig, 
768-770." 
Cymbium pepo (Lightfoot, 1786) 

Valuta pepo [Lightfoot, 1786], p. 87. 

Voluta neptuni Gmelin, 1791, p. 3467. 

Voluta navicula Gmelin, 1791, p. 3467. 

Cymbium (Cymba) pepo Solancler. — Abbott, 1962, p. 103. 

1942 "... Trochus . . . granosus from New Zealand, Martyn, vol. 
I. fig. 37. T. 2 . . . ." 
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 
vaUdated 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 
figure . . . ." 
Conchoderma virgata (Spengler, 1790) 

Lepas vittata [Lightfoot, 1786], p. 88 [nomen oblituml. 

Lepas virgata Spengler, 1790, p. 207. 

Lepas vittata Wood, 1815, p. 69. 

Cinevas vittata Leach, 1824, p. 170. 

Conchoderma virgata (Spengler). — Pilsbry, 1907, p. 99. 

The "IV" 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, 
pi. 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 Murex 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. (Lepas S.) . . . ponderosus, S. Lister. 
442. 284." 
Tetraclita squamosa stalactifera Lamarck 

Balanus ponderosus [Lightfoot, 1786], p. 89 [nomen oblitum]. 

Balanus stalactiferus Lamarck, 1818, p. 394. 

Tetraclita squamosa stalactifera Lamarck. — Pilabry, 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. Jig. 19. 1." 
Mitra terebralis Lamarck, 1811 

Valuta incom-pla [Lightfoot], 1786, p. 96 [nomen 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. 
18.1." 
Argonauta nodosa [Lightfoot, 1786] 

Argonauta nodosa [Lightfoot, 1786], p. 96. 
Argonauta tuberculata Roding, 1798, p. 71. 
Argonauta tuberculata Shaw, 1812, pi. 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. 
Vol. 3. t. 65./ 1. 2." 
Cymbium (IVlelo) 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. Jig. 2, 3, and 
4. of the Bucc. Harpa L." 
Nomen dubium 

Not Harpa testudo Donovan, 1822 {=H. ventricosa 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 {articidaris Lamarck, 1822; major Roding, 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 appHed. Donovan later applied 
the name H. testudo to still another species {Harpa ventricosa Lamarck, 
1816). 

Dall (1921, p. 98), when he identified Solander's name with Cassis 
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. G . . . ." 
Meretrix meretrix (Linnaeus, 1758) 

Venus punctata [Lightfoot, 17861, p. 98. 
Not Vemis punctata Linnaeus, 1758. 

2158 "A fine Area nodulosa, S. or studded Ark, Gualt. 87. 
E . . . . " 
Anadara granosa (Linnaeus, 1758) 

Area nodulosa [Lightfoot, 1786], p. 98. 

"Not Area nodulosa Miiller, 1776 { — Acarscabra (Poli, 1795), fide Lamy, 
1907, p. 95). 

2190 "Two curious and rare species of Mya, viz. Complanata, 
List. 150. 5. and . . . . " 
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 Area, 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. 
.... List. 414. 258." 
Panope glycymeris (Born, 1778) 
See lot 626. 

2216 "A very fine cluster of Lepas Cornu Copiae, S. D'Argenville, 
26. D. . . . rare." 
Mitella cornucopiae (Lightfoot, 1786). 

Lepas cornucopiae [Lightfoot, 1786], p. 101. 

Lepas pollicipes Gmclin, 1791, p. 3213. 

PoUicipes cornucopia Leach, 1824, p. 171. 

Pollicipes cornucopia: Darwin, 1851, p. 298. 

Pollicipes cornucopia: Gruvel, 1902, p. 223. 

Mitella pollicipes: Pilsbry, 1907, p. 5. 



NO. 3579 PORTLAND CATALOGUE — REHDER 19 

As will be seen by the above sjmonymy, the trivial name comuco'piae 
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 anmdatus [Lightfoot, 1786], p. lOL 

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 
America . . . rare." 
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 IVIartyn 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 . . . . " 
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. 
III. 917." 
Turbinella pyrum (Linnaeus, 1758) 

Valuta gravis [Lightfoot, 1786], p. 103. 
Voluta gravis Wood, 1818, p. 101. 

This is not T. rapa Lamarcls, 1822 (=ponderosa Lightfoot, 1786) 
as Dall states (1921, p. 131). 

2284 "... IVIurex plicatus, or plicated IVlurex, from Falkland's 
Islands, undescrihed. Favanne, tab. 79. /...." 

226-75&— 67 3 



20 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Trophon piicatus (Lightfoot, 1786) 

Murex piicatus [Lightfoot, 1786], p. 104. 
Buccinum lacinialum Martyn, 1784, fig. 42 (nonbinomial). 
Murex lamellosus Gmelin, 1791, p. 3536. 
Trophon laciniatus Martyn. — Strebel, 1904, p. 199. 
Murex piicatus S.— Dall, 1921, p. 99. 
Not Murex piicatus Gmelin, 1791 {="Thais" sp.). 
Not Murex piicatus Dillwyn, 1817 {—Coralliophila erosa (Roding), 
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 Bxiccinum muricatum Schroter, 1805 {—Hebra horrida Dunker, 
1847). 

Shikama (1963, pi. 49, fig. 3) follows Dall (1921, p. 125) in consider- 
ing this to be the same as Colubraria ohscura (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 
Conch. 3. 8 . . . ." 
Patella barbara Linnaeus, 1758 

Patella gorgonica [Lightfoot, 1786], p. 105. 

2302 ". . . [Patella] Pulchra, or Beauty, ibid. 2. 8 . . . ." 
Patella miniata Born, 1778 

Patella pulchra [Lightfoot, 1786], p. 105. 

2302 ". . . and a young one of [Patella] Oculus hu-ci, ibid. 2. 6." 
Patella oculus Born, 1778 

Patella oculushirci [Lightfoot, 1786], p. 105. 

2315 "A fine specimen of Voluta incrassa, S. Martyn, Vol. li- 
very rare, f. 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 Miu-ex fimbriatus, or 
furbelowed Miu-ex, from Falkland's Island, Martyn, Vol. 1. 
iig. 6. c. Favanne, 37. H.I." 



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. Alartyn. Vol. II. 676." 
Conus araneosus [Lightfoot, 1786] 
See lot 174. 

2230 "Two extremely curious and rare species oj Cypraea, viz. 
[error Pustulata, or orange warted Cowry, from China, Lister, 
for 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, pi. VII [Error for XII] fig. 15. from Madagascar . . . 
very rare.' 
Penicillus (Warnea) attrahens (Lightfoot, 1786) 

Serpula attrahens [Lightfoot, 1786], p. 106. 
Aspergillum vaginiferun Lamarck, 1818, p. 430. 
Serpula attrahens S.— Dall, 1921, p. 129. 

2481 "A fine Trochus sulcatus, from New Zealand, Naturforscher, 
Vol. IX. tab. III. Jig. 5. 6. Martyn, Vol. I. Jig. 33. [error 
for 35] r . . . ." 
Cookia sulcata (Gmelin, 1791) 

Trochus sulcatus IVIartyn, 1784, fig. 35. 
Trochus sulcatus [Lightfoot, 1786], p. 13. 
Trochus cookii 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 Gmehn, 1791 was placed on the 
"Official List of Specific Names in Zoology" in Opinion 479 (Inter- 
national Commission of Zoological Nomenclatm^e, 1957: p. 372), 
it was not reahzed that Lightfoot's name was the first vahd taxon for 
this species after Martyn. It was also overlooked that Trochus 
cookii Gmelin, 1791 is a synonym of Turbo sulcatum Gmelin. 

2516 "Isognoma rigida, S. from Pulo Condore, List. 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, 178G], 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 

Conns 7nappa [Lightfoot, 1786], p. 116. 

Not Conus mappa Crosse, 1858 (—C. eldredi Morrison, 1935). 

2961 "A large and very fine variety of Buccinum calcaratum, 
S. Gualt 31. F. . . . rare." 
Melongena (Volema) hippocastanum (Linnaeus, 1758) 

Buccinum calcaratum [Lightfoot, 1786], p. 133. 
Volema nuxmoschata Roding, 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 Mm'icidae (Thaisinae) to which the Linnean name was 
given by many later authors. 

2965 ". . . Cypraea seriata, (rather a young one of Tigiis L.) . . . ." 
Cypraea tigris Linnaeus, 1758 . 

Cypraea seriata [Lightfoot, 1786], p. 133. i 

2965 ". . . [Cypraea] nebuiosa, S. (a young one of Mauritiana, L.)." 
Cypraea mauritiana Linnaeus, 1758 

Cypraea nebuiosa [Lightfoot, 1786], p. 133. 

Not C. nebuiosa Gmelin, 1791 ( = C stercoraria Linnaeus, 1758). 

2967 "Two fine specimens of Strombus truncatus, an undescribed 
species of L. showing the different stages oj growth, from the 
E. Indies, rare . . . D'Avila, vol. I, tab. 12. 14." 
Lambis truncata truncata (Lightfoot, 1786) 

Strombus truncatus [Lightfoot, 1786], p. 133. 

Strombus bryonia GmeHn, 1791, p. 3520. 

Lambis davilae Roding, 1798, p. 66. 

Strombus truncatus Dillwyn, 1817, p. 659. 

Lambis truncata truncata (Humphrey, 1786). — Abbott, 1961, p. 155. 

2983 "Three curious species of Mya, ^dz. . . . 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. 

Unto crassa cytherea Kuster. — Haas, 1940, p. 129. 

Mya ovalis S.— 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 Kuster, 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, pi. 
2. Jig. 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. IH. 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, ibid [Seba III. 
tab. 38.], Jig. 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 perjection, 
from Guinea . . . Martyn, vol. Ill 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, I . . . very rare." 

Atrina rigida (Lightfoot, 1786) 

Pinna rigida [Lightfoot, 1786], p. 136. 

Pinna rigida Dillwyn, 1817, p. 327. 

Atrina rigida Solander. — Turner and Rosewater, 1958, p. 312. 

3041 "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 TeUina marginalis, S. List. 387." 
Tellina laevigata Linnaeus, 1758 

Tellina marginalis [Lightfoot, 1786], p. 137. 

3054 "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 oj 

Magellan, extremely scarce, D'Adla, vol. I. pi S. Jig. s." 

Adelomelon ancilla (Lightfoot, 1786) 
See lot 1873. 
3074 ". . • ^ stone with various specimens of Venus arctica, 

S. in it, List. 426. 427 .... " 

Hiatella arctica (Linnaeus, 1758) 

Venus arctica [Lightfoot, 1786], p. 138. 

3091 "A cluster of three, and a single specimen of Ostrea purpurea, 
S. from New Holland, very rare — Born. mus. caes. Tab. 6. Jig. 
11. 12." 
Ostrea cucuUata Born, 1778 

Ostrea purpurea [Lightfoot, 1786], p. 139. 

3093 "A large and fine Buccinum monodon, S. or Unicorn, from 
Terra del Fuego, Martyn, vol. I. fig. 10 e . . . . " 
Acanthina monodon (Pallas, 1774) 
See lot 372. 
3117 "A large fine pak of PateUa oculus hirci, or Goat's-eye Limpet, 
one oj them in its native state, the other polished, from the Cape 
oj 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 SheU, from China . . . rare." 
Placuna [Lightfoot, 1786] 
See lot 3028. 



I 



NO. 3579 PORTLAND CATALOGUE — REHDER 25 

3142 "A fine pair of Voluta muricata, S. one with the epidermis on, 
Lister, 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, >S^. Lister, 
841. 69. from China . . . very rare.^' 
Terebra taurinum (Lightfoot, 1786) 

Buccinum taurinum [Lightfoot, 1786], p. 142. 
Epitonium feldnianni Roding, 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. 
l.pl. 5. fig. I." 
Turbo (Batillus) cornutus [Lightfoot], 1786 

Turbo cornutus [Lightfoot, 1786], p. 147. 

Turbo cornutus Gmelin, 1791, p. 3593. 

Turbo cornutus Solander. — Shikama, 1963, pL 12. 

3307 "A fine young specimen of Strombus truncatus D'Avila, vol. I. 
tab. 12 . . . ." 

Lambis truncata truncata (Lightfoot, 1786) 
See lot 2967. 

3356 "Four curious species of Buccina, viz. . . . one of Iris, S. 
Martyn, vol. l.fig. 2. b. . . ." 
Latirus iris (Lightfoot, 1786) 
See lot 30. 

3384 "Patella Auricularia, an extremely rare species, from Amboyna 
. . . Rumyh. 40. A^." 
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 (Miiller, 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. Jig. 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 Rodiiig, 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 pimctulatus of Martyn, vol. I. Jig. 36. r. . . ." 
[error Calliostoma (Mauriella) punctulatum Martyn, 1784 

lOr Trochus punctulatus [Lightfoot, 1786], p. 155. 

3394] Trochus diaphanus GmeUn, 1791, p. 3580. 

Not Trochus punctulatus GmeUn, 1791 {—ICantharidus turriculum 
Phil., 1855). 

3494 ". . . [Trochus] granosus, ibid. 37. r. . . ." 
Modelia granosa (Mart3rn, 1784) 

See lot 1942. 

Both of the above two Alartyn 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, 
II. 687. and 688. alljrom 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. Jig. 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 ajer (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 27 

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 SolefeUina Blainville, 1824. H. and A. Adams 
(1856, p. 392) used it correctly in place of Soletellina, as did StoHczka 
(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 tjrpe, 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. Jig. 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 aUied to Helix Ovipara, List. 1055. 1. the 
country unknoum." 

Strophocheilus ovatus (Miiller, 1774) 
See lot 3388. 

3561 "A very fine specimen of Bulla Zonata, S Born. 

Mus. Caes. tab. 9. Jig. 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 Tiirton, 1834 { = Scaphander lignarius L., 1758, 
young). 



28 PROCEEDINGS OF THE NATIONAL MXTSEUM vol. 121 

The shell figured by Kira (1959, p. 104, pi. 39, fig. 15) as li. zonata, 
lacks the white band, and may be a distinct species {?H. injlata 
Dunker, 1877). 

3624 "Seven curious specimens of Solens, viz. a small one of 
Rostratus, Valentyn. Bivalves, no. 5 . . . ." 
Hiatula diphos (Linnaeus, 1771) 
See lot 3487. 

3655 "Two very fine specimens of Strombus truncatus, showing 
the dirfferent stages of growth, from China, very rare—D'Avila, 
vol. I. pi. 12 and 14." 

Lambis truncata truncata (Lightfoot, 1786) 
See lot 2967. 

3696 "A very fine pair of Voluta Incrassata, S. extremely scarce . . . 
Martyn, vol. II. 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. 
776." 

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. pi. 6. fig. 11. 12." 
Ostrea cucullata Boon, 1778 
See lot 3091. 

3744 "A very fine and large specimen of Placuna Ephippium, 

S. (Anomia, L.) 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 (Muller, 1774) 
See lot 1930. 



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 
rare . . . Rum. 18. B." 
Argonauta hians [Lightfoot, 1786] 
See lot 1055. 

3751 "A fine specimen of Voluta ^drescens, S. or brindled Music 
from Guinea, very rare . . . Martin, vol. III.^^. 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 . . . Bom, mus. caes. tab 9 
Jig. 1." 

Hydatina zonata (Lightfoot, 1786) 
See lot 3561. 

3761 "A large and fine specimen of Venus nimbosa, S. from Florida, 
very rare . . . Favanne, pi. 49, fig. I." 
Macrocallista nimbosa (Lightfoot, 1786) 

Venus nimbosa [Lightfoot, 1786], p. 175. 
Venus gigantea Gmelin, 1791, p. 3282. 
Macrocallista nimbosa Solan der. — Clench, 1942, p. 5. 

3794 "Nine curious and rare species of Hehx, among which are . 
insignita. Lister, 67, 68 . . . the last undescribed." 
Carocolus marginella (GmeHn) subsp. 

Helix insignita [Lightfoot, 1786], p. 176 [nomen oblitum]. 

3798 "A very fine specimen of Ostraea purpm-ea, S. rich in colour, 
from New Holland, exceedingly scarce. Bom. mus. caes. tab 
6. fig. 11,12." 
Ostrea cucullata Born, 1778 
See lot 3091. 

3802 "A very fine specimen of Hehx undata, a carinated umbilicated 
Land Shell, the Country unknown. Lister, 76. Favanne, 
tab. 63. ^i/. G. 3." 
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 Gmehn's name by undata 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 imdulatus, or waved Emerald 
Turbo, extremely scarce, from Van Dieman's Land, New 
Holland. Marty n. Vol. 1. Jig. 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, 
pi. 5, Jig. 5. which was taken Jrom this shell." 
Umbraculum umbraculum (Lightfoot, 1786) 

Patella umbraculum [Lightfoot, 1786], p. 178. 

Patella sinica Gmehn, 1791, p. 3705. 

Patella umbellata Gmelin, 1791, p. 3720. 

Patella umbracula Roding, 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. 
772." 
Conus pulcher pidghtfoot, 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, S.) from 
[Error China . . . extremely rare." 
for Placuna [Lightfoot, 1786] 

3847] See lot 1315. 



NO. 3579 PORTLAND CATALOGUE — REHDER 31 

3853 "A very fine specimen of Patella pulchra, Hum'phrei/s Conch. 
pi. 2. fig. 8 . . . the first is a native of the Cajpe oj Good 
Hope. . . ." 

Patella miniata Bom, 1778 
See lot 2302. 

3866 "Two fine specimens of a cm-ious 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 III. 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. 
22. 1." 
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 Dai-win, 1854 

Balanus tulipa [Lightfoot, 1786], p. 183. 

Balanus tulipa Poli, 1791, p. 24. 

Not Lepas tulipa Miiller, 1776 (=5. tintinnabulum L., 1758). 

3913 "A capital specimen of Voluta Imperialis, S. or Great horned 
Wnd Music, from Luconia, extremely scarce . . . Martin, 
vol. IV. 934. 935." 
Aulica imperialis (Lightfoot, 1786) 

Valuta imperialis [Lightfoot, 1786], p. 183. 

Voluta imperialis Lamarck, 1811, p. 62. 

Aulica imperialis Solander. — Abbott, 1962, p. 98. 



32 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

3924 "A large and finely coloured specimen of Helix Undata, an 
undescribed species of Land SnaU, from whence un- 
known . . . Favanne., pi. Q3,Jig. Q. 3." 
Solaropsis undata (Lightfoot, 1786) 
See lot 3802. 

3926 "A large and fine specimen of Valuta Nobilis, S. an extremely 
rare, and very beautiful shell of the WUd 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 unkno\^Ti, Humph. Conch, pi. 11, fig. 4. unique." 
Vermicularia tortuosa (Lightfoot, 1786) 

Serpula tortuosa [Lightfoot, 1786], p. 184. 

Vermetus costalis 'Rousseau' Chenu, 1844, Vermetus, pi. 3. 

Vermetus (Vermicularia) tortuosus Solander. — Tryon, 1885, p. 186. 

3947 "A fine specimen of Area Labiata, S. or valved Ark, called by 
the French, Coquelochon de Moine, D'Avila, vol. I. pi. 18. 
its country unknown." 
Cucullaea labiata (Lightfoot, 1786) 

Area labiata [Lightfoot, 1786], p. 185. 
Area concamera Bruguiere, 1789, p. 102. 
Area eucullus Gmelin, 1791, p. 3311. 
Cucullaea auriculifera Lamarck, 1801, p. 116. 
Area eoncamerata Dillwyn, 1817, p. 232. 
Cucullaea labiata (Solander). — Shikama, 1964, pi. 20, fig. 2. 
Not Area 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, extremely 
rare . . . Seba III. 94. The largest figure." 
Euphus 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 pan- of Helix Alba a terrestrial Shell from the E. 
[Error Indies, one of them an exceeding great variety, being left 
for handed, or turning the contrary way. — Lister 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 Hehx 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 Midler's 
species. 

3982 "A very fine Trochus tectus, the only perfect specimen known, 

from the Island oj All Saints, in the West Indies. A. bad 
specimenis Jigured in Lister, Q28. 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 Gmehn's Trochus tectum, since the latter trivial name is a noun 
(Latin tectum— rooi) in apposition to the genus, while Lightfoot 
used the adjective ^edws(= 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. Jig. 6." 
Polyphyllia talpina (Lamarclv, 1801) 

Madrepora agnus [Lightfoot, 1786], p. 188 [nomen oblitum]. 
Fungia talpina Lamarck, 1801, p. 370. 
Fungia talpa Lamarck, 1816b, p. 237. 
Polyphyllia talpina.— Thial, 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] 

Murex tribulus var. pecten [Lightfoot, 1786], p. 188. 
Murex histrix Reding, 1798, p. 145 (not Murex hystrix Linnaeus, 
1758). 



34 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Aranea gracilis Perry, 1810, pi. 47 (not Murex gracilis Montagn, 

1803). 
Aranea triremis Perry, 1811, pi. 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 Murex 
tenuispina Lamarck is tempered by the fact that Perry's name 
triremis had priority and has been used by recent workers for this j 
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, ex- | 

tremely scarce, the country unknoum. Seha III, pi. 14. Jig. 5. 6. ' 

and Favanne, pi. 58. jig. A. 2." ^ 
Plagiobrissus grandis (Gmehn, 1788) 

Echinus maximus [Lightfoot, 1786], p. 188 [nomen 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 pedoraHs, 
long in use, must yield to Gmelin's earlier name. 

4017 "A fine specimen of Conus Architalassus, S. a cmious granu- 
lated variety of the High Admiral, extremely scarce, from 
Amhoyna. Argenv. Suppl. tab. 1. jig. M. N. Martyn II. tab 
min. 2Q.jig. 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, extremely rare. Seba. III. pi. 62. jig. 3. 
Favanne 22. A. 2." 
Strombus sinuatus [Lightfoot, 1786] 

Strombus sinuatus [Lightfoot, 1786], p. 189. 
Lambis lobata Roding, 1798, p. 65. 
Strombus lacinialus 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 
figured in Favanne, pi. 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 
III. tab. 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 
mytiUformis, P.=Nacella mytilina (Helbling) 
oculus hirci, P. = Patella oculus Born 
pulchra, P. = Patella miniata Born 

Trochidae 

annulatus, Trochus=CaUiostoma annulatum (Lightfoot) 
canaHculatus, 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 imperiahs (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=Xenopliora conchyliophora (Born) 

Strombidae 

persicus, Strombus fusus= Tibia insulaechorab (Roding) 
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 miu-icata (Lightfoot) 

maculosa (Gmelin) 
pustulosum, Buccinum=Argobuccinum pustulosum (Lightfoot) 

argus (Gmehn) 

Muricidae-Muricinae 

elongatus, Murex=Pterynotus (Marchia) elongatus (Lightfoot) 

clavus (Kiener) 

pecten, Murex tribulus=Murex pecten [Lightfoot] 

triremis Perry 
tenuispina Lamarck 

Muricidae-Thaisinae 
monodon, Buccinum=Acanthina monodon (Pallas) 

Muricidae-Trophoninae 

fimbriatus, Murex =Trophon geversianus (Pallas) 
plica tus, Murex =Trophon pUcatus (Lightfoot) 

laciniatus (Martyn) 

Melongenidae 
calcaratum, Buccinum== Melongena (Volema) hippocastanum (Linnaeus) 



NO. 3579 PORTLAND CATALOGUE — REHDER 37 

Fasciolariidae 
iris, Bucciniim=Latiriis iris (Lightfoot) 

Olividaee 

incrassata, Voluta=01iva incrassata (Lightfoot) 

Mitridae 

filosa, Voluta=Mitra (Cancilla) filaris (Linnaeus) 
incompta, Voluta=Mitra terebralis Lamarck 

Turbinellidae 

angulata, Voluta=Turbinella angidata (Lightfoot) 
gravis, Vokita=Turbinella pyrum (Linnaeus) 
ponderosa, Voluta=Turbinella ponderosa (Lightfoot) 

rajpa Lamarck 

Vasidae 

muricata, Voluta=Vasum muricatimi (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 imperiahs (Lightfoot) 
melo, Voluta=Melo melo (Lightfoot) 
nobilis, Voluta=Volutocorona nobiUs (Lightfoot) 

scapha (Gmelin) 
pepo, Voluta=Cymbium pepo (Lightfoot) 
scafa, Voluta=Cymbium cymbimn (Linnaeus) 

Volutidae-Zidoninae 

arausiaca, Voluta==Harpulina arausiaca (Lightfoot) 
ancilla, Voluta=Adelomelon ancilla (Lightfoot) 

226-756—67- 4 



38 PROCEEDINGS OF THE NATIONAL MUSEUM 

Conidae 

araneosus, Conus=Conus araneosus [Lightfoot] 
architalassus, Conns =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) 

Bulimuhdae 

elongata, Voluta=Placostylus elongatus (Lightfoot) 

Hhratus (Martyn) 

Strophocheilidae 

erubescens, HelLx— Strophocheilus pudicus (Miiller) 
ovipara, Helix == Strophocheilus oblongus (MuUer) 

Sagdidae 
alveolatus, Trochus=Sagda epistylium (Miiller) 

Camaenidae 

insignita, Helix =Caracolus sp. 

Otis, HelLx=Lampadion otis (Lightfoot) 

undata. Helix =Solaropsis undata (Lightfoot) 



NO. 3579 PORTLAND CATALOGUE — REHDER 39 

Helminthogiyptidae-Cepolinae 
vitellus, Helix =Polymita picta (Born) 

Helicidae 
alba, Helix =nomen dubium 

MOLLUSCA-BIVALVIA 

Arcidae 
fusca, Area = Area sp. 
nodulosa, Area=Anadara granosa (Linnaeus) 

Cucullaeidae 

labiata, Arca=Cucullaea labiata (Lightfoot) 

Mytilidae 

castaneus, Mytilus= Modiolus vagina Lamarck 
pietus, Mytilus=Perna picta (Born) 

Pinnidae 

rigida, Pinna=Atrina rigida (Lightfoot) 

Isognomonidae 

Isognoma=Isognomon [Lightfoot] 

Isognomon [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 (Liiniaeus) 
impressum, Cardium = Corculum impressum (Lightfoot) 

humanum Roding 
protrusum, Cardium=Corculum impressum (Lightfoot) 
robustum, Cardium = Dinocardium robustum (Lightfoot) 
spinosum, Cardium=Acanthocardia echinatum (Linnaeus) 



40 PROCEEDINGS OF THE NATIONAL MUSEUM vol 

Solenidae 
pallidus, Solen=Solen sp. 

Tellinidae 

marginalis, Telliiia=Tellma laevigata Linnaeus 

Garidae 

cruenta, Tellina=Sanguinolaria cruenta (Lightfoot) 

operculata (Gnielin) 
rostratus, Solen=Hiatula diphos (Linnaeus) 
Soletellina diphos (L.) 

Solecurtidae 

antiquatus, Solen=Soleciu-tus (Azorinus) chamasolen (Da Costa) 
plebeius, Solen=Tagelus plebeius (Lightfoot) 

Veneridae 

nimbosa, Venus =Macrocallista nimbosa (Lightfoot) 
punctata, Venus = Mere trix 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, Argon auta= Argon auta 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) 

pollicipes (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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1965. The Reverend John Lightfoot, Daniel Solander and tlie Portland 
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NO. 3579 PORTLAND CATALOGUE — REHDER 45 

KoHN, Alan 

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46 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

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1929-35. Handbuch der systematischen Weichtierkunde. Jena, 2 vols, 
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^^- ^"* PORTLAND CATALOGUE — REHDER 47 

TOMLIN, J. R. LeB. 

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1956. Coloured illustrations of sea shore animals of Japan. Osaka, xvii+ 
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pp. 106-107. 



Index 



Acanthina monodon, 8, 24 
Acanthocardia echinatum, 20 
Adelomelon ancilla, 15, 24 
Anadara granosa, 18 
Aranea gracilis, 34 

triremis, 34 
Area concamera, 32 

concamerata, 32 

cucuUus, 32 

fusca, 10 

imbricata, 11 

labiata, 32 

nodulosa, 18 

tetragona, 11 
Argobuccinum pustulosuin, 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, 13, 25 

laciniatum, 20 

monoceros, 8 

monodon, 8, 24 

muricatum, 20 

prismaticum, 7 

pustulosum, 16 

taurinum, 25 

testudo, 17 
Bulimus bovinus, 9 
48 



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 



INDEX 



49 



Cookia sulcata, 21 
Corculum humanum, 26 

impressum, 26, 30 
CucuUaea auriculifera, 32 

labiata, 32 
Cymbiiim 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 
D Glabella auricularia, 25, 33 

callosa, 25 

rumphii, 25 
Dolichoschisma munita, 14 

Echinus anemonoides, 7 

grandls, 34 

maximus, 34 
Elliptic 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 



Hydatina inflata, 28 
vesicaria, 23 
zonata, 27, 29 

Isognoma, 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 

poUicipes, 18 

virgata, 16 

vittata, 16 
Limax fibratus, 9 

undulatus, 8 
Llngula 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 

calcaratus, 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 

coraplanata, 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 

Placuna, 7, 12, 23, 24, 28, 30 

ephippium, 7, 12, 23, 28, 30 

placenta, 7, 12, 14 
Plagiobrissus grand is, 34 
Plejona muta, 8 
Pleurodonte otis, 9 
Podophora atrata, 7 
PoUicipes cornucopia, 18 
Polymita picta, 8 
Polyphyllia talpina, 33 
Pterynotus elongatus, 13 
Pyrula galeodes, 22 

hippocastanum, 22 

Rostellaria curvirostris, 5 

Sagda epistylium, 12 
Sanguinolaria, 27 

bertlni, 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 

pallidas, 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 



INDEX 



51 



Strophocheilus oblongus, 15, 28 

ovatus, 25, 27 

pudicus, 33 
Subninella undulata, 8, 30 

Tagelus plebeius, 11 
Tellina eruenta, 7, 13 

cruentae, 7 

laevigata, 24 

marginalis, 24 

operculata, 7 

sanguinolenta, 7 

semiplanata, 7 
Terobra 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 

sulcatus, 21 

tectum, 33 

tectus, 33 

virgineus, 18 
Trophon geversianus, 21 

laciniatus, 20 

plicatus, 20 
Turbinella, 15 

augulata, 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 



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, 32 

pepo, 16 

polyzonalis, 8 

ponderosa, 8, 35 

scafa, 6, 10, 23 

scapha, 6 

scolymus, 14 

spectabilis, 15 

tesselata, 17 

vexillum, 8 

virescens, 8, 23, 29 
Volutocorona nobilis, 6 

Xancus, 15 

angulatus, 14 

pyrum, 8 
Xenophora conchyliophora, 6, 30 



U.S. GOVERNMENT PRINTJNG 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 oj Crustacea 



Introduction 



The Pseudophilomedidae, new family, is represented in the oceans 
of the world by only one genus, Pseudophilomedes MuUer, 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 ^vriter recently received 17 specunens 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 joveolatus Muller, 1894, the type species of Pseudo- 
philomedes, were borrowed from the Zoological Museum, BerUn, and 
the original description by Muller (1894) is supplemented in this paper. 

Specimens identified by Brady and Norman (1896) as males of 

1 



2 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

Pseudophilomedes Joveolatus M tiller (called Philomedes joveolata 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 Euphilomedes Poul- 
sen, 1962. They are identified as Euphilomedes asper (Miiller, 1894) 
and Euphilomedes sp. and are described and illustrated in this paper. 
A supplemental description is given of a syntype of Euphilomedes 
asper (Miiller) borrowed from the Zoological Museiun, Berlin. 

As a result of this study, the subfamily Philomedinae Miiller, 1912, 
has been raised to familial rank. 

I wish to thank Dr. John H. Day of the Beaufort Marine Laboratory, 
Duke Univrersity, 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 (Muller) by Brady and Norman 
(1896), and Dr. H. E. Gruner for permission to study syntypes of 
Pseudophilomedes Joveolatus Miiller and Philomedes asper Miiller in the 
collection of the Zoological Museum, Berhn, I wish also to thank 
Miss Carohne 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, Miiller (1912) established the subfamily 
Philomedinae in the Cypridinidae. He referred to the Philomedinae 
the genera Philomedes Liljeborg, 1853, Pseudophilomedes, and Ruti- 
</ermaBrady 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, Poiilsen (19G2, 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 1st 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 revicAving the genus Pseudophilomedes I find that I concur 
with Poulsen; therefore, I have estabhshed 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 Miiller, 1894. Skogsberg 
(1920, p. 348) agreed with Miiller 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 Muller (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. Muller 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 MuUer 
established the genus Pseudophilomedes. It is unfortunate that 
Brady and Norman did not have more time to study Muller's paper 
before publishing their monograph. The fact that they wongly 
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 sufl&ciently 
different from Philomedes to warrant the establishment of a new 
genus and, then, on page 670 estabhsh 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 Javosa 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, 
pi. 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 Poidsen, 1962. Claw no. 1 
of the furca 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. 
Streptoleheris javosa is probably closely related to Pseudophilomedes 
and may belong in that genus. Streptoleheris 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). Streptoleheris rectirostris Brady and Norman, 
1896, the third species that has been referred to Streptoleheris, is also 
known only from the shell and Miiller (1912, pi. 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 Philomedmae 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) PhUomedidae Pseudophttomedidae 

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) s f 

6th limb, no. of endites 4 3_4 

6th limb, end joint, no. of bristles 9-45 7-9 
7th limb, terminal with comb opposed by 2 pegs (p) or 

2 opposing combs (c) p o 

Skogsberg (1920), Poulsen (1962, 1965), and Hartmann (1964) are 
correct in considering the families Rutidermatidae Brady and Nor- 
man, 1896, Cylindroleberidae Miiller, 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 SarsielUdae as done by Sylvester-Bradley (1961) seems no longer 
tenable. Thus, the famihes contained in the superfamily Cypridi- 
nacea are: Cypridinidae Baird, 1850; Philomedidae Miiller, 1912; 
Rutidermatidae Brady and Norman, 1896; Cylindroleberidae Miiller, 
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 elongate protopodite on maxilla ... 2 

2. Mandible of female with large pincers distally Rutidermatidae 

Mandible without large pincers 3 

3. Fifth limb of female with large teeth 4 

Fifth limb of female without large teeth Sarsiellidae 

4. Maxilla with only 2 stout endites Pseudophilomedidae 

Maxilla with 3 endites ^ 

5. Lamellar prolongation of selvage with fringe of hair .... Philomedidae 
Lamellar prolongation of selvage without fringe of hair .... Cypridinidae 

* Gills are common to all Cylindroleberidae but are rare in the Cypridinidae. 

Pseudophilomedidae, new family 

Philomedinae Muller, 1912, p. 24 (part). 

Type genus. — Pseudophilomedes MiiUer, 1894. 

Diagnosis (female; male unknown) .—SheU : 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 mth 1 dorsal 
bristle; 3rd joint with 1 ventral and 1 dorsal bristle; 4th joint with 
1 distodorsal and 2 distoventral bristles; 5th joint with 1 disto ventral 
bristle having short terminal filaments; 6th joint reduced, with a 
short distomedial bristle; 7th joint reduced, with 1 short a-bristle 
%\ath 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 mth 9 joints: 2ud to 
8th joints each with 1 long bristle; 9th joint with 2-3 bristles. 

Mandible: Coxale endite bifurcate at tip. Basale mth 4 ventral 
and 3 dorsal bristles. Exopodite elongate wdth 2 terminal bristles. 
Endopodite: 1st joint Avith 1-4 bristles; 2nd joint Avith 3-4 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. 

Furca: Thu"d claw more slender than 4th. Total nimiber of claws 
6-10; all well-developed claws separated from lamella by suture. 

Distribution. — Atlantic Ocean off Morocco, on Great Bahama 
Bank, B.W.L, on Continental Shelf off North Carolina and Georgia, 
U.S.A., Gulf of Naples (Mediterranean Sea). 

Ecology. — 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 



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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 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 

Figures 1-6; Plate 1 

Pseudophilomedes ferulana Kornicker, 1959, p. 235, figs. 46, lA-B, 2A-B; 56, 
A-D.— Darby, 1965, pp. 64-70, fig. 10, pis. 11, 12. 

Holotype. — USNM 113287; male; complete dry specimen. Spec- 
imens from Atlantic Shelf, 8 specimens: USNM 113020-113022. 

Material. — The nmnber 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, Avidest posterior to center; 
posterior process slightly below middle of valve; rostriun 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. Sh). 
Inner lamella broadest behind rostrum and in front of posterior 
projection; radial pore canals numerous (fig. 3/) ; selvage with lamellar 
prolongation and fringing hairs (fig. 3^) ; inner lamella behind rostrum 
with row of 4 bristles ^nth short marginal hairs (fig. 3d) ; inner lamella 
in front of posterior process \vith row of 6 hirsute setae (fig. 3e) ; inner 



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 (pi. li). 

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 


EST 30 


1 


1.66 


1. 16 


1. 00 


eggs in brood pouch 


It 


2 


1.30 


0.81 




late instar 


It 


3 


1. 54 


1. 00 




eggs in body 


(( 


4 


1.52 


1.00 




eggs in body 


(( 


5 


1.54 


0.96 




no eggs 


GOS 1857 


1 


1.73 


1. 15 




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 ^vith forked tip and 1 distal 
and 2 proximal filaments; 1st to 4th joints with rows of slender spines 
on medial surface. 

Second antenna (figs. 4e,/) : Endopodite (fig. 4/) single jointed, vath. 
long hirsute bristle and 2 short bare bristles. Exopodite (fig. 4e) with 
9 joints: 1st joint slightly longer than total length of follo^g 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 
jomts with sparse natatory hairs and stout marginal spines near 
middle; 9th joint ^vith medium spinous bristle and 2 long bristles 
with natatory hairs; distal margins of 2nd to 8th joints ^^ith 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. 4:g~i): Coxale endite (fig. 4A) 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 
2"d antenna 



-nnaxilla 



5th limb 
- furca 



^ adductor muscles 
6»hlimb 



epipodial appendage 
(on 5^ linnb) 



7th limb 



Figure 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.) 



OSTRACOD FAMILIES — KORNICKER 



11 



odite: 1st joint with 1 short and 2 long ventral bristles distally; 
2nd joint with 1 dorsal bristle proximal to middle, 3 dorsal bristles 
near middle, and 3 short ventral bristles distally; end joint (fig. 4i) 
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, 

-^i 




Figure 2.-PseudophUomedes ferulanus: lateral view, left maxilla, Sth 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 spmes and 
2 spinous bristles. Distal edge of basale .^ath 1 spmous medial 
bristle and 2 on mner margin adjacent to low node mth 2 bnstles 
at base of coxale endite, shorter of latter bristles bare, other ^nth 
several marginal spines. Exopodite consisting of 3 lateral spmous 
bristles distaUy on basale (figs. 1, 5a). Endopodite: l^J^-t ^'^^^^ 
vvith 2 spinous bristles, 1 near middle and 1 termmally; 2nd jomt 



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-i): First joint with 2 large teeth: proximal 
tooth trident with 3 bristles (fig. 5/); distal tooth bifid (fig. 5h). 
Second joint prolonged distally forming long fangiike tooth followed 
along the inner margin by a single tooth and 2 trident teeth (fig. 5e) ; 
posterior siu-face with 2 stout spinous bristles and a minute bristle 
(fig. 5i) ; inner edge of joint with spinous bristle (fig. 6i). (It was not 
possible to discern mth 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. 5i). 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. Qb-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 mth fewer bristles) , each 
bristle with 1 to 5 bells (fig. Qd) and short marginal spines distally. 
Distribution of bristles on seventh limbs of 4 specimens from station 
BST30 areas follows: 

limb A limb B* 

specimen no. 1 

terminal bristles 6 6 

lateral bristles 8 7 

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. 

Furca (figs. 6e, J-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. 35S0 OSTRACOD FAMILIES — KORNICKER 13 

spines distally on anterior surface ; claws 2-8 witii spines along medial 
and lateral margins; lamella, in area of distal and proximal claws, 
with hairs. 

Eyes: Median eye (fig. Qj) large, pigmented. Lateral eyes (fig. 6<j) 
small, black in preserved specimens. 

Frontal organ (figs. Qj, k) : Elongate, proximal part divided into 
about 8 short segments, distal part (fig. Qk) tapered, with short spines 
near terminus. 

Eggs (fig. 6^) : 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 MuUer, 1894, pp. 211-212, pi. 3, figs. 34, 35, 45-49, 
51, 53, 54; pi. 4, figs. 127. [Not Philomedes foveolata Brady and Norman, 
1896, pp. 659-661, pi. 56, figs. 4, 5.] 

Syntypes. — Zoological Museum of Berlin, Division of Crustacea, 
catalog no. 9154. 

Material. — 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 Joveolata G. W. Midler, 
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 lameUa broadest behind rostrum and in front of posterior pro- 
jection; selvage with lameUar prolongation and fringing hairs unbroken 
around antennal sinus; inner lamella behind rostriun with row of 4 
bristles with short marginal hairs (fig. 76); inner lameUa m Ifont of 
posterior process with 1 (or 2 closely spaced?) hirsute seta (fig. 7c); 
inner lameUa below antennal sinus with 1 short bristle; outer surface 
of each valve with scattered surface hairs. ^ 

MuUer (1894, pi. 3, fig. 7) illustrates 4 bristles on the mner lamella 
behind the rostrum but does not show them to be hu-sute. 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 1st antenna illustrated by Miiller (1894, 
pi. 3, 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, pi. 3, fig. 35) is similar to 
the specimen described in this paper. Miiller apparently over- 
looked the row of short spines along the distal margin of joints 2-8 
of the exopodite. 

Mandible (figs. 7j, k) : Coxale endite (fig. 7j) bifiu-cate 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. 7A:) 
with 3 bristles and 2 claws, 1 of latter very long, other about 3rd 
length of long claw. Terminal end of 1st endopodite joint with 
spines in row. 

The mandible of the species illustrated by Miiller (1894, pi. 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 7n): 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 margui. Exopodite consisting 
of 3 lateral bristles located distally on basale. Endopodite: 1st joint 
with 1 subterminal and 1 terminal bristle; 2nd jomt with 1 short and 
2 long bristles and a slender process with annulations distally. 

Miiller (1894, pi. 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 Iknb (figs. 8a-e) : Fu-st jomt with 2 large teeth ; proxunal tooth 
trident with 2 or 3 bristles, forward tooth spmous 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 leadmg margin. Second joint prolonged distally formmg 
long fanglike tooth followed along the mner margm by a smgle tooth 
and 2 trident teeth; posterior surface with 1 long and 1 short bristle. 
Thkd joint with 2 spinous bristles near base of 4th joint. Fourth 
joint hhsute with 4 or 5 bristles. Inner margin of protopodite with 
single endite with 1 or 2 minute bristles. _ 

The taxonomicaUy unportant inner margin of the 2nd joint with its 
3 teeth is not shown in the view of this appendage illustrated by 
MuUer (1894, pi. 3, figs. 43, 46). The fanglike tooth d^tally on the 
2nd joint is much longer on the specimen I exammed than won d be 
suspected from examination of MuUer's Ulustrations. Possibly short- 
ness of that tooth in Miiller's illustration is the result of foreshortenmg, 
which results when an appendage is mounted obliquely on the slide. 



16 PROCEEDINGS OF THE NATIONAL MUSEUM vol. 121 

The 2 teeth of the 1st jomt of the specunen illustrated by Miiller and 
the one I examined are quite similar. Miiller 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 exammed. This may have been 
overlooked by Miiller, or possibly the number of bristles on the 4th 
joint varies slightly within the species. 

Sixth lunb (fig. 8/) : Fu'st 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, pi. 3, fig. 54) is quite 
sunilar 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. 8^-^): Each lameUa 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 proxknally on lateral margm, slender spines distaUy 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, pi. 3, figs. 34, 53) figures a furca with the 3rd claw 
almost equal m 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 Muller. In this regard, it is 
interesting that Brady and Norman (1896, p. 672) state that the larger 
furcal claws of Pseudophilomedes infatus are ciliated on the edge and 
in the figure of this appendage (1896, pi. 59, fig. 10) show claws 3 and 
8-10 without marginal spines. In view of MuUer'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 Muller, 1912, p. 24 (part). 

Type genus. — Philomedes Liljeborg, 1853. Gender: Masculine. 
Diagnosis. — SheU: Shell variable in outline, but generally elongate 



NO. 3580 OSTRACOD FAMILIES — KORNICKER 17 

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 disto ventral 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 ventraUy 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); 8th 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 jomt 
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 distaUy; 
joints usually with basal spmes; 2nd to 8th 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 ventraUy at least 1, but usuaUy 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 Ihnb: Exopodite: 1st jomt of female Avith 2 or more strong 
teeth; 2nd joint of female forming large squarish sclerotized process 
having large inward projecting tooth distaUy and an inner margin 
either smooth or with small nodes or 2 large teeth; 1st and 2nd ]omts 
of male without stout teeth; 3rd jomt with 3 bristles on mner 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). , • i o ^ 

Sixth Ihnb: Endites 4 in number; end jomt 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: Wleoschisma Brady, 1880: ITetragonodon Brady and 
Norman, 1896; Scleroconcha Skogsberg, 1920; Euphilomedes Poulsen, 
1962; Paraphilomedes Poulsen, 1962. 

Euphilomedes Poulsen 

Type species. — Euphilomedes nodosus Poulsen, 1962, by subsequent 
designation, Komicker (in press). Gender: masculine. 

Euphilomedes asper, new combination 

Figures 9, 10, lla-d, 12 

Philomedes aspera Miiller, 1894, pp. 210-211, pi. 3, figs. 3, 17, 21; pi. 8, fig. 1. 
Philomedes foveolata. — Brady and Norman, 1896, pp. 659-661, pi. 56, figs. 4, 5. 
[Not Pseudophilomedes foveolata Miiller, 1894.] 

Syntypes. — Zoological Museum of Berlin, Division of Crustacea, 
catalog no. 9152. 

Material. — 1. A glass slide in the collection of the British Museum 
(Natural History) labeled "Philomedes foveolata Bra & Norcf" 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. 
36962-964." The specimens in this vial are designated in this paper 



NO. 3580 OSTRACOD FAMILIES — KORNICKER 19 

as specimens A and B. Specimen A was 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. 
Miiller, Kat. Nr 9152, Fundort Napoli." The small vial contains the 
label, "9152." These specunens are syntypes; Miiller (1894) did not 
designate a holotype for the species. The whole spechnen 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-i; I2a-f): Lateral 
outline oval, elongate with greatest height near middle, prominent 
rostrmn and broad rostral incisure (figs. 9a, \2a-d)', surface of valves 
with numerous large oval pits (fig. 96). 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, i; 
12g). Inner lamella broadening at postero ventral 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 o