ARTHUR G. HUM

and JAN H. STOC

A Revision of the

Family Lichomolgidae

Kossmann, 1877,

^m Cyclopoid Copepods

f Mainly Associated with

Marine Invertebrates

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY NUMBER 127
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SMITHSONIAN CONTRIBUTIONS TO

ZOOLOGY

NUMBER 127

Arthur G. Humes A Revision of the

and Jan H. Stock -pi • i T • i_ l • J

Family Lichomolgidae

Kossmann, 1877,

Gyclopoid Gopepods

Mainly Associated with

Marine Invertebrates

SMITHSONIAN INSTITUTION PRESS

CITY OF WASHINGTON
ABSTRACT

Humes, Arthur G., and Jan H. Stock. A Revision of the Family Lichomolgidae

Kossmann, 1877, Cyclopoid Copepods Mainly Associated with Marine Invertebrates. Smithsonian Contributions to Zoology, number 127, 368 pages, 190

figures, 1973.—The poecilostome family Lichomolgidae has been divided into

five families (Humes and Stock, 1972): Sabelliphilidae Gurney, 1927; Lichomolgidae Kossmann, 1877; Urocopiidae Humes and Stock, 1972; Pseudanthessiidae

Humes and Stock, 1972; and Rhynchomolgidae Humes and Stock, 1972. These

families form the superfamily Lichomolgoidea Humes and Stock, 1972. In

the present work 76 genera are characterized, including 324 species (37 new).

Previously known species are recorded from 73 new hosts. In addition to illustrations of the new taxa, selected figures of each genus not represented by new

species have been reproduced. Keys are provided for the families, genera, and

species where possible. For each species the geographical distribution and hosts,

where known, are indicated.

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

in the Institution's annual report, Smithsonian Year.

Library of Congress Cataloging in Publication Data

Humes, Arthur Grover, 1916-

A revision of the family Lichomolgidae Kossmann, 1877.

(Smithsonian contributions to zoology, no. 127)

Bibliography: p.

1. Lichomolgidae. 2. Copepoda. I. Stock, Jan H., joint author. II. Title. III. Series: Smithsonian Institution. Smithsonian contributions to zoology, no. 127.

QL1.S54 no. 127 [QL444.C7] 595'M 75-651

For lale by the Superintendent of Documents, U.S. Government Priming Office

Washington, D.C. 20402 - Price $4-30 (paper cover)
Contents

Page

Introduction 1

Acknowledgments 2

Definition of Terms and Scope of the Work 3

Comparative External Anatomy of the Five Families of Iichomolgoidea 3

Host Specificity 4

Phylogenetic Relationships of the Five Families of Lichomolgoidea 5

Sup, i family Lichomolgoidea Humes and Stock, 1972 6

Family Sabclliphilidae Gurney, 1927 7

Calypsarion Humes and Ho, 1969 8

Calypsina Humes and Stock, 1972 9

Caribulus Humes and Stock, 1972 10

Diogenella Stock, 1968 12

Diogenidium Edwards, 1891 13

Henicoxiphium Ulg and Humes, 1971 14

Herrmannella Canu, 1891 15

Lecanurius Kossmann, 1877 40

Liehomolgidium Kossmann. 1877 41

Lichothuria Stock, 1968 44

ModioUcola Aurivillius, 1882 47

Myxomolgus Humes and Stock, 1972 61

Paranthessius Claus, 1889 66

Sabelliphilus M. Sars, 1862 66

Scambicornus Heegaard, 1944 68

Serpuliphilus Humes and Stock, 1972 73

Synapticola Voigt, 1892 80

Thamnomolgus Humes, 1969 81

Sabelliphilidae Insufficiently Described or of Uncertain Position 81

Family Lichomolgidae Kossmann, 1877 82

Acaenomolgus Humes and Stock, 1972 85

Acanthomolgus Humes and Stock, 1972 87

Amarda Humes and Stock, 1972 Ill

Anchimolgus Humes and Stock, 1972 119

Andrianellus Humes and Stock, 1972 121

Anisomolgus Humes and Stock, 1972 126

Ascetomolgus Humes and Stock, 1972 127

Ascidioxynus Humes and Stock, 1972 133

Aspidomolgus Humes, 1969 146

Astericola Rosoll, 1889 148

Colobomolgus Humes and Stock, 1972 150

Contomolgus Humes and Stock, 1972 154

Debruma Humes and Stock, 1972 161

Doridicola Leydig, 1853 165

Epimolgus Bocquet and Stock, 1956 180

Gelastomolgus Humes, 1968 181

Haplomolgus Humes and Ho, 1968 182

Indomolgus Humes and Ho, 1966 183

Kelleria Gurney, 1927 185

Lichomolgella G.O. Sars, 1918 187

Lichomolgides Gotto, 1954 188

Lichomolgus Thorell, 1859 189

Macrochiron Brady, 1872 194

Meringomolgus Humes and Stock, 1972 204

iii
IV SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

Page

Supt'i family Lichomolgoidea Humes and Stock, 1972—Continued

Family Lichomolgidae Kossmann, 1877—Continued

Metaxymolgus Humes and Stock, 1972 220

Monomolgus Humes and Frost, 1964 248

Nasamotgus Sewell, 1949 250

Octopicola Humes, 1957 251

Odontomolgus Humes and Stock, 1972 252

Panjakus Humes and Stock, 1972 255

Paradoridicola Humes and Stock, 1972 264

Paramacrochiron Sewell, 1949 271

Paramolgus Humes and Stock, 1972 272

Paraphiloconcha Yamaguti, 19S6 J75

Paredromolgus Humes and Stock, 1972 275

Pennatulicola Humes and Stock, 1972 276

Philoconcha Yamaguti, 1936 278

Plesiomolgus Humes and Stock, 1972 280

Prionomolgus Humes and Ho, 1968 280

Pseudomacrochiron Reddiah, 1969 281

Rakotoa Humes and Stock, 1972 283

Ravahina Humes and Ho, 1968 289

Schedomolgus Humes and Stock, 1972 290

Sewellochiron Humes, 1969 291

Spaniomolgus Humes and Stock, 1972 292

Stellicola Kossmann, 1877 293

Synstellicola Humes and Stock, 1972 301

Telcstacicola Humes and Stock, 1972 304

Xenomolgus Humes and Stock, 1972 309

Zamolgus Humes and Stock, 1972 314

Zygomolgus Humes and Stock, 1972 325

Lichomolgidae Insufficiently Described or of Uncertain Position 327

Copepods Placed by Various Authors in the Old Family Lichomolgidae

but Belonging Elsewhere 328

Family Urocopiidae Humes and Stock, 1972 329

Urocopia G.O. Sars, 1917 329

Family Pseudanthessiidae Humes and Stock, 1972 330

Heteranthessius T. Scott, 1904 330

Kombia Humes, 1962 331

Meomicola Stock, Humes, and Gooding, 1963 334

Pseudanthessius Claus, 1889 334

Temnomolgus Humes and Ho, 1966 341

Pseudanthessiidae Insufficiently Described or of Uncertain Position 343

Family Rhynchomolgidae Humes and Stock, 1972 343

Rhynchomolgus Humes and Ho, 1967 343

Previously Known Copepods Here Recorded from New Hosts 344

Literature Cited 345

Index to Generic and Specific Names of the Lichomolgoidea 360

Index to Host Genera 365
NUMBER 127

NEW SPECIES

Family Sabelliphilidae Family Lichomolgidae (continued)

Herrmannella bullata Doridicola ptilosarci

Myxomolgus proximus Macrochiron echinicolum

Serpuliphilus tenax Meringomolgus facetus

Serpuliphilus duplus Meringomolgus devotus

Family Lichomolgidae Meringomolgus hamatus

Acanthomolgus astrictus Metaxymolgus antheliae

Acanthomolgus bilobipes Metaxymolgus chlamydis

Acanthomolgus hales Metaxymolgus cinctus

Acanthomolgus plantei Metaxymolgus claudus

Arnarda cultrata Metaxymolgus inflatiseta

Amarda compta Panjakus hydnophorae

Andrianellus exsertidens Panjakus platygyrae

Ascetomolgus plicatus Paradoridicola sinulariae

Ascidioxynus floridanus Rakotoa proteus

Ascidioxynus bermudensis Stellicola illgi

Cohbomolgus laboutei Telestacicola angoti

Contomolgus lokobeensis Xenomolgus varius

Debruma clavelinae Zamolgus tridens

Doridicola astrophyticus Zamolgus acanthodes

SPECIES REDE-SCRIBED

Family Sabelliphilidae Family Sabelliphilidae (continued)

Herrmannella columbiae (Thompson, 1897) Modiolicola inermis Canu, 1892

Herrmannella panopeae (Illg. 1949) Modiolicola maximus (Thompson, 1893)

Herrmannella saxidomi (Illg. 1949) Myxomolgus myxicolae (Bocquet and Stock, 1958)

Herrmannella tivelae (Illg, 1949) Family Lichomolgidae

Lichomolgidium sardum Kossmann, 1877 Ascidioxynus jamaicensis (C. B. Wilson, 1921)

Lichomolgidium cynthiae (Brian, 1924) Macrochiron sargassi G. O. Sars, 1916

Modiolicola insignis Aurivillius, 1882 Family Pseudanthessiidae

Modiolicola gracilis C. B. Wilson, 1935 Kombia angulata Humes, 1962

Arthur G. Humes

and Jan H. Stock

A Revision of the

Family Lichomolgidae

Kossmano, 1877,

Cyclopoid Copepods

Mainly Associated with

Marine Invertebrates

Introduction

A variety of genera of poecilostome copepods has

been ascribed to the family Lichomolgidae since

its establishment by Kossmann in 1877. A few of

these genera, such as Anthessius Delia Valle, 1880

(=Pseudomolgus G. O. Sars, 1916), Synaptiphilus

Canu and Cuenot, 1892, Cholidya Farran, 1914,

Myicola Wright, 1885, and Rhinomolgus G. O.

Sars, 1918, have been removed from the Lichomolgidae by later authors because of their obvious

nonlichomolgid characters.

In recent years our knowledge of these copepods

has expanded considerably. From 1853 (the year

of establishment of the first lichomolgid genus,

Doridicola) to 1899, there were 13 genera described.

From 1900 to 1947 only 7 new genera were proposed. From 1948 to 1971 24 new genera were

created, and we recently added 32 new genera

(Humes and Stock, 1972). In this revision 76 genera are recognized. These contain 324 species,

37 of which are new.

Lichomolgid copepods, although well known as

common associates of marine invertebrates in

northern Europe and the Mediterranean Sea, appear to be particularly abundant on a wide variety

Arthur G. Humes, Boston University Marine Program, Marine Biological Laboratory, Woods Hole, Massachusetts,

02543. Jan H. Stock, Zodlogisch Museum, Plantage Middenloan 53, Amsterdam, The Netherlands.

of invertebrate hosts in tropical areas. Intensive

collections in warm regions have thus far been few

in number. Thompson and Scott (1903) recorded

283 species of copepods of all kinds from the Ceylon Pearl Banks (Herdman, 1904, p. 57), among

them 19 lichomolgids (including 16 new species).

Humes (1970c) and his coworkers reported 191

copepods (mostly cyclopoids) associated with marine invertebrates in the vicinity of Nosy B6,

Madagascar. Of these more than 100 were lichomolgids, the majority of them new. Other reports

by Stock (1967a, 1967b, 1968) in the Red Sea area

and by Stock, Humes, and Gooding (1963a) in the

West Indies suggest the richness of this tropical

fauna. More than seventy years ago I. C. Thompson (1900) was well aware of the diversity and

abundance of copepod associates when he wrote

(p. 140) ". . . it is evident that Sponges, Coelenterates, Echinoderms, Worms, Molluscs, Nudibranchs,

and Ascidians will all richly reward careful search;

for judging from past results, they are the hosts of

many highly organized species of Copepoda, each

specially adapted to its particular host and that

alone."

Various proposals have been made for suprageneric division of the Lichomolgidae. Kossmann

(1877) divided the family into the group Lichomolgini, characterized by a lack of deformation in

the female, and including Sabelliphilus M. Sars,

1862, Lichomolgus Thorell, 1859, and his new gen1
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

era Stellicola, Lichomolgidium, Lecanurius, and

Boholia, and the group Ergasilini, with the thorax

of the female deformed by the ovaries, and including his new genus Paclabius. Such a division seems

artificial, since deformation of the body frequently

occurs in associated or parasitic copepods, especially

in the females, where it is the expression of parasitic adaptation rather than a fundamental morphological feature.

Gurney (1927) divided the Lichomolgidae into

two subfamilies, the Sabelliphilinae, with the endopod of leg 4 composed of three segments, and

the Lichomolginae, with that endopod 2-segmented, 1-segmented, or absent. Sewell (1949)

adopted Gurney's division of the family. That division appears to reflect two evolutionary branches,

and it has served as the starting point for our revision.

In view of the greatly increased information

which we now have, we feel that the copepods included in the broad family Lichomolgidae show

more than two evolutionary lines, and that these

relationships can best be expressed by a division

of the old family Lichomolgidae into five families.

One of these, the family Sabelliphilidae, corresponds largely to Gurney's Sabelliphilinae, but the

definition has been expanded to include not only

those with a 3-segmented endopod in leg 4 but

also those with one or more of the endopods of

legs 1-4 being 2-segmented in a series from anterior to posterior. The family Lichomolgidae is

retained, but in a restricted sense, for those with

the endopod of leg 4 having fewer than three segments or being absent (with the endopods of legs

1-3 not reduced, except in leg 3 in Ravahina and

in legs 1-3 in Amarda), and with a free segment in

leg 5. The family Pseudanthessiidae shows a strong

reduction in the endopod of leg 4 (1-segmented or

absent, with this reduction in certain genera being

carried forward into the more anterior legs) and

lacks a free segment in leg 5. The family Urocopiidae accomodates the genus Urocopia, with the endopod of leg 4 composed of three segments but with

leg 5 lacking a free segment. The family Rhynchomolgidae contains the genus Rhynchomolgus, with

legs 1-4 showing very strong reduction from posterior to anterior, with leg 5 in the female lacking a

free segment, and with strong sexual dimorphism

both in body form and in the appendages (in the

male the mandible, maxilliped, leg 4, and leg 5 being absent).

Specimens that are deposited in the National

Museum of Natural History, Smithsonian Institution, Washington, D. C, are listed under the catalog numbers of the former United States National

Museum (USNM).

Acknowledgments

This revision could not have been prepared in its

present form without the help of many individuals

who have generously contributed information and

specimens.

We wish to thank particularly Dr. Ju-Shey Ho,

then at Boston University but now at California

State University, Long Beach, California, for his

substantial aid in the preparation of material. His

great patience and skill in the dissections and drawings of many species enlarged significantly the scope

of this revision.

We acknowledge with thanks the contributions

of the following:

Mr. G.H. Arnott, Monash University, Clayton. Victoria,

Australia, for information on Pseudantheuius tenuis in Australia.

Dr. F.M. Bayer, University of Miami, Miami, Florida, for

the identification of a coelenterate host of Macrochiron

sargassi.

Dr. Kenneth J. Boss, Museum of Comparative Zoology.

Harvard University, Cambridge, Massachusetts, for the currently used names of certain bivalve hosts of Herrmannella.

Dr. Thomas E. Bowman and Dr. Fenner A. Chace, Jr.,

Division of Crustacea, Smithsonian Institution, Washington.

D. C, for the loan of a paratype of Wilson's Modioticola

jamaicensis.

Mr. James F. Clark, Museum of Comparative Zoology,

Harvard University, for the identifications of several species

of Linckia, the hosts of Stellicola pollex in Hawaii.

Professor H. Engel and Dr. W. Schroevers, Amsterdam,

for the identifications of echinoid hosts of Macrochiron

echinicolum.

Dr. A. Fosshagen, Biologisk Stasjon, Espegrend, Blomsterdalen, Norway, for allowing us to include his specimens of

Meomicola amplectans from the Bahamas.

Dr. Richard U. Gooding, formerly at the University of

Singapore, for aid with the fieldwork in the West Indies

(1959) and Bermuda (1962).

Dr. Paul L. Illg, University of Washington, Seattle.

Washington, for sending us specimens of two new copepods

(Herrmannella bullata and Doridicola ptilosard) with permission to include them in this work, and also for providing

us with material of several West Coast species of Herrmannella.
NUMBER 127

M. Pierre Laboute, diver at the Centre ORSTOM de

Nosy Be, Nosy Be, Madagascar, for accompanying the first

author on SCUBA dives during June, July, and August

1967, thus making it possible to collect copepods from hosts

in depths down to 50 meters.

Dr. R. H. Millar, Marine Station, Millport, Scotland, for

the names of tunicate hosts of Macrochiron sargassi and

Ascidioxynus jamaicensis.

Dr. Marian H. Pettibone, Smithsonian Institution, Washington, D. C, for the identifications of certain West Indian

polychaetes, the hosts of Serpuliphilus tenax.

Dr. Michel Pichon, Station Marine d'Endoume, Marseille,

France, for the determination of the madreporarian host of

Kombia angulata.

Dr. Raphael Plante, Centre ORSTOM de Nosy Be, Nofy

Be, Madagascar for the specimens of the gorgonian Solenocaulon tortuosum, the host of Acanthomolgus hales.

Dr. J. Verseveldt, Zwolle, The Netherlands, for the identification of many alcyonacean hosts.

Dr. W. Vcrvoort, Rijksmuscum van Natuurlijke Historic,

Leiden, The Netherlands, for the identification of certain

gorgonian hosts.

Dr. John W. Wells for the identification of madreporarians from Eniwetok Atoll.

Dr. E. Wesenberg-Lund for the identification of polychaete hosts of Serpuliphilus from the West Indies.

Much of the fieldwork and laboratory study forming the basis for this revision could not have been

carried out without the financial assistance from

several grants (G-2899, G-4830, G-15911, GB1809, and GB-5838) made to the first author by

the National Science Foundation of the United

States. The actual preparation of the manuscript

has been aided by a current grant (GB-8381X)

from the National Science Foundation.

Definition of Terms and Scope of the Work

The brief generic descriptions that follow include

only those characters considered to be especially

diagnostic; thus, they serve to distinguish the genera one from another. The descriptive statements

apply to both sexes unless otherwise specified.

For descriptive purposes three terms have been

used to refer to the aspect of the body: (1) cyclopiform, where the aspect is that of a generalized unmodified cyclopoid (as in Paranthessius and many

Lichomolgus); (2) modified, where the form is

basically cyclopoid but is elongated and slender

(as in Octopicola), unusually broadened (as in

Aspidomolgus), or otherwise departs from the generalized form (as in Gelastomolgus); and (3) transformed, where the body, especially in the female,

is swollen, vermiform, or otherwise changed, often

with loss of external segmentation (as in Rhynchomolgus).

Although there are intergrading species for

which the application of these terms might be

difficult, the three categories are useful in a general way.

The following abbreviations are used: Alt first

antenna; A2, second antenna; L, labrura; MD, mandible, MX2, second maxilla; MXPD, raaxilliped;

Pi.5, legs 1-5; EXP, exopod; and ENP, endopod.

In the spinal and setal formulas the Roman numerals indicate spines, the Arabic numerals setae.

In the list of species under each genus the hosts

with which the copepods are associated are given in

all cases where they are known. The geographical

distribution of the species has been in some cases

noted in a general way in the interest of brevity.

For each genus where new forms are not herein

described, selected figures of a known species have

been reproduced in order to illustrate certain characters and to facilitate generic determination.

Every effort has been made to include for each

species all significant synonyms, all references containing new descriptive or interpretive material,

and all references to hosts and geographical localities. In certain cases where species names have

been used only as citations from previously recorded authors the names have not been entered in

order to shorten the lists.

Comparative External Anatomy of the

Five Families of Lichomolgoidea

In the majority of genera the body retains the

general cyclopiform fades. In certain genera, however, the body is modified or transformed. The

Sabelliphilidae and Lichomolgidae contain several

modified genera and a few which are moderately

transformed (Synapticola, Amarda, Paraphiloconcha, Ravahina). In certain cases, only the female

is modified or transformed, the male being cyclopi*

form (Lichomolgides, Epimolgus). Urocopia, the

only known genus of the Urocopiidae, is cyclopiform. In the Pseudanthessiidae there is a strong

tendency toward modification or transformation,

with only the genus Pseudanthessius having a cyclopiform facies. Rhynchomolgus, the only known

genus of the Rhynchomolgidae, is highly transformed.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Most species have a characteristic size range in

both sexes. A few, however, exist in two size forms

(Caribulus sculptus, Meomicola amplectans).

Generally, females are larger than males, though

in a few cases males are larger (Indomolgus brevisetosus, Rhynchomolgus corallophilus, Temnomolgus eurynotw).

Sexual dimorphism is most usual in the first and

second antennae, the maxillipeds, leg 1, leg 5, and

the genital segment. Sexual differences, however,

may appear in other structures; for example, in

the rostrum (Acanthomolgus varirostratus, Paramolgus insectus), in the segmentation of the first

and second antennae (Rhynchomolgus corallophilus), in the second maxilla (Meringomolgus), and

in the caudal ramus (Rhynchomolgus corallophilus). Suckers occur on the second antenna of the

male in a few species (Aspidomolgus stoichactinus,

Herrmannella poculifera). One species, Stellicola

femineus, is unique in that the male has a maxilliped much like that of the female, without the

long claw characteristic of the male.

Several pairs of species, each pair associated with

one host genus, are remarkably similar in their external anatomy and may be regarded as sibling

species (Paramolgus politus and P. simulans, Paramolgus constrictus and P. insectus, and Plesiomolgus conjunctus and P. organicus).

The genital segment varies greatly in form in

the female, less so in the male. In Calypsarion

there is a remarkable modification of this segment

in the female, where there is a ventral keel. The

genital segment in the male of this genus lacks a

keel, but a keel is present on the first postgenital

segment. In Schedomolgus lobophorus, while the

genital segment in both sexes is ordinary, there is

a ventral lobe on the first postgenital segment in

the female and on the second postgenital segment

in the male.

The number of postgenital segments is, in almost all genera, three in the female and four in

the male. In Synstellicola and in one species of

Pseudanthessius the number is two in the female

and three in the male (P. deficient). (This is

probably true also for P. dubius, where there are

two postgenital segments in the female, but the

male is unknown.)

The caudal ramus, though variable in size and

proportions, has six setae. The setae, especially

the two median terminal ones, are usually very

long, but may be very short as in Ravahina tumida. In Diogenella the two median terminal setae

are vestigial.

One species (Thamnomolgus robustus) is unique

in having a pair of ocelliform sclerotized areas on

the ventral surface of the cephalosome in the male

posterior to the bases of the maxillipeds.

The usual pattern (referred to below as lichomolgidiform) for the armature of legs 1-4 is as

follows (the Roman numerals indicating spines,

the Arabic numerals setae):

Pj coxa 0-1 basis 1-0 exp 1-0; I-l

enp 0-1; 0-1

Pt coxa 0-1 basis 1-0 exp 1-0; I-l

enp 0-1; 0-2

Pa coxa 0-1 basis 1-0 exp 1-0; I-l

enp 0-1; 0-2

P« coxa 0-1 basis 1-0 exp 10; I-l

111,1.4

13

UI3

11,13 or

IIU3

enp variable

Deviations from this arrangement are described in

the text below.

Host Specificity

Approximately three-fourths of the genera in the

five families contain at present relatively few species (less than five). Only 18 genera have five or

more species. The ten largest genera are Pseudanthessius (34 species), Metaxymolgus (23), Scambicornus (21), Lichomolgus (17), Herrmannella

(16), Doridicola (14), Acanthomolgus (13), Stellicola (10), Kelleria (9), and Macrochiron (9). Thirty

genera are monotypic. Observations on the relative

sizes of the genera have little significance, however,

since it is apparent that our present knowledge of

these families is very limited. Large numbers of

new forms remain to be described. In the collections of the present authors are numerous new

species that await study.

In the Sabelliphilidae the 18 genera are associated with particular groups of hosts, with no adventitious associations with other host groups.

(The only exception is Scambicornus armoricanus

Bocquet, Stock, and Kleeton, 1963, from a polychaete rather than from a holothurian like other

members of the genus. As mentioned in the text

below, however, this species probably belongs to a

new genus rather than to Scambicornus.) The associations of copepod genera and host groups in

this family are as follows:
NUMBER 127

with Antipatharia: Thamnomolgus

with Actiniaria: Paranthessius

with Polychacta: Myzomolgus, Sabelliphilus, Serpuliphilus

with Bivalvia: Herrmannella, Modiolicola

with Holothuroidea: Calypsarion, Calypsina, Caribulus, Diogenella, Diogenidium, Lecanurius, Lichothuria, Scambicornus (except S. armoricanus), Synapticola

with Ascidiacea: Henicoxiphium, IAchomolgidium

In the Lichomolgidae 42 genera are associated

with definite host groups, as follows:

with Scyphozoa: Paramacrochiron, Pseudomacrochiron, Sewellochiron

with Actiniaria: Aspidomolgus

with Octocorallia: Acanthomolgus, Anisomolgus, Ascetomolgus, Colobomolgus, Contomolgus, Meringomolgus, Paradoridicola, Paredroniolgus, Pennatulicola, Plesiomolgus,

Telestacicola, Zamolgus

with Madreporaria: Amarda, Anchimolgus, Andrianellus,

Haplomolgus, Monornolgus, Odontomolgus, Panjakus,

Prionomolgus, Rakotoa, Ravahina, Schedomolgus, Spaniomolgus, Xenomolgus

with Polychaeta: Acaenomolgus, Nasomolgus

with Bivalvia: Gelastotnolgus, Paraphiloconcha, Philoconcha

with Gastropoda: Epimolgus

with Cephalopoda: Octopicola

with Asteroidea: Astericola, Stellicola, Symtellicola

with Ascidiacea: Ascidioxynus, Debruma, Lichomolgides

On the other hand, several genera in the Lichomolgidae contain species associated with more than

one host group or are free-living:

with Mollusca and Ascidiacea: Lichomolgus

with Algae, Hydrozoa, Echinoidea, Ascidiacea: Macrochiron

with Alcyonacea, Zoanthidea, Actiniaria, Opisthobranchia:

Metaxymolgus

with Ascidiacea, Algae, Holothuroidea: Zygomolgus

with Algae, free-living; Lichomolgella

with Crinoidea, free-living, in burrows: Kelleria

with Zoanthidea, Actiniaria: Indomolgus

with Octocorallia, Actiniaria: Paramolgus

with Octocorallia, Actiniaria, Nudibranchia, Crinoidea: Doridicola.

Except for the genus Pseudanthessius the genera

of the Pseudanthessiidae are restricted in their host

groups as follows:

with Zoanthidea: Temnomolgus

with Madreporaria: Kotnbia

with Echinoidea: Meotnicola

with Ascidiacea: Heteranthessius

with Turbellaria, Rhynchocoela, Polychaeta, Crinoidea, Asteroidea, Ophiuroidea, Echinoidea: Pseudanthessius.

In certain cases a single-host species may have

more than one species of copepod of one genus

associated with it. These species are sometimes

strikingly similar and yet morphologically distinct.

The alcyonacean Tubipora musica, for example,

has two closely related copepod associates, Plesiomolgus conjunctus and P. organicus (Humes and

Ho, 1967a). Another example is the polychaete

Sabellastarte magnified which has four species of

copepods associated with it, Nasomolgus firmus, N.

leptus, N. parvulus, and N. rudis (Humes and Ho,

1967c). The first three of these copepods have been

recovered together from a single host. Presumably

the three species occupy different microhabitats on

the host.

Phylogenetic Relationships of the Five Families

of Lichomolgoidea

Any interpretation of possible interrelationships of

the five poecilostome families considered here is

destined to be speculative. With the recent demonstration that intensive search in a small area can

uncover many new forms (as at Nosy Be, Madagascar), it is clear that the assemblage of copepods in

the Lichomolgoidea is far from complete. Almost nothing is known about the developmental

stages of the described species.

Of all the poecilostome families the Oncaeidae

seem to approach most closely the sabelliphilid

and lichomolgid types. The oncaeid segmentation and armature of legs 1-5 resemble closely

those of the sabelliphilids. There are, however,

important features by which the Oncaeidae differ

from the five families under consideration. Chief

among these are the nonprehensile nature of the

second antenna, the form of the mandible, and the

presence of a clawed maxilliped in both sexes.

Thorell (1859) referred his new genus Lichomolgus to the Ergasilidae, and later (1860) to the Sapphirinidae. The sabelliphilids and lichomolgids

appear to be rather far removed, however, from

the Ergasilidae, which have a very prehensile second antenna (especially in the female), lack a

maxilliped in the female, and have a different segmentation and armature of legs l^i. They are also

very different from the Sapphirinidae, which have

a pair of corneal lenses.

The Sabelliphilidae apparently are the most generalized and primitive of the five families. Their

legs 1-4 usually have 3-segmented rami, and their

bodies are generally not transformed. In several

genera there is a reduction in segmentation of the
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

endopods (Calypsarion, Calypsina, Caribulus,

Lichothuria, Synapticola), such reduction being

from anterior to posterior in the series of legs. In

one genus (Synapticola) there is a slight transformation of the body.

The Lichomolgidae (in the definition used here)

may be derived from the sabelliphilid stem. There

is a reduction of the endopod of leg 4 to two segments, one segment, or none. In a few genera the

body is modified or transformed, the greatest reduction and transformation being seen in Amarda

and Ravahina.

The Pseudanthessiidae may be derived from the

lichomolgid stem. Here there is a strong tendency

toward reduction of the endopods (and in certain

cases the exopods also) of legs 1-4 from posterior

to anterior. Their bodies are often modified or

transformed. Leg 5 lacks the free segment present

in the sabelliphilids and lichomolgids.

The Rhynchomolgidae apparently have evolved

from the pseudanthessiid stem. There is a very

strong reduction of the legs and the body is highly

transformed. Again there is no free segment in

leg 5.

The relationships of the Urocopiidae are more

obscure. (The family contains only Urocopia

singular is G.O. Sars, 1917, of which only the female

is known.) Here there is no reduction in legs 1-4

and the body is cyclopiform, yet there is no free

segment in leg 5. Possibly the family evolved from

the early pseudanthessiid stem, losing the free segment of leg 5 but retaining the sabelliphilid nature

of leg 4.

The phylogenetic tree presented in Figure 1 is

intended only to suggest possible relationships

within the Lichomolgoidea.

Superfamily LICHOMOLGOIDEA

Humes and Stock, 1972

First antenna usually 7-segmented. Second antenna

either 4-segmented or 3-segmented by a fusion of

the last two segments. Mandible simple, without

terminal elements, but often attenuated into a

slender lash. Maxillipcd present in both sexes, prehensile in the male.

Legs 1-4 typically with 3-segmented exopods and

endopods, but these frequently reduced or someFIGURE 1.—Possible relationships within the Lichomolgoidea.

Key to Families of the Superfamily Lichomolgoidea

1. Exopods of legs 1 and 2 in the female 2-segmented, in the male 1-segmented

Rhynchomolgidae

Exopods of legs 1 and 2 in the female 3-segmented, in the male at least 2-segmented 2

2. Leg 5 without a free segment «. 3

Leg 5 with a free segment (except a short papilla in Synapticola) 4

3. Leg 4 endopod 3-segmented Urocopiidae

Leg 4 endopod 1-segmented, reduced to a small knob, or absent v—nAantft—HAa*

4. Leg 4 endopod 2-segmented, 1-segmented, reduced to a small knob, or absent; legs 1-3

endopods 3-segmented (except in Amarda and Ravahina); the reduction in the endopods

occurring in a posterior to anterior series lichomolgidae

Legs 1-4 endopods 3-segmented in most genera; if leg 4 endopod 2-segmented, then legs

1-3 endopods also 2-segmented; the reduction of the endopods occurring in an anterior

to posterior series Sabelliphilidae
NUMBER 127

times absent. Leg 5 with at most a single free

segment, in some cases lacking a free segment entirely; armature usually consisting of three elements, two of them on the free segment if present.

Usually associated with marine invertebrates, in

some cases evidently parasitic, with the body form

modified or transformed.

REMARKS.—The superfamily name as originally

proposed by Humes and Stock (1972) inadvertently

contained an extra syllable. The name is here corrected to Lichomolgoidea, its intended form.

Family SABELLIPHILIDAE Gurney, 1927

Subfamily SABELLIPHILINAE Gurney, 1927

Legs 1-4 with 3-segmented rami in most genera.

Certain genera with 2-segmented endopods, especially in the male, the reduction occurring in an

anterior to posterior series as in males of Scambicornus. Leg 5 present in both sexes and with a

free segment, except in Synapticola where in both

sexes it is reduced to a short papilla and in 77mmnomolgus where in the male it is reduced to a

small lobe fused with the body.

Key to Genera of the Family Sabelliphilidae

1. Second antenna 3-segmented 2

Second antenna 4-segmented 3

2. Legs 1-4 with exopods and endopods 3-segmented in both sexes; leg 5 with a free segment

Henicoxiphium

Legs 1-4 with 3-segmented exopods and 2-segmented endopods in both sexes; leg 5 without

a free segment Synapticola

3. Second antenna with one claw on third segment and three or four claws on fourth segment 4

Second antenna with claws not present on both of these segments 5

4. Rostrum bifid; second antenna with first two segments enlarged, second segment with a

toothed crest and fourth segment with four claws SabeUiphilus

Rostrum not bifid; second antenna with first two segments not enlarged; second segment

without a toothed crest and fourth segment with three claws Myxomolgus

5. Second antenna with one claw on third segment, fourth segment without daws 6

Second antenna with 1-3 terminal claws, but without a claw on third segment 11

6. Ventral keel on genital segment of female and on first postgenital segment of male;

endopods of legs 1-4 in both sexes 2-segmented Calypsarion

Without a ventral keel on genital segment of female or on first postgenital segment of

male; endopods of legs 1-4 not entirely 2-segmented in both sexes 7

7. Body much elongated with broad triangular cephalosome; legs 1-4 with all exopods and

endopods 3-segmented in both sexes; leg 4 endopod with formula 0-1; 0-1; 1,1,1,1,1;

egg sacs very long with nearly linear eggs Lecanurhu

Body not unusually elongated and cephalosome not broadened; legs 1-4 with 3-segmented

exopods, but endopods sometimes 2-segmented; leg 4 endopod with formula other than

in Lecanurius; egg sacs not unusually long and eggs not linearly arranged 8

8. Legs 1-4 in both sexes with rami 3-segmented except legs 1-3 with endopods 2-segmented;

rostrum a slender beak; endopod of leg 4 with the formula 0-0;0-0;I,IIIJ Calypsina

Legs 1-4 in female with 3-segmented rami, but in male some endopods 2-segmented;

rostrum rounded or weakly developed; endopod of leg 4 with formula other than

in Calypsina 9

9. Mandible with a large spinelike element on base; leg 4 with endopod in female 0-l;0-l;II,l,

in male 0-l;II,l,l Uchothwria

Mandible without a large spinelike element on base; leg 4 with endopod having formula

unlike Lichothuria 10

10. Legs 1-4 with 3-segmented rami, except in the male where legs 1-2 have 2-segmented

endopods and legs 3-4 have 3-segmented endopods; leg 4 with endopod in both sexes

with formula 0-l;0-l;I,IIJI SeambUornus

Legs 1-4 with 3-segmented rami, except in male where all endopods are 2-segmented; leg

4 with endopod in female 0-l;0-l;I,II,II and in male O-l;IJIJI,l Caribtdus

11. Second antenna with one terminal claw 12

Second antenna with more than one terminal claw 15
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

12. Caudal ramus with two median terminal setae vestigial Diogenella

Caudal ramus with two median terminal setae normally developed IS

13. Third segment of leg 3 endopod with four elements; leg 5 in female with a free segment,

in male with the segment fused with body Thamnomolgus

Third segment of leg 3 endopod with five elements; leg 5 in both sexes with a free

segment 14

14. Terminal lash on second maxilla much shorter than adjacent seta Diogenidium

Terminal lash on second maxilla longer than adjacent seta Herrmatmella

15. Second antenna with two terminal claws 16

Second antenna with three terminal claws 17

16. Second maxilla lacking an auxiliary lash; maxilliped in female 2-segmentcd with blunt tip

Lichomolgidium

Second maxilla with an auxiliary lash; maxilliped in female 3-segmented and prehensile

Srrpuliphilus

17. Leg 4 endopod with formula 0-1;0-2;III; leg 3 endopod with 0-1:0-2:1.11.2 Parantheuhu

Leg 4 endopod with formula 0-l;0-l;lI; leg 3 endopod with 0-l;0-l;II.H2 or I.III.2

Modiolicola

Genus Calypsarion Humes and Ho, 1969

DIAGNOSIS.—Body modified, elongated, and slender. Urosome in the female 5-segmented; in the

male 6-segmented. A ventral keel on the genital

segment of the female and on the first postgenital

segment of the male. Caudal ramus with six setae.

Rostrum rounded. First antenna 7-segmented, in

the female with the formula 4, 13, 6, S, 4 -\- 1

aesthete, 2-f 1 aesthete, and 7 -f- 1 aesthete; in the

male with the same setation but without aesthetes.

Second antenna 4-segmented, the last segment set

off laterally; formula 1, 1, 3 -j- one large claw, and

7.

Labrum with two widely diverging lobes. Mandible with the base having a few small spiniform

processes on the convex edge, the blade with its

convex side finely spinulose (not a single row of

spinules as in related genera). Paragnath a small

hairy lobe. First maxilla with two or three elements and a small knob. Second maxilla of the

usual lichomolgid type. Maxilliped of the female

3-segmented with a pointed tip; in the male 4-

segmented (assuming that the proximal part of the

claw represents a fourth segment).

Legs 1-4 in both sexes with 3-segmented exopods

and 2-segmented endopods. Legs 3 and 4 exopods

Key to Species of the Genus Calypsarion

FEMALES

1. First maxilla with one terminal seta; distal spine on posterior surface of second segment

of second maxilla club-shaped and spinulose C. Upturn

First maxilla with two terminal setae; distal spine on posterior surface of second segment

of second maxilla attenuated and laterally barbed 2

2. Genital segment in dorsal view with more or less rounded lateral margins C. carhmtum

Genital segment in dorsal view expanded laterally in anterior third and slender in posterior

two-thirds C. tentosum

MALES

1. Inner margin of second segment of maxilliped greatly produced to form a long lobe

crowned with obtuse spines and two setae C. $entosum

Inner margin of second segment of maxilliped not produced 2

2. First maxilla with one terminal seta; distal spine on posterior surface of second segment

of second maxilla dub-shaped and spinulose; inner surface of second segment of maxilliped with one row of obtuse spines and two setae C. Upturn

First maxilla with two terminal setae; distal spine on posterior surface of second segment

of second maxilla attenuated and laterally spinulose; inner surface of second segment of

maxilliped with 2-3 rows of conical spines and two setae C. carbmtum
NUMBER 1X7

with the third segment having 11,1,5. Leg 4 endopod with the formula O-1;I,III,I,1 or 0-1 ;I,

111,1. Leg 1 endopod with the same formula in

both sexes. Leg 5 with the free segment bearing

two setae. Leg 6 in the female represented by the

two setae and the process near the area of attachment of each egg sac (without these setae in C.

leprum); in the male by the ventral posterolateral

flap on the genital segment bearing two setae.

Other features as in the species below.

Endoparasitic in holothurians.

TYPE-SPECIES.—Calypsarion carinatum (Stock).

Calypsarion carinatum (Stock, 1968)

Scambicornus carinatus Stock, 1968, pp. 92-94, figs. 1-3

[from the holothurian Stichopus monotuberculatus (Quoy

and Gairaard), Dahlak Archipelago, Ethiopia].

Calypsarion leprum Humes and Ho, 1969

1 lit Kf 2

Calypsarion leprum Humes and Ho, 1969b, pp. 878-880,

figs. 1-29 [from the holothurians Actinopyga miliaris

(Quoy and Caimard), Actinopyga lecanora (Jaeger), and

Actinopyga mauritiana (Quoy and Gaimard), Nosy Hi

and vicinity, northwestern Madagascar].

Calypsarion sentosum Humes and Ho, 1969

Calypsarion sentosum Humes and Ho, 1969b, pp. 880-889,

figs. 30-56 [from the holothurian Bohadschia marmorata

Jaeger, Nosy Hi. Madagascar].

Genus Calypsina Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum a pointed

beak. First antenna 7-segmented, in the female

with the armature 4, 13, 6, 3, 4 -f 1 aesthete, 2 +

1 aesthete, and 7+ 1 aesthete. Second antenna

4-segmented, the fourth segment set off laterally;

formula 1, 1, 3 -f one large claw, and 6, the last

segment bearing only setae.

Mandible resembling that of Scambicornus. First

maxilla with three elements. Second maxilla of

the usual lichomolgid type. Maxilliped in the female 3-segmented with a pointed tip; in the male

FIGURE 2.—Calypsarion leprum Humes and Ho. Female: a, dorsal; b, anterior part of urosome;

c, second antenna; d, mandible; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes

and Ho, 1969b, figs. 1, S. 9, 10, 20, 23.) Length of female 1.22 mm, of male 139 mm.
10 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

4-segmented (assuming that the proximal part of

the claw represents a fourth segment).

In both sexes legs 1-3 with 3-segmented exopods

and 2-segmented endopods, and leg 4 with both

rami 3-segmented. Leg 4 exopod with the third

segment having 11,1,5. Leg 1 endopod 0-l;I,5,l,

without sexual dimorphism in the male. Leg 2

endopod 0-l;I,II,3,l. Leg 3 endopod 0-1 ;1,111,2,1.

Leg 4 endopod 0-0;0-0;I,III,I. Leg 5 in both sexes

with a free segment bearing two terminal elements.

Other features as in the species below.

Associated with holothurians.

TYPE-SPECIES.—Calypsina changeuxi (Stock and

Kleeton).

ETYMOLOGY.—The name is a diminutive form of

KotaniMo, a nymph who concealed Ulysses. Gender

feminine.

Calypsina changeuxi (Stock and Kleeton, 1963)

FlCURE 3

Preherrmannella changeuxi Stock and Kleeton, 1963a, pp.

697-701, figs. 9-11 [from esophagus of the holothurian

Holothuria tubulosa Gmelin, Banyuls, Mediterranean coast

of France].

"Copepode poecilostome".—Changeux, 1961, p. 55, footnotes

pp. 109, 110, 111 [from Holothuria tubulosa and Holothuria stellati Delia Chiaje, Banyuls].

Scambicornus changeuxi.—Humes, 1967a, p. 144; 1969f, p. 92.

FEMALE.—A paratypic female was studied in lactic acid without dissection. Its length is 0.80 mm

and its greatest width 0.30 mm. The genital segment is 112 x 104 n in dorsal view, broadest in its

anterior third and tapering posteriorly to 78 n

near the posterior end of the segment. The areas

of attachment of the egg sacs are located in the

anterior half of the segment.

The rostral beak is about 75 n long.

The caudal ramus is 58 x 26 u. The two long

median terminal setae are 180 M (inner) and 83 n

(outer). All the setae are naked.

Genus Caribulus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

f

FIGURE 3.—Calypsina changeuxi (Stock and Kleeton). Female: a, dorsal; b, rostrum; c, second

antenna; d, mandible; e, leg 1; /, leg 4. (From Stock and Kleeton, 1963a, figs. 9a,c,d,g, 10a,d.)

Length of female 0.77 mm, of male 0.60 mm.
NUMBER 127 11

Caudal ramus with six setae. Rostrum broadly

rounded. First antenna 7-segraented, in both sexes

with the armature 4, IS, 6, 3, 4 -f- * aesthete, 2

-j- 1 aesthete, and 7 -f- 1 aesthete. Second antenna

4-segmented with the formula 1, 1, 3 -|-

 on e

 l a rg

e

claw, and 7, with some of the terminal setae

jointed.

Labrum deeply incised medially. Mandible with

its base merging at an angle with a broad attenuated blade, serrated along the convex side and with

spinules along the concave side. Paragnath a small

hairy lobe. First maxilla with four elements. Second maxilla of the usual lichomolgid type. Maxilliped in the female 3-segmented with a pointed

tip; in the male 4-segmented (assuming that the

proximal part of the claw represents a fourth segment).

Legs 1-4 in the female with 3-segmented rami.

Leg 4 exopod in both sexes with third segment

having 11,1,5. Leg 4 endbpod in the female with

the formula 0-l;0-l;I,II,II. Legs 1-4 in the male

with 3-segmented exopods with their armature as

in the female; but these legs with 2-segmented

endopods, the second and third segments having

fused, the formula for leg 4 endopod being

0-1;1,11,11,1. Leg 1 endopod in the male with

0-1 ;1,1,5, the next to the outermost element on the

second segment being a barbed spine instead of a

seta as in the female. Leg 5 with a free segment

bearing two terminal elements. Leg 6 represented

in the female by the two setae near the genital

openings; in the male by a ventral posterolateral

flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with holothurians.

TYPE-SPECIES.—Caribulus sculptus (Humes).

ETYMOLOGY.—The name is a diminutive form

of Carib, alluding to the wide distribution of the

genus in the Caribbean Sea. Gender masculine.

Caribulus sculptus (Humes, 1969)

FlCURE 4

Scambicornus sculptus Humes, 1969f, pp. 81-93, figs. 37-78

[from the holothurians Isostichopus badionotus (Selenka),

Barbados, Puerto Rico, Jamaica, Bahamas; Ludwigothuria

FIGURE 4.—Caributus sculptus (Humes). Female: a, dorsal; b, second antenna; c, mandible; d,

leg 4 and intercoxal plate. Male: e, dorsal; f, leg 4. (From Humes, 1969f, figs. 37, 49. 52, 63,

65. 76.) Length of female 1.18 mm, of male 0.98 mm.
12 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mexicana (Ludwig), Bahamas, Jamaica, Curasao, Bonaire;

Actinopyga agassizii (Selenka), Bahamas, Jamaica; and

Ludwigothuria grisea (Selenka), Jamaica, Bonaire].

Caribulus sp.

Scambicornus sp.-Humes, 1969f, pp. 93, 94, figs. 79-85

[from the holothurians Isostichopus badionotus, Bahamas,

Barbados, Jamaica; Ludwigothuria mexicana, Bahamas,

Jamaica, Curasao; and Actinopyga agassizii, Bahamas, Jamaica].

REMARKS.—The female is unknown.

Genus Diogenella Stock, 1968

DIAGNOSIS.—Body elongated, more or less modified. Urosome in the female 5-segmented; in the

male 6-segmented. Caudal ramus with six setae,

the two median terminal setae vestigial and the

lateral seta displaced proximally. Rostrum broadly

triangular or elongated linguiform. First antenna

7-segmented, the formula for the armature being

4, 13, 6, 3, 4 -|- 1 aesthete, 2 + 1 aesthete, and

7 -f- 1 aesthete, with one seta on the second segment long and ciliated. Second antenna 4—segmented, with the formula 1, 1, 4, and 1 terminal

claw -f 5 small elements.

Labrum with the two lobes widely separated by a

shallow indentation. Mandible with the base tapering into an attenuated pectinate blade. Paragnath a

small hairy lobe. First maxilla with two elements.

Second maxilla of the usual lichomolgid type.

Maxilliped in the female 3-segmented with the

setae much reduced; in the male 4-segmented (assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami. Armature of

the usual lichomolgidiform pattern. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod with the formula O-1;O-1;II. Leg 1 endopod

with the same formula in both sexes. Leg 5 with

a free segment bearing two unequal terminal elements. Leg 6 represented by two setae near the

genital openings.

Key to Species of the Genus Diogenella

KNOWN FEMALES

1. Caudal ramus with ciliated inner and outer terminal setae; first segment of second antenna

with spinules; second maxilla with terminal and auxiliary lash (seta) slender and nearly

of equal length 2

Caudal ramus with inner terminal seta having only a few minute barbules and outer terminal seta naked; first segment of second antenna without spinules; second maxilla with

auxiliary lash (seta) stout and about half as long as terminal lash D. dtichmannae

2. Genital segment not tapered posteriorly, but of approximately same width throughout, with

areas of attachment of egg sacs near middle of segment; genital and first two postgenital

segments each with a posteroventral transverse row of small spines D. seticauda

Genital segment broadest anteriorly and tapered posteriorly, with areas of attachment of

egg sacs in anterior part of segment; genital and first two postgenital segments without

such rows of spines D. spinicauda

MALES

1. Caudal ramus with ciliated inner and outer terminal setae; first segment of second antenna

with spinules; second maxilla with terminal and auxiliary lash (seta) slender and nearly

equal in length 2

Caudal ramus with inner terminal seta having only a few minute barbules and outer

terminal seta naked; first segment of second antenna without spinules; auxiliary lash

(seta) of second maxilla stout and about half as long as terminal lash S

2. First three postgenital segments each with a posteroventral transverse row of small spines

D. seticauda

First three postgenital segments without such rows of spines D. tpinicauda

3. Rostrum linguiform and slender; ratio of greatest length to width of caudal ramus about 2:1

D. deichmarmae

Rostrum broad and subtriangular; ratio of greatest length to width of caudal ramus

about 3.6:1 D.
NUMBER 127 13

Other features as in the species below.

Lives in holothurians.

TYPE-SPECIES.—Diogenella spinicauda Stock.

REMARKS.—The female of D. impar is unknown.

The following keys are from Humes and Ho

(1970a).

Diogenella spinicauda Stock, 1968

FIGURE 5

Diogenella spinicauda Stock, 1968, pp. 97, 98, figs. 7, 8 [from

the holothurian Holothuria mexicana Ludwig, Curasao].—

Humes and Ho, 1970a, pp. 15-21, figs. 1-27 [from the

holothurians Ludwigothuria (=Holothuria) mexicana

(Ludwig), Bahamas, Puerto Rico, Jamaica; and Actinopyga agassitii (Selenka), Bahamas, Jamaica].

Diogenella deickmannae Humes and Ho, 1970

Diogenella deichmannae Humes and Ho, 1970a, pp. 25-31,

figs. 51-73 [from the holothurian Brandtothuria arenicola

(Semper). Barbados].

Diogenella impar Humes and Ho, 1970

Diogenella impar Humes and Ho, 1970a, pp. S1-S5, figs.

74-87 [from Brandtothuria arenicola (Semper), Barbados].

REMARKS.—The female is unknown.

Diogenella seticauda Stock, 1968

Diogenella seticauda Stock, 1968, pp. 98, 99, figs. 9, 10 [from

the holothurians Halodeima surinamensis (Ludwig) and

Holothuria impatiens (Forskal), Puerto Rico].—Humes and

Ho, 1970a, pp. 21-25, figs. 28-50 [from the holothurians

Semperothuria (=Halodeima) surinamensis (Ludwig)

and Brandtothuria arenicola (Semper), mixed at the time

of collection, Puerto Rico].

Genus Diogenidium Edwards, 1891

DIAGNOSIS.—Body cyclopiform, but elongated and

somewhat modified. Urosome in the female 5-segmented; in the male 6-segmented. Caudal ramus

with six setae. Rostrum a pointed beak (except

broadly rounded in D. spinulosum). First antenna

FIGURE 5.—Diogenella spinicauda Stock. Female: a, dorsal; b, caudal ramus; c, first antenna;

d, second antenna; e, mandible; /, leg 4 and intercoxal plate. Male: g, dorsal. (From Humes

and Ho. 1970a, figs. I, 6, 9. 10. 11. 19. 22.) Length of female 1.62 mm, of male 159 mm.
14 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

7-segmented, in both sexes with the armature 4,

13, 6, 3, 4 + 1 aesthete, 2 -f 1 aesthete, and 7

-\- 1 aesthete. Second antenna 4-segmented, with

the formula 1, 1, 4, and one terminal claw -|-

 s'x

setiform elements.

Labrum with two small widely separated lobes

and without a median incision. Mandible with the

basal region beyond the indentation having on its

concave side a row of slender spinules and on its

convex side a crest of stouter spinules, both rows

continuing on the long slender lash. Paragnath a

small hairy lobe. First maxilla with three elements. Second maxilla of the usual lichomolgid

form, but with the main lash much shorter than

the auxiliary lash. Maxilliped in the female 3-

segmented, in the male 4-segmented (assuming the

proximal part of the claw to be a fourth segment).

Legs 1-4 with 3-segmented rami. Armature

lichomolgidiform in pattern. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0—1;0—1;II. Leg 1 endopod with

the same formula in both sexes. Leg 5 with a free

segment bearing two terminal elements. Leg 6 represented by the setae (1 or 2 in the female, 2 in

the male) near the genital openings.

Other features as in the species below.

Lives in holothurians.

TYPE-SPECIES.—Diogenidium nasutum Edwards.

REMARKS.—The female of D. tectum is unknown.

Key to Species of the Genus Diogenidium

1. Rostrum without a pointed beak; second and third segments of rami of legs 1-4 with small

spines on posterior surface D. tpinulomm

Rostrum with a pointed beak; without such spines on rami of legs 1-4 2

2. Second segment of second antenna without fine ornamentation D. deform*

Second segment of second antenna with outer spinules S

3. Longest terminal seta on caudal ramus shorter than ramus; armature of last segment of

exopod of leg 2 usually 11,13 but sometimes 111,1.5 D. nasutum

Longest terminal seta on caudal ramus about twice the length of ramus; armature of last

segment of exopod of leg 2 always 111,13 (female unknown) D. tectum

Diogenidium nasutum Edwards, 1891

FIGURE 6

Diogenidium nasutum Edwards, 1891a, pp. 87-91, pi. 4,

figs. 12-19 [from the body cavity of the holothurian Muelleria (=Actinopyga) agassizii Selenka, Bahamas]; 1891b,

pp. 15-19, pi. 4, figs. 12-19 [from the body cavity of the

holothurian Muelleria (•=. Actinopyga) agassizii Selenka,

Bahamas].-Stock, 1968, pp. 100, 101, figs. 11, 12 [from the

body cavity of the holothurian Holothuria mexicana

(Ludwig), Puerto Rico, Curasao].—Humes and Ho, 1971,

pp. 172-177, figs, 1-24 [from Actinopyga agassizii (Selenka),

Bahamas; Ludwigothuria mexicana (Ludwig), Puerto

Rico, Jamaica; and Ludwigothuria grisea (Selenka), Jamaica].

Diogenidium deforme Stock, 1968

Diogenidium deforme Stock, 1968, pp. 102-104, figs. 15, 16

[from the holothurian Holothuria glaberrima Selenka,

Puerto Rico].—Humes and Ho, 1971, pp. 182-188, figs.

48-72 [from the holothurians Brandtothuria arenicola

(Semper), Barbados; and Ludwigothuria mexicana (Ludwig) , Puerto Rico, Bahamas].

Diogenidium spinulosum Stock, 1968

Diogenidium spinulosum Stock, 1968, pp. 101, 102, figs. IS,

14 [from the holothurian Isostichopus badionotus (Selenka), Puerto Rico].-Humes and Ho, 1971, pp. 177-182.

figs. 25-47 [from Isostichopus badionotus (Selenka), Puerto

Rico, Jamaica].

Diogenidium tectum Humes and Ho, 1971

Diogenidium tectum Humes and Ho, 1971, pp. 188-189,

figs. 73-86 [from the holothurian Actinopyga agassizii

(Salenka), Jamaica, Bahamas].

REMARKS.—The female is unknown.

Genus Henicoxiphhim Illg and Humes, 1971

DIAGNOSIS.—Body cydopiform, elongated. Urosome in the female 5-segmented; in the male 6-

segmented. Caudal ramus with six short setae.

Rostrum rounded posteroventrally. First antenna

7-segmented, in the female with the armature 4,

13, 6, 3, 4 + 1 aesthete, 2 -f 1 aesthete, and 7
NUMBER 127 15

FIGURE 6.—Diogenidium tuuutum Edwards. Female: a, dorsal; b, second antenna; e, mandible;

d, second maxilla; e, leg 4 and intercoxal plate; /, leg 5. Male: g, dorsal. (From Humes and

Ho. 1971, fig«. 1. 7, 8, 10. 16-18.) Length of female 138 mm. of male 1.49 mm.

-j- 1 aesthete; in the male with 4, IS -f- 1 aesthete,

6, 3 -f- 1 aesthete, 4 -+- 1 aesthete, 2 -j- 1 aesthete, and 7 -|- 1 aesthete. Second antenna 3-segmented, with the formula 1, 1, 3 -j- terminally one

claw, one clawlike element, and five small setiform

elements.

Labrum with two widely separated lobes. Mandible without a basal indentation; lash with a row

of spinules on the concave side and numerous

spinules on the convex side. Paragnath a small

lobe. First maxilla with three elements. Second

maxilla of the lichomolgid type. Maxilliped in the

female 3-segmented, with vestigial setae and

rounded tip; in the male 4-segmented (considering the proximal part of the claw to represent a

fourth segment).

Legs 1—4 with 3-segmented rami. Armature of

the basic lichomolgidiform type. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod with the formula 0-l;0-l;I. Leg 1 endopod

with the same formula in both sexes. Leg 5 with a

free segment bearing two unequal terminal elements. Leg 6 represented in both sexes by two

setae near the genital openings.

Associated with ascidians.

TYPE-SPECIES.—Henicoxiphium redactum Illg

and Humes.

Henicoxiphium redactum Illg and Humes, 1971

FIGURE 7

Henicoxiphium redactum Illg and Humes, 1971, pp. 570-

577, figs. 1-26 [from the ascidian Styela plicata (Lesueur),

Florida, North Carolina].

Genus Herrmannella Canu, 1891

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded or

beaked. First antenna 7-segmented, with the for-
16

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 7 —Henicoxiphium redaclum Illg and Humes. Female: a, dorsal; b, caudal ramus; e,

second antenna; d, mandible; e, leg 4 and intercoxal plate; /. leg 5. Male: g, dorsal. (From

Illg and Humes. 1971. figs. 1, 5, 9, 11, 18-20.) Length of female 1.14 mm, of male 0.90 mm.

mula 4, 13, 6, 3, 4 + 1 aesthete, 2 -f 1 aesthete,

and 7 —f- 1 aesthete; in the male 4, 13 -\- 2 aesthetes,

6, 3 -f- 1 aesthete, 4 -f- 1 aesthete, 2+ 1 aesthete, and 7 —J— 1 aesthete. Second antenna 4-segmented, with the armature 1, 1, 3, and 1 terminal

claw -j- several setae.

Labrum with the two lobes separated by a shallow indentation. Mandible slender, the region

beyond the weak indentation pectinate; lash long.

Paragnath a small hairy lobe. First maxilla with

three elements. Second maxilla of the lichomolgid

type, with the terminal lash much longer than

the auxiliary elements. Maxilliped in the female

3-segmented; in the male 4-segmented (assuming

that the proximal part of the claw represents a

fourth segment).

Legs 1-4 with 3-segmented rami. Armature having the usual lichomolgidiform pattern. Leg 4 exopod with the third segment having 11,1,5. Leg 4

endopod with the formula 0—1;0—1;II (except

0—1;0—1;I,II in H. panopeae). Leg 1 endopod with

the same armature in both sexes, as far as known.

Leg 5 with a free segment bearing two terminal

elements. Leg 6 in the female represented by one

or two setae and in the male by two setae near the

genital openings.

Other features as in the species described below.

Associated with bivalve mollusks.

TYPE-SPECIES.—Herrmannella rostrata Canu.

REMARKS.—Herrmannella parva is omitted from

the following keys because of its uncertain position. The males of H. perplexa and H. valida are

unknown.
NUMBER 127 17

Key to Species of the Genus Herrmannella

FEMALES

1. Endopod of leg 4 with the formula O-1;O-1;I,II H. panopeae

Endopod of leg 4 with the formula O-1;O-1;II 2

2. Caudal rami much shorter than anal segment and only slightly longer than wide H. valida

Caudal rami distinctly longer than anal segment, clearly longer than wide 3

3. Rostrum rounded or with a bubble-like rounded tip 4

Rostrum beaked 8

4. Rostrum with a bubble-like rounded tip H. bullata

Rostrum broadly rounded 5

5. Caudal ramus with the ratio 6:1 H. perplexa

Caudal ramus with the ratio 3.4-4:1 6

6. Free segment of leg 5 with a pronounced proximal inner expansion H. inflatipes

Free segment of leg 5 without a proximal inner expansion 7

7. Prosome not tumid; caudal ramus 135 x 34 n (4:1); free segment of leg 5, 52 x 19 n

H. protothacae

Prosome tumid in nearly all specimens; caudal ramus 102 x 30 u (3.4:1); free segment

of leg 5, 43 x 16 (i H. mesodesmatis

8. First four segments of first antenna greatly swollen in relation to last three segments;

anal segment with a pair of crescentic ventral rows of spinules H. columbine

First four segments of first antenna only slightly swollen (as in H. saxidomi) or not

swollen; anal segment without a pair of crescentic rows of spinules 9

9. Two median terminal setae on caudal ramus very short, much shorter than the ramus;

caudal ramus very elongated, a little longer than last three postgenital segments

together H. barneae

One or both median terminal setae on caudal ramus elongated; caudal ramus not longer

than last three postgenital segments together 10

10. Third exopod segment of leg 4 with three outer spines and one terminal spine 11

Third exopod segment of leg 4 with two outer spines and one terminal spine 12

11. Third exopod segment of leg 4 with formula 111,1,4; first antenna without a plumose seta

H. dissident

Third exopod segment of leg 4 with formula III,1,5; first antenna with a plumose seta

H. caribaea

12. Second segment of second antenna slender, twice or more than twice as long as wide

(length measured along the inner margin of the segment) 13

Second segment of second antenna robust (I14 to l\/2 times as long as wide) 14

13. Endopod segments 1-3 of leg 4 without lateral spiniform processes H. saxidomi

Endopod segments 1-3 of leg 4 with distinct lateral spiniform processes H. th/elac

14. Lateral side of endopod segment 3 of leg 4 with two spiniform processes; lateral spiniform

processes on endopod segments 1 and 2 of legs 1—4 long H. pecteni

Lateral side of endopod segment 3 of leg 4 with one, sometimes hardly discernible, spiniform process; lateral spiniform processes on endopod segments 1 and 2 of legs' 1-4

short 15

15. Rostrum (in lateral view) hook-shaped, recurved; lateral spiniform process on endopod

segment 3 in leg 4 vestigial H. haploceras

Rostrum (in lateral view) straight, not recurved; lateral spiniform process on endopod

segment 3 in leg 4 distinct H. rostrata

KNOWN MALES

1. Endopod of leg 4 with the formula 0-l;0-l;I,II H. panopeae

Endopod of leg 4 with the formula 0-l;0-l;II 2

2. Rostrum rounded or with a bubble-like rounded tip 3

Rostrum beaked 6

3. Rostrum with a bubble-like rounded tip H. bullata

Rostrum broadly rounded 4

4. Body length 0.85 mm; claw of maxilliped 130 n long H. inflatipes

Body length at least 1.38 mm; claw of maxilliped at least 150 \i 5
18

SMI I HSON1AN CON 1 RIBU I IONS IO ZOOLOGY

5. Body length 1.44 mm; claw of maxilliped 172 u; subtcrminal clement of leg 5, 15 \L,

spiniform H. protothacae

Body length 1.38 mm; claw of maxilliped lf>0 u; subterminal element of leg 5, 10 \i.

setiform H. metodesmatis

6. First four segments of first antenna greatly swollen in relation to last three segments;

anal segment with a pair of crescentic ventral rows of spinules H. columbiat

First four segments of first antenna only slightly swollen (as in //. saxidomi) or not

swollen; anal segment without a pair of crescentic rows of spinules 7

7. Two median terminal setae on caudal ramus very short H. barneae

One or both median terminal setae on caudal ramus of usual elongated form 8

8. Third exopod segment of leg 4 with three outer spines and one terminal spine 9

Third exopod segment of leg 4 with two outer spines and one terminal spine 10

9. Third exopod segment of leg 4 with formula 111,1,4; first antenna without a plumose seta

H. diuidens

Third exopod segment of leg 4 with formula 111,1,5; first antenna with a plumose sria

H. coribaea

10. Second segment of first antenna shorter than fourth H. tivelae

Second segment of first antenna not shorter than fourth

11. Second segment of maxilliped with one of two setae modified (stout with

recurved tip)

Second segment of maxilliped with both setae unmodified

12. Rostrum in lateral view a strongly recurved hook; terminal armature on fourth »cgmcnt

of second antenna reduced (one daw and a small clawlike seta) H. haplocera*

Rostrum flat, not strongly recurved; terminal armature on fourth segment of second

antenna normal (one claw and several setae) IS

13. Outer spiniform processes on endopods of legs 1-4 strongly developed It. pecteni

Outer spiniform processes on endopods of legs 1-4 weakly developed H. roitrata

11

slender

H. taxidomi

12

Herrmannella rostrata Canu, 1891

Herrmannella rostrata Canu, 1891a, pp. 479, 480 [from the

bivalve Mactra stultorum Linnaeus, Pointe aux Oies,

Wimereux, France]; 1891b, p. 436 [from the bivalve

Pecten opercularis Linnaeus, Boulogne coast of France];

1892, pp. 236, 237, pi. 24, figs. 1-13 [from the bivalves

Mactra stultorum, Cardium edule Linnaeus, and Pecten

opercularis, Boulogne coast of France]; 1894a, p. 3; 1899,

p. 73.-T. Scott, 1894b, p. 259; 1905, pp. 206, 207 [from

Cardium edule, Scotland]; 1906, p. 354 [from Cardium

edule, Scotland] .-Pelseneer, 1929, p. 43 [from Tapes pullaster (Montague), Channel coast of France].—Fraser, 1932,

p. 279 [from Cardium edule, Morecambe Bay, Lancashire,

England].—Atkins, 1934, p. 674 [from Cardium edule,

Plymouth, England].

Lichomolgus agilis.—T. Scott, 1892, pp. 266, 267 [from

Cardium edule, Morecambe Bay, Lancashire, England, and

Cramond Island, Firth of Forth, Scotland; nomen nudum].

- T . Scott and A. Scott, 1892a, pp. 201-203, pi. 15, figs.

1-14 [from Cardium edule, Firth of Forth, Scotland, and

Morecambe Bay, Lancashire, England].—Herdman, 1893,

p. 83.-Thompson, 1893, pp. 207, 208, pi. 25, figs. 4, 8d

[from Cardium edule, Liverpool Bay, England].

Paranthessius rostratus.—Monod and Dollfus, 1932a, pp. 147,

148 [from Mactra corallina (Linnaeus) (=Mactra stultorum Linnaeus), Chlamys (Aequipecten) opercularis

(Linnaeus), Cardium edule Linnaeus, and Tapes pullaster

(Montagu) ].-Leigh-Sharpe, 1933, p. 113 [from the testis of

Cardium edule, Plymouth, England]; 1935, pp. 47, 48 [from

the gonad of Cardium edule, the mantle cavity of Chlamys

(Aequiptcten) opercularis, and the gonad of Paphia pullastro (Montagu), England].—Fouque and Franc, 1953. pp.

24, 25 [from the astuli.ui Ascidia mentula Miiller, northern

France].—Marine Biological Association, 1957, p. 176.—

Stock, 1959a, pp. 44, 45 [from the bivalve Tapn decuaalus

Linnaeus. Gulf of Naples, Italy].—Stock and dc Vos. 1960,

p. 206 [from Cardium edule, The Netherlands]—Bruce,

Colman, and Jones, 1963, p. 128 [from Cardium edule,

Port Erin, England].-Jepsen, 1965, pp. 284, 294, 297, 304

[from Cardium edule, Germany].—Crothers, 1966, p. 52

[from Cardium edule, Wales].

Paranthessius (Herrmannella) rostratus.—Bocquet and Stock,

1959a, pp. 241-243, figs. 1, 2 [from the bivalves Cardium

edule, Solen marginatus Don., Ensis siliqua (Linnaeus),

Spisula solida (Linnaeus) , Lutraria lutraria (Linnaeus) ,

Meretrix chione (Linnaeus), and Tapes pullastra (Montagu) , Channel coast of France; and from Macoma balthica

(Linnaeus) and Spisula subtruncata (da Costa), The

Netherlands]; 1963, p. 296.-Stock, 1964a, p. 67 [from the

bivalve Venerupis pullastra (Montagu) , The Netherlands].

Paranthesius rostratus.—Green, 1968, pp. 324, 325.

Herrmannella barneae (Pelseneer, 1929)

Ischnurella barneae Pelseneer, 1929, pp. 40, 41, fig. 4 [from

the bivalve Pholas (Barnea) Candida, Wimereux, France].

Ischnurella.-Rzncurel, 1954, pp. 855, 856, fig. lib.

Paranthessius barneae.—Bocquet and Stock, 1958b, pp. 604-

609, fig. 1 [from the mantle cavity of the bivalves Barnea

Candida (Linnaeus), Langue de Chien, Ambleteuse-sur-
NUMBER 127 19

Mn (Pas-de-Calais), France; and Pholas dactylus Linnaeus, Brest, and Longues (coast of Calvados), France].—

Gotto, 1962, p. 102.-Glac.on, 1971, p. 23 (unnumbered).

Parantheisiui (Herrmannella) barneae.—Bocquet and Stock,

1959a, pp. 248, 249 [from the mantle cavity of Ostrea

edulis Linnaeus, Baie de la Foret (ouest de Concarneau),

France].

Herrmannella bullata, new species

FIGURES 8-10

TYPE MATERIAL.—119 9 9 , 67 o* cf > and 37 copepodids from 12 bivalves, Chlamys hastata hericia

(Gould), in 36 fathoms, Flat Point, San Juan Islands, Washington, 48°31'06" N, 122°55'07" W, 3

July 1950, collected by M. Dunn. USNM 187867.

Holotype 9 , allotype, and 178 paratypes (114

9 $» 64 cf d1). along with the copepodids, are

deposited in the United States National Museum

(USNM), Washington, D. C, the remaining paratypes (dissected) in the collection of A. G. Humes

(AGH).

OTHER MATERIAL EXAMINED.—From Chlamys hastata hericia: 53 9 9» 50 cftf, and 33 copepodids,

in 20-30 fathoms, "potato patch," San Juan Islands, Washington, 48°35'30//

 N, 122°51' W, 26

June 1952, collected by P. L. Illg, fairly general

infestation; and 69 9 9» 97c?cf» and 81 copepodids from 32 hosts, in 105 fathoms, President Channel, San Juan Islands, Washington, 5 August 1950,

collected by P. L. Illg. USNM 187867.

From Chlamys rubida (Hinds): 4 9 $» 1 cf» in

20-30 fathoms, "potato patch," San Juan Islands,

Washington, 48°35'3O" N, 122°51' W, 26 June

1952, collected by P. L. Illg. Very sparse infestation.

FEMALE.—The body (Figure 8a) has the prosome not unusually broadened or thickened. The

length is 1.25 mm (1.07-1.38 mm) and the greatest

width 0.50 mm (0.46-0.56 mm), based on ten specimens in lactic acid. The ratio of the length to the

width of the prosome is 1.47:1. The ratio of the

length of the prosome to that of the urosome is

1.39:1. The segment of leg 1 is separated from

the cephalosome by a transverse dorsal furrow.

The epimeral areas of the pedigerous segments are

rounded except for that of leg 4, which is slightly

truncated.

The segment of leg 5 (Figure 86) is 78 x 132 A*.

Between this segment and the genital segment

there is no ventral intersegmental sclerite. The

genital segment is 176 x 169 M, slightly lobate

laterally in its broadened anterior two-thirds, narrowed in its posterior third. The areas of attachment of the egg sacs are located dorsolaterally near

the middle of the segment. Each area (Figure 8c)

bears two naked setae, 11 M and 20 n, and a small

spinous process. The three postgenital segments

are 70 x 79 M, 55 x 70 n, and 83 x 67 n, from anterior to posterior. The genital and each of the

postgenital segments bear posteriorly a dorsal and

ventral row of minute spinules.

The caudal ram us (Figure 8d) is elongated, 114"

x 26 n, or 4.38 times longer than wide. The outer

lateral seta is 65 M, the dorsal seta 68 n, the outermost terminal seta 81 n, the innermost terminal

seta 77 M, and the two long median terminal setae

300 u (outer) and 400 M (inner), both inserted

between dorsal (unornamented) and ventral (with

a group of small spinules) flaps. All the setae are

naked. A few surficial hairs occur on the ramus.

The body surface has minute hairs (sensilla) as

in Figure 8a.

The egg sac (Figure 8a) is elongated, 900 x

220 H, in the specimen drawn, but in one female

shorter, 625 x 230 /*. The sacs reach to the ends

of the longest ramal setae and contain many eggs

about 65 v- in diameter.

The rostrum (Figure 8e) has a round, bubblelike posteroventral tip.

The first antenna (Figure 8/) is 380 n long. The

lengths of its seven segments (measured along their

posterior nonsetiferous margins) are 18 (52 v- along

the anterior margin), 120, 44, 55, 55, 35, and 19 M

respectively. The formula for the armature is 4,

13, 6, 3, 4 -f 1 aesthete, 2+ 1 aesthete, and

7 + 1 aesthete. All the setae are naked.

The second antenna (Figure 8g) has a small

seta on each of the first two segments and three

setae on the third segment. The fourth segment

is 65 /* along the outer edge, 33 n along the inner

edge, and 40 n in greatest width, bearing a terminal

claw 68 M along its axis, two clawlike, bent setae,

and four small setiform elements. All the setae

are naked.

The labrum (Figure 9a) has two, widely divergent, slender lobes ornamented with minute

spinules.

The mandible (Figure 9b) is slender and armed

as in other species of the genus. The paragnath
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE %.—Herrmannella bullata, new species. Female: a, dorsal (A) ; b, urosome, ventral (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ram us, dorsal (D) ; e, rostrum, ventral

(E) ; /, first antenna, with arrows indicating positions of additional aesthetes in male, dorsal (E) ;

g, second antenna, posterior (t.). Scale: A, 0.5 mm; B, 0.2 mm; c, 0.05 mm; D, r, 0.1 mm.
g

FIGITLE 9.—Herrmannella bullata, new species. Female: a, labrum and paragnaths, ventral (D) ;

b, mandible, anterior (c); c, first maxilla, anterior (c); d, second maxilla, posterior (D) ; e,

maxilliped, antero-inner (D) ; /, area between maxillipeds and first pair of legs, ventral (F) ;

g, leg 1 and injercoxal plate, anterior (E) ; h, leg 2, anterior (E) ; i, endopod of leg 3, anterior

(E) . Scale: c, 0.05 mm; D, E, 0.1 mm; F, 02 mm.
22 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(Figure 9a) is a small hairy lobe. The first maxilla

(Figure 9c) bears three elements. The second

maxilla (Figure 9d) has patches of small spinules

on the first segment. The second segment has a

small outer (ventral) setule, a naked surficial posterior seta, and an inner (dorsal) distal barbed

spine, and it terminates in a lash with slender

spines along one edge. The maxilliped (Figure

9e) is slender. The first segment is unornamented.

The second segment bears two naked setae and has

a finely spinulose postero-outer surface. The third

segment carries a small setule and terminates in a

finely barbed spiniform process.

The area between the maxillipeds and the first

pair of legs (Figure 9/) is slightly protuberant. A

well-defined line connects the bases of the maxillipeds.

Legs 1-4 (Figures 9g-i, 10a) have the segmentation and armature as usual for the genus. In all

four legs the inner coxal seta is long and plumose

and the inner margin of the basis has a row of hairs.

The exopod of leg 4 has the formula I—0;I—1;II,I,5.

The endopod of leg 4 (Figure 10a) has the second

and third segments weakly separated; in some specimens without a break in the sclerotization of the

wall, in others with a slight break on the inner

side, but in all specimens at the intersegmental

junction there is a row of small spinules and a

trace of an intersegmental joint. The two terminal

fringed endopod spines are 44 M (outer) and 53 n

(inner).

Leg 5 (Figure 106) has a relatively small unornamented free segment 52 x 29 M in greatest dimensions. The two terminal setae are 55 n and

94 n, both with extremely short barbules. The

adjacent seta on the body is 35 /* and naked.

Leg 6 is represented by the two setae on the area

of attachment of each egg sac (Figure 8c).

The color in life corresponds to the color of the

sex of the scallop host: yellow-orange-red copepods

in male scallops which are bright orange, and

white copepods in female scallops which are milky

white. (These color notes are from a letter from

P.L. Illg to AGH dated 6 March 1970.)

MALE.—The body (Figure 10c) resembles that

of the female. The length is 0.97 mm (0.91-1.02

mm) and the greatest width 0.35 mm (0.35-0.37

mm), based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome

is 1.57:1. The ratio of the length of the prosome

to that of the urosome is 1.40:1.

The segment of leg 5 (Figure 10d) is 34 x 78 /*.

There is no ventral intersegmental sclerite. The

genital segment is 143 x 133 M. The four postgenital segments are 57 x 75 M, 57 x 65 M, 44 x

54 M, and 61 x 50 n from anterior to posterior,

posteriorly with dorsal and ventral spinules as in

the female.

The caudal ramus is like that of the female, but

smaller, 86 x 22 M, or 3.91 times longer than wide.

The body surface has small hairs (sensilla) as in

the female.

The rostrum is like that of the female.

The first antenna resembles that of the female,

but three aesthetes are added (at points indicated

by arrows in Figure 8/), so that the formula is 4,

13-1-2 aesthetes, 6, 3 -f 1 aesthete, 4 + 1 aesthete, 2 -f 1 aesthete, and 7 + 1 aesthete.

The second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla are like

those of the female. The maxilliped (Figure 10f)

is slender. The unornamented first segment has a

distinct inner bulge. The second segment bears a

row of spinules and two setae, one slender, the

other swollen proximally as in Figure 10/. The

third segment is unarmed. The claw is 133 n along

its axis, partially divided about midway, with a

small terminal lamella and two very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs resembles that of the female.

Leg 1 resembles that of the female, except that

the inner coxal seta is only partially plumose, the

distal part having small barbs as in Figure lOg.

Leg 2 resembles that of the female, except for

longer spiniform processes on the last segment of

the endopod (Figure \0h). Legs 3 and 4 are like

those of the female.

Leg 5 (Figure 10/) has a small unornamented free

segment, 29 x 13 m the two terminal setae are

39 M and 73 I*. The adjacent seta on the body is

29(i.

Leg 6 (Figure 10;) is a posteroventral flap on the

genital segment bearing two naked setae 29 n and

39 ft and two minute spiniform processes.

The spermatophore (Figure \0k), attached to

the female in pairs, is 83 x 43 n, not including the

neck.

The color in life resembles that of the female.
a

FIGURE 10.—Herrmannclla bullata, new species. Female: a, leg 4 and intercoxal plate, anterior

(E) ; b, leg 5, dorsal (D) . Male: c, dorsal (c); d, urosome, dorsal (B) ; e, maxilliped, inner

(E) ; /, swollen seta on second segment of maxilliped, outer (H) ; g, inner coxal seta of leg 1,

anterior (c); h, third segment of endopod of leg 2, anterior (c); i, leg 5, dorsal (c); ;, leg 6,

ventral (D) ; k, spermatophores, attached to female, dorsal (D) . Scale: B, C, 02 mm; c, 0.05

mm; D, E, 0.1 mm; H, 0.02 mm.
24

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ETYMOLOGY.—The specific name bullata (Latin,

swollen like a bubble) alludes to the bubble-like

tip on the rostrum.

REMARKS.—Hermannella bullata is unique

among the known species of the genus in the form

of its rostrum. The sexual dimorphism in leg 1

(with a half-plumose, half-spinose inner coxal seta

in the male) occurs in no other species in the genus

so far as known.

Hemnannella caribaea (Humes, 1970)

Paranthessius caribaeus Humes, 1970a, pp. 615-621, figs. 26-

51 [from the bivalves Chama sinuosa Broderip, Barbados,

Jamaica, Puerto Rico; and Pseudochama radians Lamarck,

Jamaica].

Hemnannella Columbia* (Thompson, 1897)

FIGURES 11-14

Pseudolichomolgus columbiae Thompson, 1897a, pp. 87-89,

pi. 8, figs. 1-10 [in tow-net, Puget Sound].

Paranthessius columbiae.-lllg, 1949, pp. 402-413, figs. S3, 34

[from the bivalves Schizothaerus nuttalli (Conrad) and

Protothaca tenerrima (Carpenter), California].

MATERIAL EXAMINED.—213 $ $, 115 o*cf» and

154 copepodids from the mantle cavity of the bivalve Schizothaerus (z=Tresus) capax Gould, Garrison Bay, San Juan Island, Washington, 7 April

1962, collected by Jack Pearce. (Herrmannella

panopeae also in this collection.)

FEMALE.—The body (Figure lla,&) is elongated,

with the cephalosome flattened dorsally in lateral

view. The length is 1.90 mm (1.73-2.14 mm) and

the greatest width 0.61 mm (0.58-0.64 mm), based

on ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.68:1. The

ratio of the length of the prosome to that of the

urosome is 1.43:1.

The segment of leg 5 (Figure lie) is 120 x 352

n. There is no ventral intersegmental sclerite between that segment and the genital segment. The

genital segment is 335 n long, a little broader in

its anterior half (290 n) than in its posterior half

(218 u), the two halves separated dorsally by a

transverse line. The areas of attachment of the

egg sacs are located laterally on the anterior half.

Each area (Figure lid) bears a single naked seta,

21 n. The three postgenital segments are 174 x

190 M, 153 x 161 u, and 130 x 121 u from anterior to posterior. The anal segment has a pair

of crescentic rows of spinules anteriorly placed on

the ventral surface (Figure We) and carries a row

of minute spinules on the posteroventral margin

on both sides.

The caudal ramus (Figure 11*) is 164 x 11 n in

greatest dimensions, or 4.8 times longer than wide.

The outer lateral seta is 62 M and finely barbed,

the dorsal seta 33 n and naked, the outermost

terminal seta 78 M and finely barbed, the innermost

terminal seta 65 n and naked, and the two long

median terminal setae 240 M (outer) and 500 u

(inner), both naked and inserted dorsally over a

small ventral flap ornamented with a few spinules.

The body surface has a few hairs (sensilla) as

in Figure 1 la.

The egg sac (Figure lla) is elongated, 1180 x

290 M, reaches nearly to the ends of the longest

ramal setae, and contains many eggs about 65 u in

diameter.

The rostrum (Figure 11/) has a long, pointed

beak arising from a base with lateral, slightly

knobbed expansions.

The first antenna (Figure llg) is 411 n long.

The lengths of its seven segments (measured along

their posterior nonsetiferous margins) are: 33 (75

fi along the anterior margin), 75, 37, 63, 61, 56, and

44 M respectively. The formula for the armature

is 4, 13, 6, 3, 4 + 1 aesthete, 2 -f- 1 aesthete, and

7 -(- 1 aesthete. All the setae are naked except

one on segment 5, one on segment 6, and four on

the last segment which are haired.

The second antenna (Figure Wh) has one seta

on the first segment, one seta and an outer row

of small spinules on the second segment, and three

setae on the third segment. The fourth segment

is 59 M along its outer edge, 37 •* along its inner

edge, and 24 u wide, with a claw 58 M along its

axis, two bent setae, and four other setae. All the

setae are naked.

The labrum (Figure 12a) has two, widely divergent, posteroventral lobes.

The mandible (Figure 126) is slender and bipectinate. The paragnath (Figure 12a) is a small

lobe with a few hairs. The first maxilla (Figure

12c) has three elements. The second maxilla is as

shown in Figure 12d. The maxilliped (Figure

12e) is unusually small and 3-segmented. The first
IIUHID 197

FIGURE \\.—Herrmannella columbine (Thompson). Female: a, dorsal (A); b, lateral (A); C,

urosome, dorsal (B) ; d, area of attachment of egg sac, lateral (c); e, caudal ramus and part of

anal segment, ventral (D) ; /, rostrum, ventral (E); g, first antenna, with arrow indicating

position of additional aesthete in male, ventral (D) ; h, second antenna, posterior (F) . Scale:

A, 1.0 mm; B, D, E, 02 mm; c, F, 0.1 mm.
FIGURE \Z.—Herrmannella columbine (Thompson). Female: a, labrum, with paragnaths indicated by broken lines, ventral (c); b, mandible, posterior (c); c, first maxilla, anterior (c);

d, second maxilla, posterior (F) ; e, maxilliped, antero-inner (c); /, area between maxillipeds

and first pair of legs, ventral (D) ; g, leg 1 and intercoxal plate, anterior (D) ; h, leg 2, anterior

(D) ; i, endopod of leg S, anterior (D) . Scale: c, r, 0.1 mm; o, 02 mm; c, 0.05 mm.
NUMBER 127 27

segment is broadly joined with the body and unarmed. The second segment is slender with two

setae. The third segment bears a small setule and

has an attenuated, finely barbed tip.

The ventral area between the maxillipeds and

the first pair of legs (Figure 12/) is not protuberant. A sclerotized bar connects the bases of the

maxillipeds.

Legs 1-4 (Figures I2g-i, 13a) are segmented and

armed as in other species of the genus. In all four

legs the inner coxal seta is long and plumose. In

legs 1-3 the inner margin of the basis has a row

of slender spines, but in leg 4 this margin has a

row of hairs. In legs 1-3 the anterior surface of

the basis between the rami has a similar row of

long slender spines, but in leg 4 these are reduced

to a few small spinules. In all four legs the first

and second endopod segments are without outer

distal spiniform processes, and the second segment

has a distal outer group of spinules. The two terminal fringed spines of the fourth endopod are

73 n (outer) and 140 M (inner); there is a row of

slender spinules on the segment near their insertions.

Leg 5 (Figure 13fc) has a moderately slender free

segment, 92 x 28 n in greatest dimensions (ratio

of 3.29:1), that is unornamented except for a row

of minute spinules near the insertions of the two

terminal naked elements which are 78 n and 99 MThe adjacent seta on the body is about 60 /* and

naked.

Leg 6 is represented by the single seta on the

area of attachment of each egg sac (Figure lid).

The color in life is unknown.

MALE.—The body (Figure 13c) is slender. The

length is 1.82 mm (1.62-1.93 mm) and the greatest width is 0.48 mm (0.45-0.50 mm), based on ten

specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.75:1. The ratio

of the length of the prosome to that of the urosome is 1.06:1.

The segment of leg 5 (Figure 13d) is 52 x 177 n.

There is no ventral intersegmental sclerite. The

genital segment is 220 x 243 n. The four postgenital segments are 122 x 156 M, 117 X 139 n, 104 x

122 \i, and 107 x 101 M from anterior to posterior.

The anal segment has ventral spinules as in the

female.

The caudal ramus is similar to that of the female, 166 x 36 n, with the ratio of 4.6:1.

The body surface has hairs as in the female.

The rostrum (Figure 13c) shows sexual dimorphism in the presence of two long spiniform processes extending posteroventrally from the lateral

margins of the basal area.

The first antenna is similar to that of the female

but has an additional aesthete on the fourth segment (at the location indicated by the arrow in

Figure llg), so that the formula is 4, 13, 3, 3 + 1

aesthete, 4 -{- 1 aesthete, 2+ 1 aesthete, and

7 + 1 aesthete.

The second antenna resembles that of the female but there are prominent additional spinules

along the inner side of the second segment (Figure

13/).

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 13g) is 4-segmented, assuming that the proximal part of the claw represents a fourth segment. The first segment is short

and broad, unarmed except for an inner knob.

The second segment inwardly has a slender seta,

a modified seta with a stout base and slender tip

(Figure 13/i), and two rows of spinules. The third

segment is small, obscure, and unarmed. The claw

is 230 M along its axis, weakly divided about midway, with a small terminal lamella, and with two

very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs (Figure 13») has a sclerotization slightly different from that in the female.

Legs 1-4 are segmented as in the female, with

the spinal and setal formula as in that sex. The

intercoxal plate of leg 1 (Figure 14a) is wider and

shorter than in the female (as in the following

three legs also). The coxa and basis of all four

legs are ornamented as in the female, except that

in leg 4 the anterior surface of the basis between

the rami bears a row of long slender spines as in

the previous legs (not reduced as in the female).

The exopods of legs 1-4 are ornamented as in the

female. The endopod of leg 1 (Figure 14&) has

the first segment with a distal outer group of

spinules, the second segment with a distal outer

spiniform process and a reduced group of spinules,

and the third segment with an outer spiniform

process near the insertion of the spine and a few

terminal spinules and small spinous processes. The

endopod of leg 2 (Figure 14c) lacks distal outer

spinules on the first segment, bears an outer spini-
FIGURE M.—Herrmannella columbine (Thompson). Female: a, leg 4 and intercoxal plate,

anterior (D) ; b, leg 5, dorsal (F) . Male: c, dorsal (A) ; d, urosome, dorsal (B) ; e, rostrum,

ventral (D) ; /, first and second segments of second antenna, posterior (F); g, maxilliped,

posterior (c); h, modified seta on second segment of maxilliped, posterior (H) ; i, area between maxillipeds and first pair of legs, ventral (D) . Scale: A, 1.0 mm; B, D, 02 mm; c, r, 0.1

mm; H, 0.02 mm.
NUMBER 127 29

e

Ficimc YA.-Herrmannella columbine (Thompson). Male: a, intercoxal plate of leg 1, anterior

(c); b, endopod of leg 1 anterior (F) ; c, endopod of leg 2, anterior (F) ; d, leg 5, dorsal (F) ;

e, leg 6, ventral (c). Scale: c, r, 0.1 mm.

form process on the second segment, but has only

a trace of an outer spiniform process on the third

segment. The endopod of leg 3 is like that of leg

2. The endopod of leg 4 is also similar to that of

leg 2, but the outer spiniform process on the second segment is smaller, and this process is absent

on the third segment.

Leg 5 (Figure \4d) has an unornamented free

segment, 48 x 16 n. The two terminal elements

are 44 M and 66 ii, and the adjacent seta on the

body is 44 p.

Leg 6 (Figure He) is a posteroventral flap on the

genital segment bearing two naked setae, 44 n and

56 M, and ornamented with two parallel rows of

spinules. In five specimens the anterior row had

an average number of 22 spinules (20-25) and the

posterior row averaged 15 (12-19).

The spermatophore was not observed.

The color in life is unknown.

REMARKS Several similar features suggest a

close relationship between H. columbiae and H.

saxidomi (IHg). Among such features in the female are the single seta on the area of attachment

of the egg sac, the enlarged four proximal segments of the first antenna, the reduction in size

of the maxilliped, the general ornamentation of

legs 1-4, and the appearance of leg 5. In the male

the similarities include the modified seta on the

second segment of the maxilliped, details of the

ornamentation of the endopods of legs 1 and 2,

and the two rows of spinules associated with leg 6.

Herrnumnella dissidens (Humes, 1970)

Paranthessius dissidens Humes, 1970a, pp. 605-613, figs. 1-25

[from the bivalves Chama sinuosa, Barbados, Jamaica, and

Puerto Rico; and Pseudochama radians, Jamaica and

Puerto Rico].

Herrnumnella haploceras (Bocquet and Stock,

1959)

Paranthessius (Herrmannella) haploceras Bocquet and Stock,

1959a, pp. 246-248, fig. 5 [from the bivalve Cardium erassum Gmelin, "La Vieille," Baie de Morlaix, Roscoff; Lucsur-Mer, Normandy, France].

Paranthessius colmani Reddiah, 1960, pp. 440-443, fig. 1

[from the bivalve Laevicardium crassum (Gmelin), Port

Erin, Isle of Man, England].-Bruce, Colman, and Jones,

1963, p. 128 [from Laevicardium crassum, Port Erin].
30 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Herrmannella inflatipes (Humes and Cressey,

1958)

Modiolicola inflatipes Humes and Cressey, 1958a, pp. 936-

938, figs. 101-126 [from the bivalve Mytilus perna Linnaeus,

Pointe Noire, Moyen Congo, West Africa].—Cheng, 1967,

pp. 315, 343.

Herrmannella mesodesmatis (Humes, 1967)

Paranthessius mesodesmatis Humes, 1967b, pp. 6-9, figs. 34-

43 [from the bivalve Mesodesma donacium Lamarck, Valparaiso, Chile].

Herrmannella panopeae (Illg, 1949)

FIGURES 15-16

Paranthessius panopeae Illg. 1949, pp. 413-417, figs. 35a-g,

36a-g, [from the bivalve Panope generosa Gould, California].—Stout, 1970, p. 68 [from Tresus (=Schizothaerus)

nuttalli, California].

The descriptive notes given below (and the accompanying figures) are based upon the study of

a freshly dissected male and female from a new

bivalve host, Clinocardium nuttalli (Conrad), at

Mitchell Bay, San Juan Island, Washington

(USNM 90700, collected by G. Comita, 7 August

1949, P. L. Illg. det.). An examination of the holotype 9 (USNM 85343) and the allotype (USNM

85344), both dissected and mounted on slides, and

three paratypes (1 ?, 2 d"o"), sent by Dr. Paul

L. Illg, showed a close similarity in the observable

features with the specimens from Clinocardium.

Features identical with Illg's original description

are omitted from the following partial characterization. Additional specimens (35 $ $, 2 cf cf. sent

to us by P. L. Illg) from the bivalve Schizothaerus

capax Gould at Garrison Bay, San Juan Island,

Washington, a second new host, have been studied.

FEMALE.—The length of the body (not including

the setae on the caudal rami) is 1.77 mm and the

greatest width is 0.51 mm, based on a specimen in

lactic acid. The segment of leg 5 (Figure 15a) is

90 x 180 M. The genital segment is 260 x 187 n,

slightly expanded in its anterior two-thirds. The

areas of attachment of the egg sacs are located

dorsolaterally near the middle of the segment.

Each area (Figure 15&) bears a naked seta 21 M

long, a small process, and a hyaline spot (the site

of a seta now lost?). The three postgenital segments are 107 x 118 M, 91 x 100 i*. and % x 83

u from anterior to posterior. The posteroventral

margin of the anal segment is unornamented.

The caudal ramus (Figure 15a) is 190 x 27 n

(the width taken just in front of the outer lateral

seta) with the ratio 7:1. All six setae are naked.

The ventral flap near the insertions of the two long

median terminal setae has a smooth margin.

The first antena is 330 n long. The lengths of

the seven segments (measured along their posterior

nonsetiferous margins) are 17 (44 M along the anterior margin), 72, 33, 55, 45, 42, and 39 »i respectively. The formula is 4, 13. 6, 3, 4 + 1 aesthete,

2 -f 1 aesthete, and 7 + 1 aesthete.

The second antenna (Figure 15c) has an inner

seta on the first segment, an inner seta and a row

of outer spinules on the second segment, and three

inner setae and a row of minute spinules on the

third segment. The fourth segment bears a stout

claw, 57 n along its axis, two slightly spiniform

setae, and four small hyaline elements.

The labrum, mandible (Figure 15d), and paragnath are much like those of H. protothacae

(Humes). The first maxilla (Figure 15<r) has three

elements. The slender maxilliped (Figure 15/) has

two setae on the second segment. The third segment bears two setae and has an attenuated, spiniform, finely barbed tip.

Legs 1-4 resemble closely those of H. protothacae and have the same spinal and setal formula.

The spines on the last segment of the endopod of

leg 2 (Figure 15g) are 28, 35, and 54 M from outer

to inner; these spines in leg 3 have proportionally

similar lengths. Leg 4 (Figure 15h) has three

fringed spines on the last segment of the endopod,

their lengths from outer to inner being 43, 54, and

82 M, and the greatest length of the segment 70 p.

Leg 5 (Figure 15t) has a free segment, 49 x 37 n,

ornamented with a few spinules at its outer (anterior) distal corner. Of the two naked terminal

elements, one is hyaline and 54 »* and the other is

spiniform and 138 ft. The naked seta adjacent to

the free segment is 55 n.

MALE.—The length of the body (without the

ramal setae) is 1.46 mm and the greatest width

0.40 mm, based on a specimen in lactic acid. The

segment of leg 5 (Figure 16a) is 65 x 117 M-. The

genital segment is 221 x 178 M. The four post-
FIGURE 15.—Herrmanella panopeae (Illg). Female: a, tiresome, dorsal (A) ; b, area of attachment of egg sac, dorsal (B) ; c, second antenna, posterior (c); d, mandible, anterior (B) ; e,

first maxilla, anterior (o); /, maxilliped, antero-inner (B) ; g, third segment of endopod of leg

2. anterior (c); h, leg 4, anterior (E) ; i, leg 5, dorsal (c). Scale: A, 03 mm; B, D, 0.05 mm;

c, E, 0.1 mm.
32

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 16.—Herrmannella panopeae (Illg). Male: a, urosome, dorsal (r); b, second antenna,

posterior (c); c, maxilliped, posterior (E) ; d, modified seta on second segment of maxilliped,

posterior (c). Scale: c, E, 0.1 mm; r, 02 mm; c, 0.02 mm.

genital segments are 81 x 96 n, 83 x 83 M, 60 x

73 ji, and 78 x 60 M from anterior to posterior.

The caudal ramus is like that of the female, but

smaller, 140 x 24 n.

The formula for the armature of the first antenna is 4, 13, 6, 3 -f 1 aesthete, 4 -f 1 aesthete,

2 -f- 1 aesthete, and 7+ 1 aesthete, one aesthete

having been added on the fourth segment, but

none on the second.

The second antenna (Figure 166) is more slender than in the female, the second segment bears

short spines along its inner surface, and the terminal claw is 62 n along its axis.

The maxilliped (Figure 16c) has on the inner

surface of the second segment two rows of spinules

and two naked setae, one of them modified (Figure

16d) with a swollen base. The claw is 211 /* along

its axis including the small terminal lamella.

The last segment of the endopod of leg 1 has

the same formula as in the female. The second

segment of the endopod of legs 1-3 shows an enlarged outer distal spinous process, as described by

Illg. Leg 4 is like that of the female.

Leg 5 (Figure 16a) has a rectangular free segment, 33 x 11 M. Its two terminal naked elements

are 39 |i and 50 M.

Leg 6 (Figure 16a) bears a row of spinules as

described by Illg and two naked setae, 22 n and

34 ft.

REMARKS.—Although H. panopeae differs from

all other species of Herrmannella'in having three

spines instead of two on the third segment of the

fourth endopod, its other features conform to the

generic concept. Perhaps the future discovery of

similarly armed species will indicate generic separation, but for the present it seems best to place

H. panopeae within Herrmannella.
NUMBER 127

Hemnannella parva Norman and T. Scott, 1905

Herrmannella parva Norman and T. Scott, 1905, pp. 299,

300 [from Plymouth Sound, England]; 1906, pp. 199, 200,

pi. IS. fig. 15, pi. 15, figs. 7-10, pi. 16. fig. 12, pi. 19,

fig. 5 [from Plymouth, England].—G. O. San, 1918, pp.

176, 177. pi. 99 [from Norway].

Hermannella parva.—Marine Biological Association, 1957, p.

176.

Paranthessius parvus.—lllg, 1949, p. 401.

Paranthessius T parvus.—Stock and Kleeton, 1963b, pp. 259,

260, fig. 8 [from the gorgonian Eunkella stricta (Bert.),

Banyuls, Mediterranean coast of France].

REMARKS.—The position of this species in Herrmannella is questionable. The terminal armature

of the second antenna, with three setiform claws,

sets it apart from other species in the genus.

Herrmannella pecteni (Sowinski, 1884)

Lichomolgus pecteni Sowinski, 1884, pp. 249-253, pi. 7, figs.

35. 36 [from Pecten sulcalus Born (-Chlamys glaber (Linnaeus) var. sulcata Born), Black Sea]; 1904, p. 94.

Hermannella rostrata.—Canu, 1891b, p. 436 [part; from

Pecten opercularis Linnaeus, Boulogne coast of France;

see Bocquet and Stock, 1959a, p. 245].

Herrmannella rostrata.-Cznu, 1892, pp. 236, 237, pi. 24,

figs. 1-13 [part; from Pecten opercularis, Boulogne coast

of France; see Bocquet and Stock, 1959a, p. 245].

Herrmannella pectint.—Stock, 1969, p. 221.

Pseudolichomolgus pectinis.—Pesti, 1909, pp. 263-265, pi. 2,

figs. 8-10, pi. 3, figs. 11-16 [from Pecten glaber (Linnaeus),

Barcola Canal, near Trieste, Italy].

Pestalichomolgus.—Wilson, 1932, p. 587 [new name for

Pseudolichomolgus Pesta, 1909].

Paranthessius pectinis.—Monod and Dollfus, 1932a, p. 148

[from Pecten glaber].—Illg, 1949, p. 401.—Bruce, Colman.

and Jones, 1963, p. 128 [from Chlamys opercularis,

Chlamys tigerina (Muller), and Pecten maximus (Linnaeus) , Isle of Man, England].

Paranthessius rostratus— Leigh-Sharpe, 1935, p. 47 [part;

from Pecten opercularis, England].

Paranthessius {Herrmannella) pectinis— Bocquet and Stock,

1957a, p. 427; 1959a, pp. 243-246, figs. 3, 4 [from Pecten

tigerinus Muller forma obsoletus (Penn.), Pecten opercularis, Pecten varius (Linnaeus), and Anomia ephippium

Linnaeus, northwestern France].

Lichomolgus pectini.—StoA, 1969, p. 220.

Herrmannella saxidomi (Illg, 1949)

FIGURES 17-19

Paranthessius saxidomi Illg, 1949, pp. 420-423, fig. 37a-h

[from Saxidomus nuttalli Conrad, California].

MATERIAL EXAMINED.—We have studied the holotype ? (USNM 85341) and the allotype (USNM

85342), both dissected and mounted on slides. In

addition, Dr. Paul L. Illg has generously provided

us with a para type of each sex. These have been

dissected and form the basis for the following redescription and figures.

FEMALE.—The length of the body in lactic add,

not including the setae on the caudal rami, is 1.71

mm; the greatest width is 0.66 mm. The prosome

(Figure 17a) is moderately broad. The urosome in

this specimen was damaged and thus could not be

drawn entirely. The genital segment (Figure 176)

is 260 x 203 M in greatest dimensions. The areas

of attachment of the egg sacs are situated laterally

in the wide anterior part of the segment. Each

area (Figure 17c) bears a single naked seta, 24 H.

The three postgenital segments are 133 x 126 M,

101 x 109 u, and 99 x 83 M from anterior to posterior.

The caudal ramus (Figure lid) is 140 M long,

27 u in greatest width proximally, and 19 M in least

width distally. The ratio of the length to the

greatest width is 5.2:1. All the setae are naked.

The rostrum (Figure lie) has a long slender

beak.

The seven segments of the first antenna (Figure

17/), with the first four segments a little swollen

as in H. columbiae, have the following lengths

(measured along their posterior nonsetiferous margins): 28 (63 u along the anterior margin), 62, 28,

54, 36, 32, and 23 M respectively. The formula for

the armature is 4, 13, 6, 3, 4 -j- 1 aesthete, 2+ 1

aesthete, and 7 -|- 1 aesthete. All the setae are

naked.

The second antenna (Figure 17g) has a row of

spinules along the outer margin of the second segment. The fourth segment, 48 A along its outer

edge, 30 u along its inner edge, and 23 M wide,

bears terminally a claw, 50 i* along its axis, and

five setae, two of them very small.

The labrum (Figure 17 h) has two divergent

posteroventral lobes. Its ventral surface is ornamented with retractile bosses.

The mandible (Figure 17i) is slender. The

paragnath (Figure \7h) is a small lobe with hairs.

The first maxilla (Figure 17;) bears three elements. The second maxilla (Figure 17ft) has retractile bosses on the outer surface of the first

segment. The lash on the second segment bears a

row of long slender spines. The dorsal (inner)

seta is strongly barbed. The maxilliped (Figure
FIGURE 17.—Hcrrmannella saxidomi (Illg). Paratypic female: a, prosome, dorsal (A) ; b, genital

and postgenital segments, ventral (B) ; c, area of attachment of egg sac, lateral (c); d, caudal

ramus, dislodged from anal segment, dorsal (c); e, rostrum, ventral (D) ; /, first antenna,

with arrows indicating positions of additional aesthetes in male, ventral (c); g, second

antenna, posterior (E) ; h, labrum and paragnaths, ventral (E) ; i, mandible, anterior (r); ;,

first maxilla, anterior (F) ; *, second maxilla, outer (F) . Scale: A, B, OJS mm; c, E, 0.1 mm;

D, 0J2 mm; r, 0.05 mm.
NUMBER 1*7

FICUIE \%.—Herrmannella saxidomi (Illg). Paratypic female: a, maxilliped, antero-inner (r) ;

b, leg 1 and intercoxal plate, anterior (c); c, leg 2, anterior (c); d, third segment of endopod

of leg 3, anterior (c); e, leg 4 and intercoxal plate, anterior (c); /, leg 5, dorsal (r). Scale:

c, 0.1 mm; r, 0.05 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 19.—Herrmannella saxidomi (Illg). Paratypic male: a, dorsal (B) ; b, second antenna,

posterior (F) ; c, second maxilla, outer (F) ; d, maxilliped, inner (F) ; e, modified seta on second

segment of maxilliped, inner (c); /, endopod of leg 1, anterior (F) ; g, endopod of leg 2, anterior

(F) ; h, leg 5, dorsal (F) ; i, leg 6, ventral (c). Scale: B, 0.5 mm; c, 0.1 mm; r, 04)5 mm; c.

02
NUMBER. 127

18<z), greatly reduced in size, has a large first segment. The second segment bears two setae. The

third segment bears a small seta and has an attenuated tip with spinules.

Legs 1-4 (Figure 186-e) have the same spinal and

setal formula as in other species in the genus (for

example, H. protothacae Humes). The inner margin of the basis is ornamented with a row of spinules in legs 1-3, but with hairs in leg 4. A row of

spinules is present on the lobe of the basis between

the rami, but this row is absent in leg 4. The

short naked seta on the third segment of the endopod of leg 1 in Figure 186, present on both

endopods of the paratype dissected, is considered

to be an abnormality. The last segment of the

endopod of leg 4 is 46 x 21 ii in greatest dimensions, with the two terminal fringed spines 37 M

(outer) and 90 p (inner). The distal outer margin

of the second segment of the endopod of legs 1-4

lacks a spiniform process.

Leg 5 (Figure 18/) has an unornamented free

segment 47 n long, 20 M wide at the proximal expanded area, and 14 M wide distally. The two

naked terminal setae are 75 P (inner) and 62 /*

(outer). The adjacent seta on the body is 55 /*

and feathered.

Leg 6 is probably represented by the seta near

the attachment of each egg sac.

The color in life is unknown.

MALE.—The body (Figure 19a) resembles in

general form that of the female. The length (without the ramal setae) in lactic acid is 1.10 mm and

the greatest width 0.35 m. The ratio of the length

to the width of the prosome is 1.53:1. The ratio

of the length of the prosome to that of the urosome is 1:1.

The genital segment is 164 x 161 /*. The four

postgenital segments are 83 x 99 n, 79 x 90 M,

70 x 77 /*, and 65 x 65 n from anterior to posterior.

The caudal ramus resembles that of the female,

but is somewhat shorter, 96 x 23 i», with the ratio

4.2:1.

The rostrum is similar to that of the female.

The first antenna is like that in the female, but

three aesthetes are added (Figure 19a), so that the

formula is 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete,

4 -f- 1 aesthete, 2-4- 1 aesthete, and 7 + 1 aesthete.

The second antenna (Figure 196) resembles that

of the female, but the second segment bears small

spines along its inner margin.

The labrum has two posteroventral lobes much

like those of the female. The ventral surface of the

labrum lacks the retractile bosses seen in the opposite sex.

The mandible, paragnath, and first maxilla resemble those of the female. The second maxilla

(Figure 19c) is much like that of the female, but

the first segment is ornamented with spinules, and

the second segment is not indented as it is in the

female. The maxilliped (Figure 19d) has a spiniform process on the distal inner (ventral) area

of the first segment. The second segment bears

stout spinules, arranged as illustrated in the figure,

and two setae, one slender, the other highly modified (Figure 19tf), stout, with a slender recurved

tip. The third segment is small. The claw is 156 M

along its axis, is weakly divided about midway,

and bears proximally the usual two very unequal

setae.

Legs 1-4 have the same spinal and setal formula

as in the female. The endopods differ from those

of the female chiefly in having a spiniform process

on the outer distal area of the second segment.

Such a process is most pronounced in leg 1 (Figure

19/), a little less so in leg 2 (Figure 19g), and still

smaller in legs 3 and 4. The exopods are like those

of the female.

The unornamented free segment of leg 5 (Figure

I9h) is 31 x 9 M.

Leg 6 (Figure 19i) is a posteroventral flap on the

genital segment bearing two naked setae, 28 M and

39 n, and ornamented with two oblique rows of

spinules (27 in the anterior row, 18 in the posterior

row).

The color in life is unknown.

Herrtnannetta tvvelae (Illg, 1949)

FIGURES 20, 21

Paranthessitu tivelae Illg, 1949, pp. 417-420, figs. S5h-j,

36h-m [from the bivalve T'wela stultorum (Mawe), California].

MATERIAL EXAMINED.—We have studied the dissections of the holotype (USNM 85346) and the

allotype (USNM 85345). The following redescription of the female is based on a paratype (from
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGUKE 20.—Herrmannella tivelae (Illg) . Paratypic female: a, dorsal (A) ; b, area of attachment

of egg sac, dorsal (B) ; c, posteroventral border of first postgenital segment, ventral (B) ; d,

caudal ramus, ventral (c); e, rostrum, ventral (c); /, first antenna, dorsal (c); g, second antenna, posterior (D) ; h, labrum, ventral (B) ; i, first maxilla, anterior (B) ; ;', second maxilla,

posterior (B) . Scale: A, 0.5 mm; B, 0.05 mm; c, D, 0.1 mm.
NUMBER it? 59

FIGURE 21.—Herrmannella tivelae (lllg). Paratypie female: a, maxilliped, posterior (a); b,

leg 1 and intercoxal plate, anterior (c); c, leg 2, anterior (c); d, third segment of endopod of

leg 3, anterior (c); e, leg 4 and intercoxal plate, anterior (c); /, leg.5, dorsal (B) . Scale: B,

0.05 mm; c, 0.1 mm.
40

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the bivalve Tivela stultorum Mawe, La Selva

Beach, California, 13 July 1943) sent to us by Dr.

Paul L. Illg.

FEMALE.—The length of the body (Figure 20a)

in lactic acid, excluding the setae on the caudal

rami, is 1.29 mm and the greatest width is 0.45

mm. The prosome is only moderately broad, the

ratio of the length to the width being 1.63:1. The

ratio of the length of the prosome to that of the

urosome is 1.26:1.

The genital segment is 174 x 138 /x. The areas

of attachment of the egg sacs are located dorsolaterally on the widest part of the segment. Each

area (Figure 20b) has a single naked seta 12 /*•

The three postgenital segments are 104 x 78 \i, 96

x 63 n, and 81 x 55 n from anterior to posterior.

The posteroventral borders of the genital segment

and the first two postgenital segments bear a row

of hyaline spinules (Figure 20c), each about 11 M

long, but the posteroventral border of the anal

segment is smooth.

The caudal ramus (Figure 20d) is 143 x 18 /t in

greatest dimensions, the ratio of the length to the

width being 7.9:1. The two long median terminal

setae are 154 M (outer) and 328 ji (inner). All six

setae are naked.

The rostrum (Figure 20^) has a pointed beak.

The first antenna (Figure 20/) is notable in having the second segment shorter than the fourth.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are: 8 (30 M

along the anterior margin), 46, 26, 55, 35, 42, and

33 n respectively. The formula for the armature is

the same as in H. saxidomi. All the setae are

naked.

The second antenna (Figure 20g) has small spinules along both outer and inner margins of the

second segment. The fourth segment, 48 n along

its outer edge, 25 /* along its inner edge, and 21 n

wide, bears a terminal claw 38 M along its axis

and five setae.

The labrum (Figure 20A), weakly sclerotized and

lacking the refractile bosses seen in H. saxidomi,

has two very divergent posteroventral lobes with

hyaline margins.

The mandible and paragnath are the same as in

H. saxidomi. The first maxilla (Figure 20i) has

three elements. The second maxilla (Figure 20;)

has a row of long spinules along the convex (ventral) border of the first segment. The maxilliped

(Figure 21a) resembles that of H. saxidomi, but

the last segment is small and globose rather than

attenuated and is smooth except for one minute

seta.

Legs 1-4 (Figure 2\b-e) have the same spinal and

setal formula as in H. saxidomi and other species

of the genus. The coxa of leg 1 has an outer and

slightly posterior protuberance. The coxa of leg 4

has sparser ciliation on the inner seta than in legs

1-3. The inner margin of the basis of leg 1 has a

row of broad spinules, but in legs 2-4 these spinules are replaced by hairlike spinules. On the

lobe of the basis between the rami in legs 1-3

there are a few stout spinules. In leg 4 such spinules are absent, and instead there is a row of

minute spines. The outer distal area of the first

and second segments of the endopods of all four

legs has a spinous process and an adjacent row of

spinules [these being much more slender and hairlike on the second segment of the endopod of leg

3 (Figure 2Id) and on the endopod of leg 4 (Fig

ure 21^)]. The last segment of the endopod of leg

1 is 61 x 20 n in greatest dimensions and has two

small outer spiniform processes. The two terminal

fringed spines are 52 M (outer) and 94 » (inner).

Leg 5 (Figure 21/) has an unornamented free

segment 39 n long, 17 /* wide proximally and 13

n wide distally. The two terminal elements are

76 n (inner) and 68 M (outer).

Leg 6 is represented by the seta near the attachment of each egg sac.

The color in life is unknown.

Herrmannella valida G. O. Sars, 1918

Herrmannella valida G. O. Sars, 1918, pp. 175, 176, pi. 98

[from Norway].

REMARKS.—The male is unknown.

Genus Lecanurius Kossmann, 1877

DIAGNOSIS.—Body modified, elongated, cephalosome broad and triangular. Urosome in the female

5-segmented; in the male 6-segmented. Caudal

ramus with six moderately short setae. Rostrum

not well defined posteroventrally. First antenna

7-segmented, in both sexes with the formula 4, 13,

6, 3, 4 -\- 1 aesthete, 2 -f- 1 aesthete, and 7

-f- 1 aesthete. Second antenna 4—segmented, with

the formula 1,1,3 —J— one large claw, and 7.
NUMBER 127 41

Labrum incised medially. Mandible with the

base not indented but having distally on its convex edge a blunt process and on its concave side

a pointed process; blade bipectinate. Paragnath

a small lobe with spinules. First maxilla with four

elements. Second maxilla with a short lash about

as long as the adjacent seta. Maxilliped in the

female 3-segmented with a pointed tip; in the

male 4-segmented (taking the proximal part of

the claw as a fourth segment).

Legs 1-4 in the female with 3-segmented rami.

Leg 4 exopod with the third segment having 11,1,5.

Leg 4 cndopod with the formula O-1;O-1;I,1,1,1,1.

Legs 1-3 in the male having the same formula as

in the female but showing sexual dimorphism in

other respects (the endopods of legs 1 and 2 being

2-segmented). Leg 5 in both sexes with a free segment bearing two setae. Leg 6 in the female

represented by the two setae near the area of attachment of each egg sac, in the male by the ventral

posterolateral flap on the genital segment bearing

two setae.

Other features as in the species below.

Lives inside holothurians.

TYPE-SPECIES.—Lecanurius intestinalis Kossmann.

REMARKS.—A key to the females has not been

prepared because the female of L. intestinalis is

unknown.

Key to Species of the Genus Lecanurius

MALES

Average body length 1.97 ram; ccphalosome laterally indented; claw of maxilliped with one

pointed prominence L. kossntannianus

Average body length 1.46 mm; ccphalosome approximately semicircular in outline, not indented;

claw of maxilliped with two blunt protuberances L. intestinalis

Lecanurius intestinalis Kossmann, 1877

Lecanurius intestinalis Kossmann, 1877, pp. 20-22, pi. 5, fig.

1 [from the intestine of the holothurian Muelleria Ucanora

Jaeger, Bohol, Philippine Islands].—Changeux, 1961, p. 55.

REMARKS.—The female is unknown.

Lecanurius kossntannianus Humes, 1968

FIGURE 22

Lecanurius kossmannianus Humes, 1968a, pp. 181-190, figs.

1-32 [from the holothurians Actinopyga Ucanora (Jaeger)

and Actinopyga miliaris (Quoy and Gaimard), region of

Nosy Be", Madagascar].

Lecanurius sp.

Lecanurius sp., copepodid.—Humes, 1968a, p. 190 [from the

holothurian Synapta maculata (Chamisso and Eysenhardt),

Ambariobe, near Nosy Be1, Madagascar].

REMARKS.—This form has been omitted from

the key because of its immaturity.

Genus Lichomolgidium Kossmann, 1877

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum linguiform.

First antenna 7-segmented, in the female with the

formula 3, 10, 6, 3, 4 -f 1 aesthete, 2 -f 1 aesthete,

and 6 -}- 1 aesthete. Second antenna 4-segmented,

with the armature 1, 1, 4, and II -|- 5, the two

terminal claws being slender and setiform.

Mandible with the basal area tapering to a

broad, fringed blade. First maxilla with three elements. Second maxilla of the usual lichomolgid

type except that it lacks the auxiliary lash. Maxilliped in the female 2-segmented with a blunt tip;

in the male 4-segmented (assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod with 0-l;0-l;II. Formula for the armature

of the typical lichomolgidiform type and similar

in both sexes. Leg 5 with a free segment bearing

two terminal elements.

Other features as in the species below.

Associated with ascidians.

TYPE-SPECIES.—Lichomolgidium sardum Kossmann.

REMARKS.—The genus Lichomolgidium, in its

present conception, is characterized chiefly by the

3-segmented endopod of leg 4, armed with two

elements on its distal segment, and by the absence

of an auxiliary lash on the second maxilla.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 22.—Lccanurius kossmannianus Hume*. Female: a, dorsal; b, second antenna; e, mandible; d, leg 3; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes, 1968a, figs. 1, 9,

11, 20, 22, 24.) Length of female 2.64 mm, of male 1.97 mm.

Key to Species of the Genus Lichomolgidhim

FEMALES

Terminal spiniform processes on leg 1 endopod rather strongly developed; leg 4 endopod with

the third segment not much tapered or constricted; leg 5 with the free segment 63 x 45 ft

L. sardum

Terminal spiniform processes on leg 1 endopod weakly developed; leg 4 endopod with the third

segment constricted and tapered distally; leg 5 with the free segment 68 x 40 \i .. JL cynthiae

Lichomolgidium sardum Kossmann, 1877

FIGURE 23

Lichomolgidium sardum Kossmann, 1877, pp. 19, 20, pi. 4,

fig. 3 [from the ascidian Cynthia (=Pyura) microcosmus

(Savigny), Mediterranean Sea].—Carus, 1885, p. 351.

Although the genus Lichomolgidium was described

and illustrated in a very superficial way by Kossmann,1

 the host, the locality, and the available

1

 Kossmann (1877, p. 19) himself admitted the "incompleteness" of his description: "Leider kann ich nur eine sehr

unvollkommene Darstellung geben, weil ich nur ein Exemplar in la Maddalena selbst unter schwierigen Umst&nden

untersuchen konnte."

descriptive notes indicated in the original publication give us a clue as to which copepod was described as Lichomolgidium sardum.

The host was called Cynthia microcosmus by

Kossmann, a tunicate now known as Pyura microcosmus (Savigny, 1816) or as P. saxngnyi (Philippi,

1843). We have examined material from this host

from the Mediterranean and the Atlantic as well

as from the closely related Halocynthia papillosa

(Linnaeus, 1758) from the Mediterranean. The

copepods from these different hosts and different

localities agree with the original description in the

more essential details: general structure of the legs

(with a 3-segmented endopod in leg 4) and finely
NUMBER 1*7 43

FIGURE 2S.—Liehomolgidium sardum Koumann. Female: a, endopod of leg 1 (E) ; b. leg 4

(E) ; c, kg 5 (D) . Scale: D, E, 0.05 mm.

toothed mandibular blade.2 We have ventured,

therefore, to consider the correspondence in host

and general morphology a sufficient basis to identify our material as L. sardum Kossmann.

It appears that L. sardum is very similar to a

copepod described from the Atlantic Ocean under

the name of Herrmannella cynthiae Brian (1924;

see also Gotto, 1961b). We are in doubt whether

the Atlantic material is conspecific with the Mediterranean samples, since a few minor differences

exist. The differences are so slight that, although

we treat here the Atlantic material as a different

species, more detailed studies based on more material from different localities and different hosts are

necessary to determine whether the differences

found are constant or due to intraspecific variation.

MATERIAL EXAMINED.—1 $, 1 o*» and 1 copepodid from Pyura microcosmus (Savigny), at the pier,

depth about 10 m, Banyuls, France, 8 August 1962;

4 ? 9» ^ o"o*»

 an d 5 copepodids from Halocyn-

"It appears to us that Kossmann actually described not

the mandible but the lash of the second maxilla.

thia papillosa (Linnaeus), at the pier, depth about

10 m, Banyuls, France, 11 August 1961, both collections made by JHS.

REMARKS.—The material from the Mediterranean mentioned above is so similar to the Atlantic specimens that it suffices to mention the

differences. Only three appendages show minor

differences, viz., legs 1-3. In the remaining appendages no differences whatsoever have been observed.

Leg 1 differs from the corresponding appendage

of L. cynthiae in having the spiniform processes on

the endopod more strongly developed (Figure

23a). This is especially distinct in the terminal

spiniform process. The endopod of leg 4 (Figure

236) is shorter than in L. cynthiae. The distal endopod segment is more cylindrical, less tapered,

and less constricted in the middle. The terminal

endopod spines are shorter (69 v- and 35 M. against

75 M and 50 n in L. cynthiae).

The free segment of leg 5 in the female (Figure

23c) is wider than in L. cynthiae (63 x 45 M, against

68 x 40 u in L. cynthiae).
44

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The body length of L. sardum ranges from 1.26-

1.42 mm for females and from 0.93-0.97 mm for

males.

Lichomolgidium cynthiae (Brian, 1924)

FICURES 24, 25

Hermannella cynthiae Brian, 1924, pp. 5-7, figs. 1-4 [from

ascidians (?Cynthia), Mauritania].

Paranthessius cynthiae.—Gotto, 1961b, pp. 88-91, fig. 2a-l

[from the pharynx of the ascidian Styela clava Herdman

(=S. tnammiculata Carlisle), Plymouth, England].

MATERIAL EXAMINED.—1 <j>, 1 <$ from Pyura

microcosmus, Roscoff, France, 1961 (gift of C. Monniot); 2 copepodids from Pyura sp., dredged in

about 25 m, Chateau du Taureau, Baie de Morlaix, France, 23 August 1954, collected by JHS.

FEMALE.—The length of the body is 1.51 mm.

The general shape has been illustrated adequately

by Gotto (1961b, fig. 2a). [Only minor additions

to Gotto's description and illustrations of the appendages are necessary. The mandible and first

maxilla, not mentioned in his paper, are described

here, and the sexual dimorphism found in the

male is treated.]

The ventral surfaces of urosomal segments 2-4

are armed at their posterior margins with a row

of small cuticular spinules (Figure 24a). Still

smaller spinules are present on the posterior margin of the fifth urosomal segment (anal). The

caudal rami are as described by Gotto.

The rostrum (Figure 24&) is elongated and

tongue-shaped.

The first antenna (Figure 246) is 7-segmented.

The armature of the segments is as follows: 3, 9

-f- 1 spinule, 3 -|- 3 spinules, 3, 4 -f- 1 aesthete,

2 + 1 aesthete, and 6 -f 1 aesthete. The second

antenna (Figure 24c) agrees with Gotto's account.

The mandible (Figure 24d) has a broad basal

portion tapering gradually into a medium-long

lash, armed on both sides with a hyaline, finely

denticulated fringe. The fringe on the lateral (concave) side of the lash is very transparent and hardly

visible by normal light microscopy; it shows up

very clearly, however, by the use of phase contrast.

The first maxilla (Figure 24d) is a tapering

lobe, armed with one terminal and one subterminal spine and with a short lateral seta. The second

maxilla (Figure 24e) is remarkable in that it lacks

an auxiliary lash. Thus, the second segment is

armed only with a subbasal smooth spine and a

medium-long apical lash, which has a hyaline,

sharply toothed medial margin and a smooth lateral margin.

The maxilliped (Figure 24/) shows up differently

in different positions. It is 2-segmented, with the

distal segment bearing two setae and having a

finely spinulose tip.

Legs l-A are as illustrated in Figures 24g, 25a-c.

The exopod of leg 3 is similar to that of leg 2.

The third segment of the exopod of leg 4 has the

armature 11,1,5. The formula of the endopod of

leg 4 is 0—1 ;0—1 ;II, the two terminal spines being

75 and 50 n in length.

Leg 5 (Figure 25d) has an ovate free segment

68 ii long and 40 M in greatest width, medially produced into an obtuse projection.

Leg 6 is represented by the two short, wide spinules near the genital openings.

MALE.—The length of the body is 1.06 mm. The

only male appendages that show marked sexual

dimorphism are the maxilliped and leg 5. The

maxilliped (Figure 25<r) has two large proximal

segments, the second of which is armed with two

spines and ornamented with numerous small cuticular spinules. The third segment is small and

unarmed. The fourth segment is probably represented by the proximal part of the long, strongly

recurved terminal claw with a short basal setule.

Leg 5 (Figure 25/) is not widened as in the female. Its median margin has a shallow notch.

Distally the segment bears a fiat short spine and

a long seta.

Genus Lichothuria Stock, 1968

DIAGNOSIS.—Body modified, elongated and slender. Urosome in the female 5-segmented; in the

male 6-segmented. Caudal ramus with six setae.

Rostrum weakly developed. First antenna 7-segmented, in both sexes the formula being 4, 13, 6,

3, 4 -f 1 aesthete, 2 -f- 1 aesthete, and 7 + 1

aesthete. Second antenna 4-segmented, with the

armature 1, 1, 3 -f- one claw, and 6 (three of these

setae being jointed and clawlike).

Labrum with a shallow posteroventral indentation. Mandible composed of a basal part and a

distal bipectinate blade at the junction of which

on the convex side there is a large spinelike element. Paragnath a small hairy lobe. First maxilla
NUMBER 127 45

FIGURE 2i.—Lichomolgidium cynthiae (Brian). Female: a, three posterior urosomal segments

and caudal rami, ventral (A) ; b, first antenna and rostrum (B) ; c, second antenna (B); d,

mandible and first maxilla (c); e, second maxilla (D) ; /, maxilliped (D); g, leg 1 and intercoxal plate (B) . Scale: A, B, 0.1 mm; c, 0.02 mm; D, 0.05 mm.
46

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 25.—Lichomolgidium cynthiae (Brian). Female: a, leg 2 and intercoxal plate (B) ; b,

endopod of leg 3 (B) ; c, leg 4 and intercoxal plate (B) ; A, leg 5 (D) . Male: e, maxilliped (D) ;

/, leg 5 (D) . Scale: B, 0.1 mm; D, 0.05 mm.
NUMBER 127 47

with two setae. Second maxilla of the usual lichomolgid type. Maxilliped of the female S-segmented

with a pointed tip; in the male 4-segmented (assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 in the female with 3-segmented rami;

in the male with 3-segmented exopods but 2-

segmented endopods. Leg 4 endopods in the female

with the formula 0—1;0—1;II,1; in the male with

0-l;II,l,l. Leg 1 endopod with the same formula

in both sexes. Leg 5 in both sexes with a free

segment bearing two elements. Leg 6 represented

in the female by the two setae and the spiniform

process near the area of attachment of each egg

sac; in the male by the ventral posterolateral flap

on the genital segment bearing two setae.

Other features as in the species below.

Endoparasitic in holothurians.

TYPE-SPECIES.—Lichothuria mandibularis Stock.

Lichothuria mandibularis Stock, 1968

FICURE 26

Lichothuria mandibularis Stock, 1968, pp. 95-96, figs. 4-6

[from the holothurian Halodeima atra (Jaeger), Eilat,

Gulf of Aqaba, Israel].-Humes and Ho, 1969b, pp. 889-894,

figs. 57-82 [from the holothurians Holothuria scabra

Jaeger, Holothuria aff. fuscopunctata Semper, Argiodia

nobilis (Selenka), and Ludwigothuria (=Halodeima) atra

(Jaeger), region of Nosy Be, Madagascar].

Genus Modiolicola Aurivillius, 1882

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, with

the armature 4, 13, 6, 3, 4 -f 1 aesthete, 2 -f 1

aesthete, and 7 + 1 aesthete; in the male 4, 13

-f 2 aesthetes, 6, 3 + 1 aesthete (or 3 + 2 aesFIGLRE 26.—Lichothuria mandibularis Stock. Female: a, dorsal; b, second antenna; c, mandible;

d, leg 4 and intercoxal plate. Male: e, dorsal; /, leg 1; g, endopod of leg 4. (From Humes and

Ho, 1969b, figs. 57, 63, 64, 73, 75, 78, 81.) Length of female 1.61 mm, of male 1.32 mm.
48 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

thetes), 4 -f- 1 aesthete, 2 -f- 1 aesthete, and 7

-(- 1 aesthete. Second antenna 4-segmented, with

the formula 1, 1, 3, and three terminal claws -ffour setae.

Labrum incised medially. Mandible with a slender base which, distal to the slight indentation,

has on its concave side a row of long spinules and

on its convex side a row of spinules; lash long and

pectinate. Paragnath a small hairy lobe. First

maxilla with four elements. Second maxilla of the

usual lichomolgid type, but the auxiliary lash short

and naked. Maxilliped in the female 3-segmented,

the third segment small, the setae vestigial; in the

male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami. Leg 3 endopod with the third segment having 11,11,2 or 1,111,2.

Leg 4 exopod with the third segment having 11,1,5.

Leg 4 endopod with the formula 0—I ;0—1;II. Leg

5 with a free segment bearing two terminal elements. Leg 6 represented by two setae near the

genital openings.

Other features as in the species below.

Associated with bivalve mollusks.

TYPE-SPECIES.—Modiolicola insignis Aurivillius.

REMARKS.—Detailed information on which to

construct a key for the males is not available, but

in general the following key to females will serve

for males also, allowing for somewhat smaller body

sizes.

Key to Species of the Genus Modiolicola

FEMALES

1. Second antenna slender with very short setae (easily overlooked) on one or both of fust

and second segments 2

Second antenna robust with conspicuous setae on both first and second segments S

2. Caudal ram us 2.58:1; spinules on concave side of base of mandible of one thickness, but

longest at either end of row M. hutgnit

Caudal ramus 4.2:1; spinules on concave side of base of mandible of two thicknesses, proximal

group slender, distal group stout M. gracilii

3. Caudal ramus distinctly less than 9:1 4

Caudal ramus clearly more than 9:1 5

4. Caudal ramus 5:1; body length 1:45-1.48 mm M. bifidus

Caudal ramus 6:1; body length 0.83 mm M. trabalis

5. Body length 1.90 mm; concave side of base of mandible with about 19 spinules of two sizes;

caudal ramus 11.9:1 M. inermis

Body length 2-40 mm; concave side of base of mandible with 9-11 spinules of one size;

caudal ramus 11:1 M. maximus

Modiolicola insignis Aurivillius, 1882

FIGURE 27

Modiolicola insignis Aurivillius, 1882a, pp. 40-44, pi. 6, figs.

1-10 [from the gill lamellae of the bivalve Modiola vulgaris

Linnaeus, Gullmarsfjord, Bohuslan, Sweden]; 1882b, pp.

43-48, pi. 13, figs. 1-8 [from the bivalves Modiola vulgaris

and Mytilus edulis Linnaeus, Bohuslan]; 1883, pp. 10-14,

pi. 2, figs. 1-10 [from Modiola xmlgaris, Gullmarsfjorden,

Bohuslan, pp. 39-44, pi. 4, figs. 1-8 [from Modiola vulgaris

and Mytilus edulis, Bohuslan].—Canu, 1891a, p. 482 [from

the bivalve Modiola modiolus, between Le Varne and Le

Colbart, northern France]; 1892, pp. 238-240, pi. 24, figs.

14-26 [from Modiola modiolus, between Le Varne and Le

Colbart, in the middle of the straits at Pas-de-Calais,

France]; 1899, p. 73.-T. Scott, 1893, pp. 207-209, pi. 4,

figs. 13-24 [from the bivalve Mytilus modiolus, Scotland];

1894b, p. 233; 1905, p. 207 [from Mytilus modiolus (=

Modiola modiolus), Scotland]; 1906, p. 354 [from Mytilus

modiolus, Firth of Forth, Scotland] Herdman, 1896, pp.

42, 44 [in washings of dredged material. Port Erin,

England].—A. Scott, 1896a, p. 153 [from Mytilus modiolus,

England]; 1896b, p. 78 [from Mytilus modiolus, Liverpool

Bay, England] G. O. Sars, 1918, pp. 173, 174, pi. 97 [from

Modiola modiolus, Norway].—Monod and Dollfus, 1932a,

p. 152 [from Modiolus modiolus (Linnaeus), Mytilus edulis

Linnaeus, and Mytilus galloprovincialis Lamarck].—Stock

and de Vos, 1960, p. 206 [from Mytilus edulis, The Netherlands].— Bresciani and Lutzen, 1962, p. 377.—Bruce, Colman,

and Jones, 1963, p. 128 [from Modiolus modiolus, Isle of

Man, England].—Porumb and Andriescu, 1964, pp. 94, 95,

figs. 1-19 [from Mytilus galloprovincialis Lamarck, Black

Sea].-Stock, 1964a, p. 67.

Modiolicola insignes.—T. Scott, 1894b, p. 259.

Modiocola insignis.—Marcus, 1970, p. 10.

Lichomolgus insignis.—Raffaele and Monticelli, 1885, pp.

302-307, figs. 13-16 [from the bivalve Mytilus galloprovincialis Lamarck, Naples, Italy].
FIGURE 27.—Modialicola insignis Aurivillius. Female: a, area of attachment of egg sac, dorsolateral (A); b, posterior border of first postgenital segment, ventral (A) ; c, second antenna,

anterior (inner) (A) ; d, mandible, anterior (B) ; e, second maxilla, posterior (B) ; /, maxilliped,

inner (B) ; g, endopod of leg 3, anterior (c); h, endopod of leg 4, anterior (A) ; i, abnormal

third segment of endopod of leg 4, anterior (B) ; /, leg 5, dorsal (B) . Scale: A, C, 0.1 mm; B,

0.05 mm.
50 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The descriptions and figures of this species given

by Canu (1892) and Sars (1918) are reliable in

most respects. There are, however, a few details

which need correction or addition. In the brief

notes below we attempt to provide this information. Our observations and drawings are based on

a single female from Modiolus barbatus. No males

were available to us at the time of the study.

MATERIAL EXAMINER.—From Modiolus barbatus

(Linnaeus): 2 ? $, 1 copepodid, Pte. du Binde,

Rade de Brest, France, 30 September 1955; 2 $ ? ,

Pte. du Binde, Rade de Brest, 26 March 1956, collected by JHS.

FEMALE.—The dimensions of the body (without

the ramal setae) are 1.56 x 0.49 mm (measurements

of a single specimen in lactic acid). The area of

attachment of each egg sac (Figure 27a) bears two

long slender setae, 36 u. and 38 ii, with a spiniform

process between them. The posteroventral edges of

the genital segment and the first two postgenital

segments bear a fringe of small spines (as in Figure 276).

The caudal ramus is 62 x 24 n in greatest dimensions, or 2.58:1.

The lengths of the seven first antennal segments

are 17 (39 n measured along the anterior edge),

78, 22, 41, 48, 40, and 21 M respectively.

The second antenna (Figure 27c) has the usual

armature, contrary to the figures of both Canu and

Sars. The single seta on the first and second segments and the three setae on the third segment are

extremely small (approximately 2.5 M long), and

visible only with very high magnification (oil immersion objective). The fourth segment bears three

jointed claws, the longest 44 n along its axis, and

four setae. The segment is 67 /* along its outer

edge, 56 n along its inner edge, and 16 M wide at

the middle.

The mandible (Figure 27d) has the row of spinules on its concave side, at either end, composed

of spinules longer and stouter than in the middle

of the row.

The second maxilla (Figure 27c) has on the second segment a very small setule near the outer

(ventral) margin, a superficial posterior naked

seta, and a small naked seta on the inner margin

(rather difficult to see in some cases). The lash

is of the usual type.

The maxilliped (Figure 27/) is 3-segmented,

with two minute setae on the second segment.

The four spines on the last segment of the endopod of leg 3 (Figure 27g) are 13, 20, 22, and

38 u from outer to inner. The endopod of leg 4

is shown in Figure 27/». The right endopod in

this female had, abnormally, three spines (Figure

27i) instead of two.

Leg 5 (Figure 27;) has an unornamented free

segment (25 x 15 M) bearing two unequal naked

terminal elements (52 n and 99 M).

REMARKS.—M. insignis may be recognized by the

rows of small spines on the posteroventral borders

of the postgenital segments (segments 1, 2 in the

female, 1-3 in the male) and by the reduction of

the armature of the second antenna. The mandible is distinctive in the nature of the spinules

on the concave margin.

Modiolicola bifidus Tanaka, 1961

Modiolicola bifidus Tanaka. 1961. pp. 266-269. pi. 32. figs.

6-11. pi. 33, figs. 1-12. pi. 34, fig*. 1-9, pi. 35. figi. 1-10

[from the mantle cavity of the bivalve Paphia, Saiebo Bay,

Japan; misspelled at Modiloicola.]

Modiolicola bifida.-K6, 1969a, p. 5; 1969b. pp. 83. 84, 86.

90, 91 [from Tapes japonica, Sasebo Bay. Japan].

Paranthessius sp. ?—K.6. Murakami, and Daidu, 1962, pp. 114,

116 [from the bivalves Tapes japonica Dethayei and Maetra sulcataria Reeve, Japan].

Modiolicola gracilis C B. Wilson, 1935

FIGURES 28-30

Modiolicola gracilis C. B. Wilson, 1935, pp. 781-782. pi.

25, figs. 8-12 [from the gill filaments of the bivalve Mytilus

californianus Conrad, Elkhorn Slough, Monterey Bay,

California; host erroneously reported by Wilson as

Mytilus edulis Linnaeus, but see below].—MacGinitie and

MacCinitie, 1949, p. 257-Ricketu and Calvin, 1952, p.

205.

MATERIAL EXAMINED (all from Mytilus californianus Conrad).—19 9 9 , 2 cf cf» Hopkins Marine Station, Pacific Grove, California, 7 November

1942, collected by P. L. Illg; 3 9 9 , 2 #<$, Bodega Bay, California, 23 January 1944, collected

by K. A. Hok; 5 9 9, 2 c? o*. two miles north of

Montara, San Mateo County, California, 25 October 1942, collected by P. L. Illg.

Although Wilson's type specimens were reported

from Mytilus edulis, this represents an error in labeling; the correct host is Mytilus californianus,

fide P. L. Illg.
length, dorsal (D) ; /, rostrum, ventral (c); g, first antenna, ventral (c); h, second antenna,

anterior (inner) (c); i, labrum, ventral (c); ;', mandible, posterior (E) ; k, first maxilla, anterior (E) . Scale: A, D, 0.5 mm; B, C, 0.1 mm; E, 0.05 mm.
52 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The male of this species is here described for

the first time.

FEMALE.—The average size is 1.47 x 0.52 mm,

based on five females in lactic acid. The form of

the body is like that in Wilson's figure 8 of plate

25. The segment of leg 5 (Figure 28a) is 104 x

174 n. The genital segment is distinctly bipartite,

the anterior part 120 x 174 M, the posterior part

101 x 130 »i. The three postgenital segments are

112 x 109 n, 101 x 86 n, and 68 x 78 fi from anterior

to posterior. The posteroventral borders of the

genital segment (posterior part) and of the first

two postgenital segments bear a row of fairly strong

spines (Figure 286) somewhat resembling those in

M. insignis. The anal segment has a posteroventral

row of very small spinules on each side. The areas

of attachment of the egg sacs are situated dorsolaterally and posteriorly on the anterior section of

the genital segment. Each area (Figure 28c) bears

two long setae with a small spiniform process between them.

The caudal ramus (Figure 28d) is 130 x 31 n.

The outer lateral seta is 59 M, the dorsal seta 23 n,

the outermost terminal seta 43 n, and the innermost

terminal seta 40 n. The two long median terminal

setae are 120 n (outer) and 250 M (inner), both

inserted between very short and unornamented

dorsal and ventral flaps.

The egg sac (Figure 28e) is elongated, 750-790

x 264 n, reaches a little beyond the caudal rami,

and contains many eggs each about 85 M in diameter.

The rostrum (Figure 28/) is broadly rounded

posteroventrally.

The first antenna (Figure 28g) is 312 M long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 18 (51 n

along the anterior margin), 86, 22, 31, 50, 48, and

24 n respectively. The formula for the armature

is the same as in Af. maximus. All the setae are

naked.

The second antenna (Figure 28/i) resembles in

its general form that of M. insignis. The setae on

the first three segments are short; the seta on the

first segment is 19 /* and the one on the second

segment is 4.5 n; and the three setae on the third

segment are 6, 8, and 4 /t. The fourth segment,

which is 66 n along its outer edge, 56 /* along its

inner edge, and 17 M wide, bears three claws (the

largest 40 n along its axis) and four setae. All the

elements are naked.

The labrum (Figure 28i) has two broad posteroventral lobes.

The mandible (Figure 28;) is similar in many

ways to that of M. insignis, but the spinules on the

concave edge, though similarly ranged in two

groups according to length, are of two sizes—the

proximal group slender, the distal group stout.

The paragnath is a small lobe with hairs as in

other species in the genus. The first maxilla (Figure 28A) has four elements, the longest very finely

barbed along one edge. The second maxilla (Figure 29a) resembles that of Af. insignis. The maxilliped (Figure 29b) has the second segment more

elongated than in Af. insignis, with apparently only

one small seta on this segment instead of two as in

that species.

The ventral area between the maxillipcds and

the first pair of legs is slightly protuberant.

Legs 1-4 (Figures 29c-/) have the same segmentation and armature as in Af. maximus, Af. inermis,

and Af. insignis. (The formula given by Wilson

erroneously includes both spiniform processes and

articulated spines.) The four spines on the last

segment of the endopod of leg 3 are 19, 32, 50, and

58 n from outer to inner. The inner seta on the

coxa of leg 4 is only 22 M and naked, instead of

long and feathered as in legs 1-3. The two spines

on the last segment of the endopod are 38 M and

77 n.

Leg 5 (Figure 29g) has an unornamented free

segment 34 x 19 M, armed terminally with a large

seta, 140 n, bearing a narrow fringe along one edge

and with a more slender, delicately feathered seta,

72 M. The adjacent seta on the body is 55 n and

lightly feathered.

Leg 6 is probably represented by the two setae

near the attachment of each egg sac (Figure 28c).

MALE.—The size is 1.08 x 0.34 mm, based on two

males in lactic acid. The body (Figure 30a) is

similar in general form to that of the female. The

ratio of the length to the width of the prosome is

1.64:1. The ratio of the length of the prosome to

that of the urosome is 1.07:1.

The segment of leg 5 is 60 x 143 M. The genital

segment is 190 x 213 n. The four postgenital segments are 75 x 120 ii, 81 x 91 n, 75 x 74 fi, and

64 x 67 n from anterior to posterior, ornamented

as in the female.
NUMBER 127

FIGURE 29.—Modiolicola gracilis C. B. Wilson. Female: a, second maxilla, posterior (B) ; b,

maxilliped, inner (E) ; c, leg 1 and intercoxal plate, anterior (c); d, leg 2, anterior (c); e,

endopod of leg 3, anterior (c); /, leg 4 and intercoxal plate, anterior (c); g, leg 5, dorsal (B) .

Scale: B, C, 0.1 mm; E, 0.05 mm.
54

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE SO.—Modiolicola gracitis C. B. Wilson. Male: a, dorsal (A) ; b, maxilliped, inner (B) ; e,

leg 5, dorsal (E) ; d, leg 6, ventral (c). Scale: A, 05 mm; B, C, 0.1 mm; E, 0.05 mm.

The caudal ramus is similar to that of the female, but smaller, 106 x 29 n.

The rostrum resembles that of the female.

The first antenna is similar to that of the female,

but two aesthetes are added on the second segment and one on the fourth segment, so that the

formula is 4, 13 + 2 aesthetes, 6, 3 -f 1 aesthete,

4 -f 1 aesthete, 2 + 1 aesthete, and 7+ 1 aesthete.

The second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla are like

those in the female.

The maxilliped (Figure 306) has two setae and

three rows of spinules on the second segment. The

claw is 164 n along its axis and bears proximally

two very unequal setae.

The ventral area between the maxillipeds and

the first pair of legs is similar to that in the female.
NUMBER 127 55

Legs 1-4 are like those of the female, with no

observed sexual dimorphism in armature or ornamentation.

Leg 5 (Figure 30c) has an unornamented free

segment (32 x 18 /x) with the larger terminal seta

(83 n) having a conspicuous fringe and the smaller

terminal seta measuring 52 /*•

Leg 6 (Figure 30d) is a posteroventral flap on

the genital segment having a plumose seta (23 M)>

a naked seta (42 n), and a spiniform process.

The spermatophore was not observed.

REMARKS.—This species seems to be more closely

related to M. insignis than to other species of

Modiolicola. This affinity is indicated by similarities in the second antenna, the division of the genital segment in the female, and the mandible.

Modiolicola inermis Canu, 1892

FlCURE 31

Modiolicola inermis Canu, 1892, pp. 240, 241, fig. 19 [from

the bivalve Pecten opercularis Linnaeus, Channel coast of

France]; 1894a, pp. 10-14 [from the bivalves Pecten maximus Linnaeus and Pecten opercularis Linnaeus, Channel

coast of France; specimens from Pecten maximus probably

are Modiolicola maximus; see Reddiah and Williamson,

1959, and below]; 1899, p. 74.-T. Scott, 1905, p. 207.-

Monod and Dollfus. 1932a, pp. 152. 153 [from Chlamys

(Aequipecten) opercularis (Linnaeus) and Pecten maxiFIGURE 31.—Modiolicola inermis Canu. Female: a, urosome, dorsal (A) ; b, posterior border of

first postgenital segment, ventral (B) ; c, mandible, posterior (c); d, third segment of endopod

of leg 4, anterior (D) ; e, leg 5, dorsal (B) . Male: /, urosome, dorsal (E) . Scale: A, 0J> mm;

B, D, 0.1 mm; c, 0.05 mm; E, 0.2 mm.
56 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

mus (Linnaeus) ].-Leigh-Sharpe, 1935, pp. 47, 48 [from

the mantle cavity of Chlamys (Aequipecten) opercularis

and Pecten maximus, England; specimens from Pecten

maximus probably are Modiolicola maximus, see below].—

Reddiah and Williamson, 1959. pp. 691-696, figs, la-f, 2al

[from the bivalve Chlamys opercularis, Isle of Man,

England].—Gotto, 1962, p. 103.—Bruce, Colman, and Jones,

1963, p. 128 [from Chlamys opercularis, Isle of Man,

England].

"Lichomolgide nouveau."—Canu, 1891b, p. 436 [from Pecten

opercularis, Boulogne-sur-Mer, France].

fLichomolgus maximus.—T. Scott, 1897a, p. 154 [from Pecten

opercularis, Loch Gair, off Inveraray, Upper Loch Fyne,

Scotland].

Hermannella maxima.—Norman and T. Scott, 1906, p. 199

[from Pecten opercularis, England].

Herrmannella maxima.—T. Scott, 1906, p. 354 [from Pecten

opercularis, Inchkeith, Firth of Forth, Scotland].

Paranthessius (Modiolicola) inermis.—Bocquet and Stock,

1957a, p. 427.

Paranthessius inermis.-Stock, 1960c, pp. 247, 248 [from

Pecten opercularis forma audouini Payr., Banyuls, Mediterranean coast of France].

MATERIAL EXAMINED.—10 $ <j>, 1 o* from Pecten

opercularis Linnaeus, in ± 20 m, Cochons Noirs,

northwest of La Vieille, near Roscoff, France, 27

July 1954; and 1 ? , 1 copepodid from the same

host, in ± 20 m, Cochons Noirs, northwest of Le

Cerf, near Roscoff, 27 July 1954; both collections

made by JHS.

Our specimens agree well with the description

and figures of Reddiah and Williamson (1959).

We offer here a few notes on selected features,

chiefly those differentiating this species from Af.

maximus.

FEMALE.—The dimensions of one female are 1.90

x 0.65 mm (in lactic acid). The segment of leg 5

(Figure 31a) is 104 x 195 n. The genital segment

is 234 x 224 fi in dorsal view, broadest near the

middle of the segment instead of posteriorly as in

Af. maximus. The three postgenital segments are

135 x 121 M, 107 x 94 u, and 153 x 82 u from anterior to posterior. The posteroventral borders of

the genital segment and the first two postgenital

segments bear a fringe of slender spinules (Figure

316) much as in Af. maximus.

The caudal ramus is 297 x 25 fi.

The mandible (Figure 31c) has about 19 long

spinules of two sizes on the concave side instead

of about 10 or 11 of one size as in Af. maximus.

The three spines on the last segment of the endopod of leg 2 are 20, 24, and 45 M (instead of

26, 26, and 62 u as in Af. maximus); the four spines

on the last segment of the endopod of leg 3 are

23, 28, 51, and 66 M (instead of 32, 39, 73, and

86 u); and the two spines on the last segment of

the endopod of leg 4 (Figure 3Id) are 37.5 /* and

78 u (instead of 45 M and 101 u).

The free segment of leg 5 (Figure 5\e) is 40 x

25 ji. The two terminal elements are 50 M and

105 u, more unequal than in Af. maximus where

they are 81 M and 112 n.

MALE.—The dimensions of one male are 1.30

x 0.43 mm. The segment of leg 5 is 50 x 121 M

(Figure 31/). The genital segment is 160 x 200 n.

The four postgenital segments are 73 x 122 M,

88 x 81 u, 70 x 65 M, and 96 x 60 n from anterior

to posterior.

The caudal ramus is 195 x 23 u.

The first antenna has the same arrangement of

aesthetes as in Af. maximus.

The claw of the maxilliped is 177 n along its

axis.

The two elements on leg 5 are 44 M and 69 n.

REMARKS.—Although Af. inermis and Af. maximus have many similarities, they may be readily

distinguished by careful study (which may necessitate dissection in order to see the mandible). The

chief points of difference are mentioned under Af.

maximus below.

Modiolicola maximus (Thompson, 1893)

FIGURES 32-34

Lichomolgus maximus Thompson, 1893, pp. 208-210, pi. 25

[from the bivalve Pecten maximus Linnaeus, Port Erin

Bay, England].-Herdman, 1894, p. 24; 1896. p. 50.-T.

Scott, 1897a, p. 154 [from Pecten opercularis, Scotland].—

d-Arcy Thompson, 1901, p. 47.

Modiolicola inermis.—Canu, 1894a, pp. 10-14. [See above

under M. inermis.]

Modiolicola maxima.—Reddiah and Williamson, 1959, pp.

696-699, figs, lg-h, 2m-x [from Pecten maximus, Port

Erin, England].-Gotto, 1962, p. 103; 1966, p. 194.-Bruce,

Colman, and Jones, 1963, p. 128 [from Pecten maximus,

Isle of Man, England].

Herrmannella maxima-T. Scott, 1905, p. 206 [from Pecten

maximus and Pecten opercularis, Scotland; specimens from

Pecten opercularis probably are Modiolicola inermis, see

above].

Hermannella maxima.—Farran, 1913, p. 6 [from Pecten maximus, Ballynakill Harbour, Ireland].—Williams, 1954, p73 [from Pecten maximus, Strangford Lough, Ireland].—

Marine Biological Association, 1957, p. 176.
NUMBER 127 57

The brief redescriptions below, based entirely on

specimens from Pecten maximus, supplement

previous descriptions of this species by Canu

(1891a, 1892) and by Reddiah and Williamson

(1959). Figures of the entire animal have been

omitted, since they have been adequately supplied

by the latter authors.

MATERIAL EXAMINED.—From Pecten maximus

(Linnaeus): 5 9 ? , S d*cf» Baie de la Foret, west

of Concarneau, France, 2 November 1955; 1 $ ,

Pte. du Binde, Rade de Brest, France, 27 March

1956; and 5 9 9 , 2 o* o* from two hosts, in ± 25 m,

Chateau du Taureau, Baie de Morlaix, Bretagne,

France, 18 September 1969; all collections made by

JHS.

From Pecten opercularis forma audouini Payr.: 1

9 , in 50-95 m, trawled between Banyuls and Cerbere, 9 October 1959, collected by JHS.

FEMALE.—The dimensions of the body in two

specimens measured in lactic acid are 2.40 x 0.80

mm and 2.24 x 0.77 mm. The segment of leg 5

(Figure 32a) is 117 x 226 it, The genital segment is

270 x 315 n. The three postgenital segments are

177 x 172 n, 140 x 115 M, and 198 x 101 M from anterior to posterior. The posteroventral borders of

the genital segment and of the first two postgenital

segments bear a row of fine spinules resembling a

fringe (Figure 32fe,c). The anal segment bears a

posteroventral row of minute spinules on each side.

The areas of attachment of the egg sacs are situated dorsolaterally in the anterior half of the genital segment. Each area (Figure 32d) bears two

setae, one 20 M and naked and the other 36 M- and

lightly feathered, with a spiniform process between

them.

The caudal ramus (Figure 52e) is 390 x 35 M,

or about 11 times longer than wide. The setae are

naked except for the feathered dorsal seta and the

innermost and outermost terminal setae which are

bilaterally haired proximally. The two long median terminal setae are inserted between dorsal

(unornamented) and ventral (with a marginal row

of minute spinules) flaps.

The elongated egg sac (Figure 32/) is 968 x 335

/*, reaches nearly to the end of the caudal ramus,

and contains many eggs about 91 n in diameter.

The rostrum (Figure 32g) is broadly rounded

posteroventrally.

The first antenna (Figure 32/i) is 465 n long.

The lengths of the seven segments (measured along

the posterior nonsetiferous margins) are 29 (72 I*

along the anterior margin), 145, 41, 74, 61, 42,

and 30 M respectively. The formula for the armature is 4, 13, 6, 3, 4 -f- 1 aesthete, 2+ 1 aesthete,

and 7 + 1 aesthete. All the setae are naked except one on the fifth, one on the sixth, and four

on the seventh segment which are feathered.

The second antenna (Figure 32t) has on the

fourth segment two unequal claws 69 /* and 39 i*

along their axes, a bent clawlike element, and four

setae. The fourth segment is 88 y- along its outer

edge, 52 n along its inner edge, and 44 /* wide. All

the elements of the appendage are naked except

the terminal bent clawlike element which has a

few minute distal barbs.

The labrum (Figure 32g) has two posteroventral lobes.

The mandible is shown in Figure 33a. The

paragnath is a small lobe with a few hairs and a

lamblike tip (see Figure 34e of the male). The

first maxilla (Figure 336) has four elements. The

second maxilla (Figure 33c) has on the second segment an extremely small setule near the outer (ventral) margin, a surficial posterior naked seta, and

a naked seta on the inner margin (this seta being

delicate and sometimes broken off). The terminal

lash has a row of graduated spinules. The maxilliped (Figure 33d) has three segments, the third

very small. The first segment is unarmed. The

expanded second segment bears two small naked

setae and a row of slender spinules. The minute

third segment bears only a very small terminal

hyaline setule.

The ventral area between the maxillipeds and

the first pair of legs (Figure $$e) is only slightly

protuberant.

Legs 1-4 (Figures $$f-h, 34a) have the same armature as described by Reddiah and Williamson

(1959).

Leg 5 (Figure 346) has an unornamented free

segment, 52 x 32 n. The two naked terminal elements are 81M and 112 n. The adjacent seta on

the body is 25 n and feathered.

Leg 6 is probably represented by the two small

setae near the attachment of each egg sac (Figure

32d).

MALE.—The dimensions of the body in two

specimens in lactic acid are 1.62 x 0.47 mm and

1.66 x 0.52 mm. The segment of leg 5 (Figure
FIGURE 52.—Modiolicola maximus (Thompson). Female: a, urosome, dorsal (A) ; b, posterior

part of genital segment and postgenital segments, ventral (B) ; c, posterior border of first

postgenital segment, ventral (c); d, area of attachment of egg sac, dorsal (c); e, caudal ramus,

dorsal (D) ; /, egg sac with side of urosome, dorsal (E) ; g, rostrum and labrum, ventral (F) ; h,

first antenna, ventral (F) ; i, second antenna, anterior (inner) (c). Scale: A, B, 0.5 mm; c, c,

0.1 mm; D, T, 0J2 mm; z, 1.0 mm.
NUMBER 127

FIGURE SS.—Modiolicola maximus (Thompson). Female: a, mandible, posterior (H) ; b, first

maxilla, anterior (H) ; c, second maxilla, posterior (c); d, maxilliped, inner (c); e, area between maxillipeds and first pair of legs, ventral (F) ; /, leg ) and intercoxal plate, anterior (F) ;

g, leg 2, anterior (F) ; h, endopod of leg S, anterior (F) . Scale: c, 0.1 mm; F, 02 mm; H,

0.05 mm.
60

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE H.—Modiolicola maximus (Thompson). Female: a, leg 4 and intercoxal plate, anterior

(F) ; b, leg 5, dorsal (c). Male: c, urosome, ventral (B) ; d, segments 1-4 of first antenna, ventral

(c); e, paragnath, ventral (i); /, maxilliped, inner (c) ; g, endopod of leg 1, anterior (c); h,

third segment of endopod of leg 2, anterior (c); i, leg 5 dorsal (H) ; /, leg 6, ventral (c). Scale:

B, 05 mm; c, c, 0.1 mm; F, 02 mm; H, 0.05 mm; i, 0.4 mm.
NUMBER 127 61

34c) is 52 x 143 n. The genital segment is 180 x

216 ii. The four postgenital segments are 91 x

156 n, 112 x 106 M, 83 x 82 M, and 123 x 75 »i

from anterior to posterior and are ornamented as

in the female.

The caudal ramus is similar to that of the female, but smaller, 93 x 11 /*.

The rostrum is like that of the female.

The first antenna resembles that of the female,

but two aesthetes are added on the second segment and two on the third segment (Figure 34d),

so that the formula is 4, 13 -{- 2 aesthetes, 6, 3

-f- 2 aesthetes, 4 + 1 aesthete, 2 -j- 1 aesthete,

and 7 + 1 aesthete.

The second antenna, labrum, mandible, paragnath (Figure 34e), first maxilla, and second maxilla

are like those in the female. The maxilliped (Figure 34/) has a row of weak striations on the first

segment. The second segment bears two naked

setae and two rows of spinules. The claw is 206 n

along its axis, indistinctly divided midway, and

armed with two very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Leg 1 resembles that of the female except that

the two spinous processes at either side of the spine

on the last segment of the endopod (Figure 34g)

are relatively longer. Leg 2 shows slight sexual

dimorphism in the nature of the spiniform processes on the last segment of the endopod (Figure

34A) but otherwise resembles that of the female.

Legs 3 and 4 are like those of the female.

Leg 5 (Figure 34i) has an unornamented free

segment that is 31 x 18 M in greatest dimensions.

The shorter terminal naked element is 52 n and

the longer element, fringed along the outer side, is

75 /*. The adjacent seta is 18 n and feathered.

Leg 6 (Figure 34;) is a posteroventral flap on the

genital segment bearing a plumose seta, 30 fi, a

naked seta, 45 n, and a finely barbed spine, 19 MThe spermatophore was not observed.

REMARKS.—Af. maximus may be distinguished

from Af. inermis, with which it appears to be closely

related, by the shape of the genital segment in the

female, the relative lengths of the two terminal

setae in leg 5 of the female, the size and number

of spinules on the concave edge of the base of the

mandible, and the size of the body.

Modiolicola trabalis Humes, 1959

Modiolicola trabalis Humes, 1959, pp. 315-319, figs. 250-268

[from the bivalve Area decussata Sowerby, Nosy Be,

Madagascar].

Genus Myxomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum elongated

and triangular. First antenna 7-segmented in the

female, with the armature 4, 13, 6, 3, 4 -j- 1 aesthete, 2+ 1 aesthete, and 7 -|- 1 aesthete; one of

the elements on the first segment and two on the

second segment enlarged to form stout denticulated

spines. Second antenna 4—segmented, the formula

being 1, 1, 3 -f- one claw, and 4 -{- three claws.

Labrum deeply indented medially. Mandible

with the blade at nearly a right angle to the base

and provided along its outer side with spinules

and on its inner side with lamellate teeth. First

maxilla with two or three elements. Second maxilla

of the usual lichomolgid type, with the auxiliary

lash about as long as the lash itself. Maxilliped

in the female 3-segmented, the third segment forming a short recurved claw; in the male 4—segmented

(assuming that the proximal part of the claw rep

resents a fourth segment).

Legs 1-4 with 3-segmented rami. Leg 4 exopod

with the third segment 11,1,4 or 11,1,5. Leg 4 endopod with the formula 0—l;0-l;II. Armature of

the lichomolgidiform type, with the formula similar in both sexes. Strong sexual dimorphism in

leg 2 endopod. Leg 5 with a free segment armed

with two elements. Leg 6 in the female represented by a seta and a spine near the genital openings; in the male by a ventral posterolateral flap

on the genital segment bearing two elements.

Other features as in the species below.

Associated with polychaete annelids.

TYPE-SPECIES.—Myxomolgus myxicolae (Bocquet

and Stock).

ETYMOLOGY.—The name is derived from the

Greek words pv%a (mucus) and HoXyo;. It alludes

to the fact that both known species of this genus

live in the mucous tube of the annelid rather than

on the host itself. Gender masculine.

REMARKS.—The genus is closely related to Diogenidium, Herrmannella, and Paranthessius but it

differs from these by the following characters: (1)
62 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

segments 1 and 2 of the first antenna have strong

flat spines; (2) segments 3 and 4 of the second antenna have one and three claws respectively; (3)

the blade of the mandible is medially strongly

toothed; and (4) the fifth leg has a strong basal

swelling.

Key to Species of the Genus Myxomolgus

FEMALES

Third segment of exopod of leg 4 with 11,1,4; weak cuticular teeth on the urosome; free

segment of leg 5 108 x 49 fi Af. myxicolae

Third segment of exopod of leg 4 with 11,1,5; strong cuticular teeth on the urosome; free

segment of leg 5 70 x 43 y. M. proximus

Myxomolgus myxicolae (Bocquet and Stock, 1958)

FIGURE 35

Paranthessius myxicolae Bocquet and Stock, 1958a, pp. 244-

253, figs. 1-4 [from the mucous tubes of the polychaete

Myxicola infundibulum (Renier), Penpoull, Finistere,

France].—Bocquet, Stock, and Kleeton, 1963, pp. 31-32

[between the layers of the mucous tube of Myxicola infundibulum, Roscoff, France, and Plymouth, England].

Myxomolgus proximus, new species

FIGURES 36-38

MATERIAL EXAMINED.—1 9 (holotype) embedded

between the mucous layers of the tube of the polychaete Myxicola aesthetica (Claparede), intertidal

in crevices of rocks, Locquirec, Finistere, France, 3

November 1955, collected by JHS. The holotype

is preserved in the Zoologisch Museum, Amsterdam (ZMA Co.102.218).

FEMALE—The body (Figure 36a) is slender. The

total length is 1.16 mm, without the ramal setae.

The first pedigerous segment is separated only by

a faint line from the cephalosome. The urosome

consists of five segments: the fifth pedigerous segment, the genital segment, and three postgenital

segments. Urosomal segments 2-4 are ventrally

provided at their posterior margins with 5-7 strong

sclerotized teeth (Figure 366), stronger than those

of Af. myxicolae (Figure 35a). The anal segment

bears ventrally, at its posterior margin, a row of

minute spinules on each side.

The caudal ramus (Figure 366) is 68 x 27 u,

distinctly longer than the anal segment. The lateral seta is implanted slightly proximal to the middle of the ramus. The four terminal setae are

smooth, the central two being very long.

The rostrum (Figure 36d) is elongated and triangular. Supporting structures, as in M. myxicolae,

have not been observed in the single specimen

available, but this point needs verification.

The first antenna (Figure 36c) is very similar to

that of M. myxicolae, both in segmentation and in

armature. The fifth segment in the new species is

slightly more slender than in M. myxicolae. The

second antenna (Figure 36c) is constructed as in

M. myxicolae, viz., with four segments, armed with

1 seta, 1 seta, 3 seta -f 1 daw, and 4 setae -f 3

B

FICUKE 35.—Myxomolgus myxicolae (Bocquet and Stock). Female: a, distal part of uroaome and

caudal rami (setae omitted), ventral (A) ; b, terminal portion of second maxilla (B) . Scale: A,

0.1 mm; B, 0.05 mm.
NUMBER 1X7 63

FIGURE 36.—Myxomolgus proxitnus, new species. Female: a, entire animal, dorsal (total length

1.16 mm) ; b, urosome, ventral (A) ; c, first antenna (c) ; d, rostrum (c); e, second antenna (D) ;

/, mandible (B) ; g, first maxilla (D) ; h, second maxilla (B) . Scale: A, 0.1 mm; B, C, D, 0.05 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 37.—Myxomolgus proximus, new species. Female: a, maxillipcd; b, endopod of leg 2; c,

endopod of leg 3; d, leg 4 and intercoxal plate. Scale: D, 0.05 mm, applies to all drawings.
NUMBER 127 65

claws respectively. In Af. proximus the fourth segment is about as long as wide, whereas in Af.

myxicolae it is much longer than wide. On the

other hand, the terminal claws in Af. proximus are

longer than the fourth segment, but in Af. myxicolae shorter than that segment.

The mandible (Figure 36/) is very similar to

that of the type-species. The first maxilla (Figure

36g) bears three elements, among them a small

medial setule not observed in Af. myxicolae.

The second maxilla (Figure 36A) differs in several respects from that of Af. myxicolae. The distal

lash is armed with elongatedly triangular teeth

(less elongate in Af. myxicolae); the most proximal

tooth is shorter than the second tooth (longer in

Af. myxicolae). A curiously toothed bladelike spine

arises near the base of the auxiliary lash; it bears

five strong teeth on one side and three smaller

teeth on the other. [In the original description of

Af. myxicolae a structure homologous to this is not

mentioned, but re-examination of the type material has revealed its presence (Figure 35&); it is

much smaller than in Af. proximus, and instead of

being toothed it is bicuspidate.] The maxilliped

(Figure 37a) corresponds rather closely with that

of Af. myxicolae.

Legs 1-4 (Figures $7b-d, 38a) show two distinguishing characters: the spiniform processes on the

outer side of the endopod (in particular in legs

1-3) are more strongly developed in Af. proximus;

and the chaetotaxis formula of the exopod of leg

4 is different in the two species—I—0;I—1;II,I,5 in

Af. proximus and I-0;I-l;II,I,4 in Af. myxicolae.3

The free segment of leg 5 (Figure 38&) is 70 n

long and, at a level just below the shortest distal

spine, 43 n wide; this segment is thus shorter, and

much less slender than the corresponding segment

* In the original description of M. myxicolae (Bocquet

and Stock, 1958a) the chaetotaxis formula of a somewhat

abnormal female was used for leg 1. Instead of I—0;I—1;II,I,3

for the exopod, one usually finds I-0;I-l;II,I,4, as has been

corrected by Bocquet, Stock, and Rleeton (1963, p. SI).

FIGURE 38.—Afyxomolgus proximus, new species. Female: a, leg 1 (D) ; b, leg 5 (c). Scale: c, D,

0.05
66 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in Af. myxicolae (108 x 49 v). The two distal elements in Af. proximus are 108 n and 95 n long, in

Af. myxicolae 135 ji and 97 j*. This means that

these two distal elements are more unequal in

length in Af. myxicolae than in Af. proximus.

Leg 6 (Figure 366) is represented by a seta and

a very short spine near the genital openings.

The male is unknown.

REMARKS.—This new species is closely related

to Af. myxicolae, and the hosts of these two copepods are closely related tubicolous polychaetes,

Myxicola infundibulum and Myxicola aesthetica.

The main differences between the two copepod

species are as follows: stronger cuticular teeth on

the urosome in Af. proximus; the less slender fourth

segment and the longer terminal claws in the second antenna of Af. proximus; the length of the

teeth and the structure of the proximal bladelike

spine on the second maxilla; the length of the

spiniform processes in legs 1-3; the different armature of the third exopod segment in leg 4; and the

length-to-width ratio in leg 5.

In our opinion these differences warrant the

creation of a new species for the present material,

although there is little doubt that the two species

of Myxomolgus are closely related. The proposed

specific name proximus (Latin, closely akin) alludes to this fact.

Genus Paranthessius Claus, 1889

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented; in the male 6-segmented.

Caudal ramus with six setae. Rostrum linguiform.

First antenna 7-segmented. Second antenna 4-segmented, with the formula 1, 1, 3, and three terminal claws -f- several setae.

Labrum incised medially. Mandible slender with

the basal region beyond the small indentation having on the concave side a row of spinules and on

the convex side an area with a few spinules followed by a serrated fringe; lash long. First maxilla

with three elements. Second maxilla of the lichomolgid type but the lash and the two auxiliary

elements about equal in length, forming a tripartite

tip. Maxilliped in the female 3-segmented; in the

male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami. Armature

lichomolgidifonn in pattern. Leg 4 exopod with

the third segment 11,1,5. Leg 4 endopod with the

formula 0—1;0—2;III. Leg 1 endopod in the male

with the third segment having 1,1,4 instead of 1,5

as in the female. Leg 5 with a free segment bearing two terminal elements.

Other features as in the species below.

Associated with actiniarians.

TYPE-SPECIES.—Paranthessius anemoniae Claus.

Paranthessius anemoniae Claus, 1889

FIGURE 39

Paranthessius anemoniae Clam, 1889, p. 17, pi. 4, figs. 8-15

[from the actiniarian Anemonia sp., Trieste, Italy].—Voigt,

1892, p. 35-Canu. 1894b. p. 1S7 [from Isles Chituey.

northern France]; 1898. pp. 415-415. pi. 10. figs. 1-8 [from

Isles Chausey. FranceJ.-Gadeau de Kerville. 1898, p. 436 -

Craeffe, 1900, p. 41 [from Anemonia sulcata (Pennant).

Trieste. Italy].-De Zulueta. 1912. pp. 12. 50.—Illg. 1949.

p. S97.-Bocquet and Stock. 1959b. pp. 44-53. fig*. 1-5

[from the actiniarian Anemonia sulcata (Pennant), Channel coast of France].-Bouligand. 1966. p. 269.

REMARKS.—This species is unusual among lichomolgidiform copepods in having two inner setae

(instead of one seta) on the second segment of the

fourth endopod.

Genus Sabelliphilus M. San, 1862

DIAGNOSIS.—Body cyclopiform, elongated. Urosome in the female 5-segmented; in the male 6-

segmented. Caudal ramus with six setae. Rostrum

bifid. First antenna 7-segmented, the first two segments enlarged. Second antenna 4-segmented, with

a toothed crest on the second segment, the formula

being 1, 1, one claw -f- 2 small setae, and four

terminal claws -f- several small setae.

Labrum incised medially. Mandible with area

beyond the indentation having fine spinules on the

concave side and coarse spinules along the convex

side, both rows continuing on the long lash. Paragnath a small hairy lobe. First maxilla with three

elements. Second maxilla of the usual lichomolgid

type. Maxilliped in the female 3-segmented with

a pointed tip; in the male 4-segmented (assuming

that the proximal part of the claw represents a

fourth segment).

Legs 1-4 with 3-segmented rami. Armature of

the basic lichomolgidifonn type. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod
NUMBER 127 67

f

FICUIIE 59.—Paranthe$sius anemoniae Claus. Female: a, dorsal; b, rostrum; c, second antenna;

d, mandible; e, second maxilla; /, endopod of leg 4. (From Bocquet and Stock, 1959b, figs, la.c,

2b, 3a,c. 5c.) Length of female 1.5-1.6 mm, of male 0.95-1 mm.

with the formula 0-l;0—1;II. Leg 1 endopod of the

male with the third segment having 1,1,4 instead

of 1,5 as in the female. Leg 5 with a free segment

bearing two terminal elements. Leg 6 represented

by the two setae near the genital openings.

Other features as in the species below.

Associated with polychaete annelids.

TYPE-SPECIES.—Sabelliphilus elongatus M. Sars.

Key to Species of the Genus Sabelliphilus

FEMALES

Body length 1.56-1.87 mm; two teeth of rostrum separated basally; crest on second segment

of second antenna with 4-10 spines (average 6); middle endopod segment of legs 1-4 with

one dentiform process S. elongatus

Body length averaging 0.90 mm (up to 1.17 mm); two teeth of rostrum with bases contiguous; crest on second segment of second antenna with 1-4 spines (average 2); middle

endopod segment of legs 1-4 with two dentiform processes S. tarsi

MALES

Body length 0.8-0.9 mm; first two postgenital segments with posteroventral denticulations

S. elongatus

Body length 0.60-0.90 mm; first two postgenital segments with posteroventral borders smooth

S.
68 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sabclliphilus elongatus M. San, 1862

Sabelliphilus elongatus M. San. 1862, pp. 1S9, 140 [from the

polychaete Sabella sarsii Kr0yer (=Sabella pavonina Savigny), Norway].-De Saint-Joseph, 1894, p. 266.-McIntosh, 1905, p. 86.-G. O. Sars, 1918, pp. 188, 189, pi. 107

[from Norway]-Lang, 1949, pp. 9, 10, figs. 12-15 (on the

gills of Sabella pavonia Savigny, Rddberget, Sweden].—

Rullier and Cornet, 1951, p. 48.-Bocquet, 1953, pp. 283-

286.—Gotto, 1954a, p. 133 [from Sabella pavonina, Ringhaddy Sound, Strangford Lough, County Down, Northern

Ireland]; 1960a, pp. 619-628, fig. 1, pi. 1 [from Sabella

pavonina, Ireland]; 1961a, p. 266; 1966, p. 194.—Bocquet

and Stock, 1957a, p. 427; 1958a, p. 243 [from the plume

of Spirographis spallanzanii Viviani and Sabella pavonina

Savigny, Penpoull, Finistere. France]; 1963, p. 295; 1964.

pp. 157-180, figs. 2, 4, 6, 9-22, 32, 37, 39, 46, 48-52, 58-67

[from the plume of Sabella pavonina, Spirographis spallanzanii, and Spirographis spallanzanii var. brevispira De

Quatrefages (=Sabella pavonina var. bicoronata Hornell),

Channel coast of France; also from Sabella and Spirographis at Banyuls, southern France].—Stock, 1960c, p. 248

[from the plume of the polychaete Spirographis spallanzanii, Banyuls, Mediterranean coast of France].—Breseiani

and Lutzen, 1962, p. 377.—Bocquet, Stock, and Kleeton,

1963, p. 21 .-Carton, 1966b, p. 816.-Crothers, 1966, p. 52

[from the branchiae of Sabella, Wales].—Glac.on, 1971, p.

23 (unnumbered).

Sabelliphilus sarsii.—Kossmann, 1877, pp. 16, 17, pi. S, fig. 2

[from Sabella, La Spezzia, Italy].—Canu, 1891b, pp. 436,

437 [from Sabella pavonina, Boulogne-sur-Mer, France].—

Thompson, 1893, p. 210, pi. 25, fig. 5 [from Sabella, Puffin Island, Liverpool Bay, England; synonym, Lichomolgus

sabellae, p. 210].

Sabelliphilus sarsii Claparede var. massiliensis Gourret, 1889,

pp. 475-477, pi. 34, figs. 1-6 [from the plume of Spirographis spallanzanii, Gulf of Marseille, France].

Sabelliphilus sarsi.-Canu, 1892, pp. 234, 235 [from Sabella

pavonina, Roches Bernard, Boulogne coast of France].—T.

Scott, 1897b, p. 196 [from Sabella, Clyde, Scotland]; 1897c,

p. 256 [from Sabella, Firth of Forth, Scotland]; 1905, p.

203 [from the plume of Sabella, Scotland]; 1906, p. 335

[from the plume of Sabella sp. (?S. pavonina), St. Monans, Firth of Forth, Scotland].

Sabelliphilus sarsi var. branchialis (Delia Valle).—Rioja,

1935, pp. 239-244, figs. 1-16 [from the plume of Sabella

pavonina, Bay of Pontevedra, near Marin, northwestern

Spain].

Lichomolgus sabellae Thompson, 1888a, pp. 32-33 [from

Sabella, Beaumaris, England]; 1888b, pp. 68, 69, pi. 2,

figs. 1-9 [from Sabella, Liverpool Bay, England]; in Herdman, 1888, p. 56 [from Sabella penicillus, Liverpool Bay,

England]; 1889, p. 182.-Herdman, 1890, p. 65.-De SaintJoseph, 1894, p. 266.

non Bispiraphilus tenax De Saint-Joseph, 1894, p. 292

[=Gastrodelphys].

REMARKS.—This species always lives on the

branchial crown of the polychaete host

Sabelliphilus sarsi Claparede, 1870

FIGURE 40

Sabelliphilus sarsii Claparede, 1870, pp. 10-18, pi. 7. figs.

1-8 [from the polychaete Spirographis spallanzanii Viviani, Naples, Italyj.-Claus, 1876. pp. 161-165. pi. 10

[from Spirographis spallanzanii, Naples, Italy]; 1889, pp.

14-15, pi. 1, figs. 8-10 [from Spirographis spallanzanii,

Trieste, Italy].-Stossich. 1880, p. 251 [on the skin of

Spirographis spallanzanii, Trieste, Italy].—Aurivillius, 1883.

p. 9.-Carus, 1885, p. 352.-De Saint-Joseph, 1894, p. 266.-

Graeffe, 1900, p. 41 [from Spirographis spallanzanii, Trieste,

Italy].

Sabelliphilus sarsii var. branchialis.—Carus, 1885, p. S52.

Sabelliphilus sarsi.-T. Scott, 1894b. p. 258-Mclntosh, 1905,

p. 86.-Bocquet, 1953, pp. 283-286.-Bocquet and Stock,

1957a, p. 427; 1958a, p. 243 [from the body of Spirographis

spallanzanii, Penpoull, Finistere. France]; 1963, p. 295;

1964, pp. 157-180. figs. 1, 3, 5. 7. 8. 23-36, 38, 40-45. 47.

53-57, 68, 69 [from the body of Spirographis spallanzanii,

Penpoull, northern France, and Banyuls, Mediterranean

coast of France].—Bocquet, Stock, and Kleeton, 1963, p.

21.-Bouligand. 1966, p. S02.-Carton. 1966a. pp. 427-442.

figs. 1, 2, tables 1. 2 [from Spirographis spallanzanii, Sabella pavonina, and Spirographis spallanzanii var. brevispira, Channel coast of France, and from Spirographis

spallanzanii var. bicoronata, Plymouth, England]; 1966b,

pp. 807 et seq.; 1967, pp. 387 et seq.; 1968a. pp. 123 et

scq.; 1968b. pp. 269 et seq.; 1968c. pp. 545 et seq.

Lichomolgus sarsii.-DeWa Valle, 1880a, pp. 87-97, pi. 5. fig*.

1-26 [from Spirographis spallanzanii, Naples, Italy]; 1880b,

pp. 110-118, pi. 1, figs. 1-26 [from Spirographis spallanzanii, Naples, Italy].—Lo Bianco, 1888, p. 405 [from Spirographis spallanzanii, Naples, Italy].

Lichomolgus sarsii var. branchialis Delia Valle, 1880a, p. 97

[from Spirographis spallanzanii, Naples, Italy]; 1880b, p.

117 [from Spirographis spallanzanii, Naples, Italy].

Lichomolgus (Sabelliphilus) sarsii.—Raffaele and Monticelli,

1885, p. 307.

REMARKS.—This species always lives on the body

of the polychaete host.

Genus Scambicornus Heegaard, 1944

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented; in the male 6-segmented.

Caudal ramus with six setae. Rostrum broadly

rounded or not well defined posteroventrally. First

antenna 7-segmented, the formula in the female

being 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete,

and 7 —f- 1 aesthete; in the male with a similar

formula or in S. lobulatus with an additional small

aesthete on the second segment. Second antenna

4-segmented, the fourth segment offset; formula 1,
NUMBER 127 69

FIGUUE: AO.-Sabelliphilus sarsi Claparedc. Female: a, dorsal; b, rostrum; c, first antenna; d,

second antenna (a fourth terminal claw is hidden); e, mandible; /, second maxilla; g, leg 4.

Male: h, dorsal. (From Bocquet and Stock, 1964, figs. 1, 3, 5, 7, 35, 38, 44, 47.) Length of female

0.90-1.17 mm, of male 0.60-0.68 mm.

1, 3 -\- one large claw, and 7, with all the terminal

elements being slender setae.

Labrum deeply incised medially. Paragnath a

small hairy lobe. First maxilla with four elements.

Second maxilla of the usual lichomolgid type.

Maxilliped in the female 3-segmented with a

pointed tip; in the male 4-segmented (if the proximal part of the claw is considered as the fourth

segment).

Legs 1-4 in the female with both rami 3-segmented. Leg 4 exopod with the third segment having 11,1,5. Leg 4 endopod with the formula

0-l;0-l;I,II,II. Legs 1 and 2 in the male with 3-

segmented exopods and 2-segmented endopods;

legs 3 and 4 with both rami 3-segmented. (In

S. armoricanus the endopods of legs 1—4 in the

male are 3-segmented; see below.) Distal segment

of leg 1 endopod of the male with 1,5,1 (though

the outermost seta on leg 1 endopod may be partly

feathered and partly barbed as in S. lobulatus);

spines on the distal endopod segments of male legs

2 and 3 strongly modified. Leg 5 with a free segment bearing two terminal setae. Leg 6 in the

female represented by the two setae and the process

near the genital openings, in the male by a ventral

posterolateral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated mostly with holothurians so far as

known.
7 0 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TYPE-SPECIES Scambicornus hamatus Heegaard. practical to prepare a key to the males of ScambiREMARKS.—The species Scambicornus armori- cornus, since in seven species the males are

canus is omitted from the following key because unknown and in several others the males are not

the female is unknown. We have thought it im- described in sufficient detail.

Key to Species of the Genus Scambicornus

KNOWN FEMALES

1. Leg 5 bell-shaped, much wider distally than proximally; genital segment with a pair of

dorsolateral toothlike protuberances S. campanulipet

Leg 5 subrectangular, not much wider distally than proximally; genital segment lacking

a pair of toothlike protuberances 2

2. Caudal ramus elongated, at least 4:1 3

Caudal ramus not more than 3:1 8

3. Claw on third segment of second antenna slender and setiform; fourth segment of second

antenna elongated 4

Claw on third segment of second antenna stout and unguiform; fourth segment of second

antenna short and almost quadrate 5

4. Caudal ramus 4:1 S. finmtnchicus

Caudal ramus 12:1 $. Umtkaudit

5. Caudal ramus between 43:1 and 5.5:1 6

Caudal ramus 7:1 S. peHti

6. Genital segment in dorsal view widest near middle, with sides smooth, not indented

5. prthtntUU

Genital segment in dorsal view widest posterior to middle, with sides indented 7

7. Body length 1.16 mm; ratio of length to width of prosome about 1.5:1; caudal ramus

about 5:1 S. terendibiau

Body length 1.23 mm; ratio of length to width of prosome about 1.18:1; caudal ramus

5.45:1 S. idonnu

8. Genital area with two short spiniform elements S. tuberahu

Genital area with two slender setae 9

9. Second antenna with claw on third segment rather slender and not strongly unguiform;

fourth segment of this appendage not offset 10

Second antenna with claw on third segment stout and distinctly prehensile; fourth segment

offset 11

10. Caudal ramus about 1:1 S. brevicauda

Caudal ramus 25:1 S. propinquw

11. Genital segment in dorsal view markedly expanded laterally, with a short slender posterior

part 12

Genital segment in dorsal view not much expanded laterally with gently rounded sides ... 17

12. Expanded sides of genital segment near genital openings rounded S. hamatus

Expanded sides of genital segment near genital openings angular 13

13. Body length 1.3-1.44 mm 14

Body length 0.6-1.18 mm 16

14. Mandible with a small terminal lash S. adduensis

Mandible without a terminal lash 15

15. Genital segment in dorsal view with sides tapering posteriorly; caudal ramus 2.9:1; free

segment of leg 5, 1.9:1 S. lobulatus

Genital segment in dorsal view with sides rounded; caudal ramus 25:1; free segment of

leg 5, 2.1:1 S. tubgrandu

16. Caudal ramus 2.7:1 S. modestus

Caudal ramus 1.67:1 S. rubtilis

17. Second antenna with fourth segment elongated, ratio about 3.8-4.0:1 18

Second antenna with fourth segment short, nearly quadrate 19

18. Caudal ramus 1.5:1; free segment of leg 5 about 2.25:1 S. nicobaricus

Caudal ramus 1.19:1; free segment of leg 5 with ratio of 1.65:1 S. pocuttferm

19. Caudal ramus 2:1; body length 1.1 mm S. rebuttut

Caudal ramus 1.6:1; body length 0.6 mm S. brachysetosus
NUMBER 127 71

Scambicornus ham at us Heegaard, 1944

FlCURE 41

Scambicornus hamatus Heegaard, 1944, pp. 360-366, figs.

1-10 [from the tentacles of the holothurian Thyonidium

alexandri Fisher, in 400 m, Okinose, Sagami Sea, Japan].—

Changeux, 1961. p. "..-Stock. 1964b, pp. 183-191. figs.

l-S.-Humes. 1967a. p. 144; 1969f, p. 92.

Scambicornus adduensis (Sewell, 1949)

Preherrmannella adduensis Sewell, 1949, pp. 85-89. figs.

20a-g, 21a-i [in weed-washings, Addu Atoll, Maldive

Archipelago, and Nankauri Harbor, Nicobar Islands].

Scambicornus adduensis.—Humes, 1967a, pp. 144-145; 1969f.

p. 92.

REMARKS.—The male is unknown.

Scambicornus armoricanus (Bocquet, Stock, and

Kleeton, 1963)

Preherrmannella armoricana Bocquet, Stock, and Kleeton,

196S, pp. S2-S7, figs. 6-9 [from the polychaete Polymnia

nebulosa (Mont.), Roscoff, northern France].

REMARKS.—The female is unknown. It is probable that this species belongs to a new genus. Legs

1-4 of the male have 3-segmented endopods, a

situation unlike any other Scambicornus. The fact

that this species is associated with a polychaete instead of with a holothurian suggests that it is different from Scambicornus. Clarification of the

position of this species, however, must await the

discovery of the female.

Scambicornus brachysetosus Reddiah, 1968

Scambicornus brachysetosus Reddiah, 1968c, pp. 126-131,

figs. 1, 2 [from the holothurian Holothuria atra Jaeger,

Kilakarai, Ramnad District, Madras State, India].

Scambicornus brevicauda (Sewell, 1949)

Preherrmannella brevicauda Sewell, 1949, pp. 82-85, fig.

19a-o [in weed-washings, Addu Atoll, Maldive Archipelago].—Ummerkutty. 1968, p. S14 [in weed-washings, Gulf

of Mannar, India].

Scambicornus brevicauda.—Humes, 1967a, p. 144; 1969f, p. 92.

e

g

FIGURE 41.—Scambicornus hamatus Heegaard. Female: a, dorsal; b, second antenna; c, mandible;

d, leg 4. Male: e, dorsal; /, endopod of leg 1; g, endopod of leg 2. (From Stock, 1964b, figs. la,c,e,

2a, Sd-f.) Length of female 1.13 mm, of male 0.97 mm.
72 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Scambicornus campanulipes (Humes and Cressey,

1961)

Preherrmannclla campanulipes Humes and Cressey, 1961a,

pp. 45-49, figs. 75-92 [from the holothurians Actinopyga

mauritiana (Quoy and Gaimard), Actinopyga echinites

(Jaeger), Actinopyga sp., and Halodeima atra (Jaeger),

and from the echinoid Diadema setosum (Leske), region

of Nosy Be\ Madagascar].

Scambtcornus campanulipes.—Humes, 1967a, pp. 148-150,

figs. 36-41 [from the holothurians Actinopyga lecanora

(Jaeger), Actinopyga mauritiana, Actinopyga miliaris

(Quoy and Gaimard), and Actinopyga echinites, region

of Nosy Be\ Madagascar]; 1969f, p. 92.

Scambicornus modestus.—Humes, 1967a, pp. 152-154, figs.

61-66 [from the holothurians Stichopus chloronotus, from

either Microthele difficilis (Semper) or Urodemas ehrenbergi Selenka, and Brandtothuria impatiens (Forskal),

region of Nosy Be1, Madagascar]; 1969f. p. 92.

Scambicornus nicobaricus (Sewell, 1949)

Preherrmannella nicobarica Sewell, 1949, pp. 89-91, fig.

22ag [in weed-washings, Nankauri Harbor, Nicobar Islands].

Scambicornus nicobaricus.-Humes, 1967a, p. 144; 1969f, p. 92.

REMARKS.—The male is unknown.

Scambicornus finmarchicus (T. Scott, 1903)

fHerrmannella finmarchica T. Scott, 1903b, pp. 28-30, pi.

4, figs. 14-19 [from Bog Fjord, East Finmark].

Herrmannella finmarchica.-G. O. Sars. 1918, pp. 179, 180,

pi. 101 [from Norway].

Scambicornus finmarchicus.—Humes, 1967a, p. 144; 1969f,

p. 92.

Scambicornus petiti (Stock and Klccton, 1963)

Preherrmannella petiti Stock and Klccton. 1963a, pp. 690-

696, figs. 5-8 [from the holothurian Stichopus regalis

(Cuvicr), Banyuls, Mediterranean coast of France, and

Cape Creus, Spain].

Scambicornus petiti.-Humet, 1967a, p. 144; 1969f, p. 92.

Scambicornus idoneus (Humes and Cressey, 1961)

Preherrmannella idonea Humes and Cressey, 1961a, pp.

26-34, figs. 1-27 [from the holothurians Halodeima atra

(Jaeger), Actinopyga echinites (Jaeger), and Holothuria

leucospilota (Brandt), and from the echinoid Diadema

setosum (Leske), region of Nosy Be\ Madagascar].

Scambicornus idoneus.—Humes, 1967a, p. 150, figs. 42-48

[from the holothurians Ludwigothuria (=Halodeima)

atra (Jaeger), Argiodia nobilis (Selenka), and Holothuria

scabra Jaeger, region of Nosy Be\ Madagascar]; 1969f,

p. 92.

NEW HOST.—2 <j> <j>, 1 cf from 15 specimens of

Holothuria edulis (Lesson), in 2 m, Ambariobe,

near Nosy B6, Madagascar, 25 May 1967, collected

by AGH.

Scambicornus poculiferus (Humes and Cressey,

1961)

Preherrmannella poculifera Humes and Cressey, 1961a, pp.

57-62, figs. 129-157 [from the holothurian Synapta maculata (Chamisso and Eysenhardt). Nosy Be". Madagascar].

Scambicornus poculiferus.-Humes, 1967a, pp. 150-152, figs.

49-60 [from Synapta maculata, region of Nosy B4, Madagascar]; 1969f, p. 92.

Scambicornus prehensilis (G. O. Sars, 1918)

Hermannella prehensilis G. O. Sars, 1918, p. 178, pi. 100

[from Norway].-Heegaard, 1944, pp. 361-363. fig. 11 (after

G. O. Sars, 1918).

Scambicornus prehensilis.—Humes, 1967a, p. 144; 1969f, p. 92.

REMARKS.—The male is unknown.

Scambicornus lobulatus Humes, 1967

Scambicornus lobulatus Humes, 1967a, pp. 136-148, figs.

1-25 [from the body surface of the holothurian Bohadschia

graeffei (Semper), region of Nosy Be", Madagascar].

Scambicornus propinquus (NichoIIs, 1944)

Paranthessius propinquus NichoIIs, 1944, pp. 53-54, figs.

23-24 [from South Australia].

Scambicornus propinquus.—Humes, 1967a, p. 144; 1969f,

p. 92.

Scambicornus modestus (Humes and Cressey, 1961) REMARKS.—The male is unknown.

Preherrmannella modesta Humes and Cressey, 1961a, pp.

39-43, figs. *)6-74 [from the holothurians Stichopus monotuberculatus (Quoy and Gaimard), Stichopus chloronotus

Brandt, Stichopus variegatus Semper, Holothuria pardalis

Selenka, and Bohadschia drachi Cherbonnier, region of

Nosy Be", Madagascar].

Scambicornus robustus (Thompson and A. Scott,

1903)

Hermannella robusta Thompson and A. Scott, 1903, p. 282,

pi. 17, figs. 1-8 [from washings of invertebrates, Ceylon].
NUMBER 127 73

Lichomolgus robustus.—Stock, 1957, p. 382.

Macrochiron robustus.-Stock, 1957, p. 382.

Scambicornus robustus.—Humes, 1967a, p. 144; 1969f, p. 92.

REMARKS.—The male is unknown.

Scambicornus serendibicus (Thompson and A.

Scott, 1903)

Hermannella serendibica Thompson and A. Scott, 1903, pp.

282-283, pi. 17, figs. 9-11 [in washings of sponges, Gulf of

Mannar, Ceylon].

Scambicornus serendibicus.—Humes, 1967a, p. 144; 1969f,

p. 92.

Preherrmannella serendibica.—Ummerkutty, 1968, p. 314 [in

weed washings, Gulf of Mannar].

REMARKS.—The male is unknown.

Scambicornus subgrandis (Humes and Cressey,

1961)

Preherrmannella subgrandis Humes and Cressey, 1961a, pp.

53-57, figs. 110-128 [from the holothurian Holothuria rugosa Ludwig, region of Nosy B6, Madagascar].

Scambicornus subgrandis.—Humes, 1967a, p. 144; 1969f, p. 92.

Scambicornus subtilis (Humes and Cressey, 1961)

Preherrmannella subtilis Humes and Cressey, 1961a, pp.

49-53, figs. 93-109 [from the holothurians Halodeima

edulis (Lesson), Opheodesoma grisea (Semper), and Holothuria curiosa Ludwig, Nosy B£, Madagascar].

Scambicornus subtilis.—Humes, 1967a, p. 144; 1969f, p. 92.

Scambicornus tenuicaudis (G. O. Sars, 1918)

Hermannella tenuicaudis G. O. Sars, 1918, pp. 180, 181, pi.

102 [from Norway].

Scambicornus tenuicaudis.—Humes, 1967a, p. 144; 1969f,

p. 92.

REMARKS.—The male is unknown.

Scambicornus tuberatus (Humes and Cressey, 1961)

Preherrmannella tuberata Humes and Cressey, 1961a, pp.

34-39, figs. 28-55 [from the holothurians Bohadschia sp.,

Bohadschia koellikeri (Semper). Thelenota ananas

(Jaeger), Opheodesoma grisea (Semper), and Bohadschia

cousteauxi Cherbonnier, region of Nosy Bi, Madagascar].

Scambicornus tuberatus.—Humes, 1967a, p. 154, figs. 67-73

[from the holothurian Bohadschia marmorata Jaeger, region of Nosy Be\ Madagascar]; 1969f, p. 92.

NEW HOST.—3 $ $ , 2 d*o*. and 1 copepodid

from a single specimen of Bohadschia vitiensis

(Semper), in 17 m, in pass between Nosy Komba

and Nosy Be1, Madagascar, 10 August 1967, collected by AGH.

Genus Serpuliphilus Humes and Stock, 1972

DIAGNOSIS.—Body cydopiform. Urosome in the

female 5-segmented; in the male 6-segmented.

Caudal ramus with six setae. Rostrum broadly

rounded. First antenna 7-segmented, slender. Second antenna 4-segmented, with two strong claws

on the fourth segment.

Mandible having on its convex side a serrated

fringe and on its concave side a row of spinules;

tapering gradually into a relatively short lash. First

maxilla with three elements. Second maxilla of

the usual lichomolgid type. Maxilliped in the female 3-segmented and prehensile, but the last

segment very long and clawlike; in the male 4 -

segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami. Armature

lichomolgidiform in pattern. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0—1;0—1;II. Leg 1 endopod with

the same formula in both sexes. Leg 5 with a free

segment bearing two terminal elements. Leg 6 represented by two setae near the genital openings.

Other features as in the species described below.

Associated with serpulid polychaetes.

TYPE-SPECIES Serpuliphilus tenax, new species.

ETYMOLOGY.—The generic name is compounded

from Serpulidae and qaXos, loving. Gender masculine.

REMARKS.—It is not possible to construct a key

for the males of Serpuliphilus, since the male of

5. duplus is unknown.

Key to Species of the Genus Serpuliphilus

Caudal ramus about 13 times as long as wide; maxilliped with a long slender daw, resembling

that of a male *• tenax

Caudal ramus about 4 times as long as wide; maxilliped reduced with a short terminal claw

S. duphu
74

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Serpuliphilus tenax, new species

FIGURES 42, 43, 44a

MATERIAL EXAMINED.—All from the gills of the

sedentary polychaete Spirobranchus giganteus (Pallas): Curasao: 22 9 9 , 36 o* cf. and 19 copepodids

(including $ holotype and allotype) from eight

polychaetes, depth 1-2 m, Piscadera Bay, 10 October 1958, deposited in ZMA (Co. 102.274A), collected by JHS; 9 9 9, 7 cf cf, and 12 copepodids

from one polychaete, depth 2 m, same locality, 14

October 1958, collected by JHS.

Bonaire: 21 9 9 , 48 cfcf. and 1 copepodid,

depth 0-1 m, pier of Kralendijk, 28 December

1958, collected by JHS.

Puerto Rico: 2 cf cf from one polychaete, in 1.8

m, on edge of lee side of Margarita Reef, near La

Parguera, 8 August 1959; 13 cf cf from 9 polychaetes, Terremoto Reef, southeast of Magiieyes

Island, near La Parguera, 9 August 1959; 20 9 9 ,

39 cf cf from 14 polychaetes, Laurel Reef, south

of Magiieyes Island, 13 August 1959. All collected

by AGH and R. U. Gooding (RUG).

Jamaica: 1 9. 1 cf» and 4 copepodids from three

polychaetes, in 15 m, southeastern corner of Maiden Cay, off Kingston Harbor, 6 September 1959;

1 9» ^ cf cf» and 1 copepodid from 3 polychaetes,

Rackham's Cay, off Kingston Harbor, 7 September 1959. Collected by AGH and RUG.

Barbados: More than 50 specimens from 20

polychaetes, depth 90-180 cm, off Six Men's Bay,

St. Peter, 30 June 1959; 7 9 9 , 6 cf cf from five

polychaetes, 120-180 cm, reef off St. James Church,

St. James, 30 June 1959; 9 9 9, 4 cf o \ on wreck,

Carlisle Bay, 6 July 1959. Collected by AGH and

RUG.

Bahamas: 23 9 9 > 6 cf cf» and 3 copepodids, on

wreck, three miles south of South Bimini, 8 June

1959; 5 9 9» ^ cf cf» and 4 copepodids from one

polychaete, west of southern end of Alec Bay,

Bimini Lagoon, 9 June 1959. Collected by AGH

and RUG.

The description and figures are based on a single

female and a single male from Curacao, except

for Figure 44a, which was made from a Puerto

Rican specimen. The polychaetes from Curacao

and Bonaire were identified by Dr. E. WesenbergLund, those from Puerto Rico, Jamaica, Barbados,

and the Bahamas by Dr. Marian E. Pettibone.

FEMALE.—The body form (Figure 42a) is that

of a generalized Herrmannella. The size, not including the setae on the caudal rami, is 0.85 mm

(0.81-0.87 mm) x 0.39 mm (0.37-0.40 mm), based

on five specimens.

The first pedigerous segment is fairly clearly demarcated from the cephalic segment. It has the

same width as the cephalic segment; pedigerous

segments 2 and 3 diminish regularly, but slightly,

in width; segment 4 is suddenly much narrower,

and segment 5 narrower again. The genital segment (Figure 426) has a larger diameter than the

fifth pedigerous segment, its greatest width being

reached at the level of two roughly triangular lateral expansions that carry the genital openings on

their posterior margin. The genital segment carries one spinelike seta and a flexible setule near

the sexual openings. The egg sacs (Figure 42a)

are elongate, reach to the tip of the setae on the

caudal rami, and contain rather numerous eggs in

multiserial arrangement. The size of the egg sac

is 586 n (515-612 M) X 121 n (113-137 M). A spermatophore found attached to the genital segment

measured 98 x 47 n.

The total number of urosomal segments is five.

The anal segment does not carry spinules. The

caudal ramus (Figure 426) is 24 x 18 » to 32 x

23 n, thus about 1.5 times as long as wide; its

lateral seta arises near the middle of the outer margin. The lateral seta, the four terminal setae, and

the dorsal seta are smooth and not transformed.

The rostral fold (Figure 43/) is a flat, rounded

lobe, not accompanied by auxiliary structures.

The first antenna (Figure 42c) is composed of

seven distinctly articulated segments. The third

segment bears a trace of a subdivision, but is certainly not subdivided into two articulated units.

The segments bear 4, 14, 5, 3, 4, 2, and 7 setae

respectively; in addition, each of segments 5-7

bears an aesthete.

The second antenna (Figure 42d) is very stout.

It consists of the usual four segments, but the third

segment is reduced to a narrow, triangular wedge

carrying three normal setae and a longer seta with

a flattened tip. The fourth segment is armed with

three lateral setae, one long outer terminal seta,

one very short and one long inner terminal seta,

and two unusually powerful, curved claws.

The mandible (Figure 42e) has the median edge

of the lappet finely serrated, the lateral edge armed
NUMBER 127

FIGURE 42.-Serputiphilus tenax, new species. Female: a, dorsal (A) ; b, urosome, ventral (B) ; e,

first antenna, with an arrow indicating the position of an additional aesthete in the male (c); d,

second antenna (D) ; e, mandible (F) ; /, first maxilla (E) ; g, second maxilla (E); h, maxilliped

(D) . Scale: A, 02 mm; B, 0.1 mm; c, D, 0.05 mm; E, 0.02 mm.
76

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE ii.—Serpuliphilus tenax, new species. Female: a, leg 1 (c); b, second and third segments

of endopod of leg 2 (c); c, leg 3 and intercoxal plate (c); d, leg 4 (c); e, leg 5 (E) ; /, rostrum,

ventral (B) . Male: g, urosome, ventral (B) ; h, maxilliped (D) . Scale: B, 0.1 mm; c, D, 0.05 mm;

E, 0.02 mm.
NUMBER 127 77

with long, fine spinules. The apical lash is relatively short.

The first maxilla (Figure 42/) is a simple lobe

armed with three short terminal setae.

The second maxilla (Figure 42g) bears two articulated elements: a smooth spine and, more distally, a large, feathered, annulated seta. The terminal lash bears numerous regular serrations on its

median edge.

The maxilliped (Figure 42h) has attained an

unusual development for this family and this sex.

It is a slender, prehensile organ consisting of a

smooth basal segment, a tapering second segment,

and a slender claw. The second segment bears a

conical projection on the outer side and two large

spines and a row of minute spinules on the inner

side.

Legs 1-4 (Figures 43a-d) are biramous, each

ramus being 3-segmented. The spinal and setal

formula of die rami is as follows:

1*.

p,

p.

The endopod of leg 4 is slender and weak compared to the exopod.

The fifth leg (Figure 4Se) consists of a single

segment 19 x 11 n. The two terminal setae are

smooth.

MALE.—The size (based on five specimens) is

0.76 mm (0.74-0.79 mm) x 0.29 mm (0.29-0.31

mm). Apart from the configuration of the urosome, of the first antenna, and of the maxilliped,

there are no structural differences between the two

sexes. The genital segment is of the normal,

roughly circular shape (Figure 43g); four urosomal

segments follow after the genital segment, against

three in the female. The first antenna is armed

and constructed as in the female, but has an extra

aesthete on segment 4, implanted near the most

proximal seta. The male maxilliped (Figure 43/J)

has a stronger claw, which is armed, moreover,

with two basal setae; the spinules in the row on

segment 2 are larger than in the female. A very

slight sexual difference exists in the second antenna

vxp

enp

exp

enp

exp

cup

exp

enp

I-O;

0-1;

1-0;

0-1;

1-0;

0-1;

1-0;

0-1;

1-1;

0-1;

i-l;

0-2;

I-l;

0-2;

I-l;

0-1;

MI.I.4

13

111.1,5

1,11,3

IH.I3

1,11,2

11,1,5

II

where the terminal seta is as long as the claw in

the male, but shorter than the claw in the female.

COLOR.—In the Curasao specimens the body is

clear as glass, the eye is red, the intestine is ivorycolored. The two egg sacs are white with a slightly

pinkish hue. Other females, from the Bahamas,

have brown or white reproductive organs and a

brown gut. A female from Puerto Rico was colorless except for a bright red eye and a very slight

rose tint on the thoracic appendages and the genital

segment.

ANOMALOUS STRUCTURE.—A female specimen collected at Terremoto Reef, Puerto Rico, shows a

curious anomaly. On the exopod of the first leg

(Figure 44a) one of the setae is replaced by an

additional small segment which carries two short,

plumose terminal setae. The corresponding leg on

the other side of the body is normally built.

AFFINITIES.—Within the Sabelliphilidae, the present species stands well isolated by the presence of

two strong claws on segment 4 of the second antenna, whereas segment 3 of the second antenna is

devoid of claws. Also, the great development of

the female maxilliped is a most remarkable feature

of this new species.

ETYMOLOCY.—The specific name tenax (Latin,

tenacious) alludes to the strong development of

the antennal and maxillipedal claws.

Serpuliphilus duplus, new species

FIGURES 446-/, 45

MATERIAL EXAMINED.—From the gills of the

sedentary polychaete Pomatostegus stellatus

(Abildg.): Curasao: ovigerous <j>, the holotype

(ZMA Co. 102.274), from a polychaete fixed on a

coral in about 2 m depth, Santa Martha Bay, 12

October 1958, collected by JHS. Puerto Rico: 1 $

from five polychaetes, depth about 0.5 m, landing

at Magiieyes Island, off La Parguera, 4 February

1963, collected by JHS.

FEMALE (holotype).—The body (Figure 446) is

fairly slender; the first pedigerous segment is

vaguely separated from the cephalic segment. The

size, not including the setae on the caudal rami, is

0.76 x 0.31 mm. The metasome segments diminish

regularly in width. The genital segment has a

characteristic shape (Figure 44c); the total number

of urosomal segments is five; the anal segment is
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FICURE 44.—Scrpuliphilus tenax, new species. Female: a, abnormal exopod of leg 1 in a specimen

from Puerto Rico (D). Scrpuliphilus duplus, new species. Female: b, dorsal, with egg sac

(F) ; c, urosome, ventral (B) ; d, rostrum, ventral (B) ; e, first antenna (c); /, second antenna

(D) ; g, mandible (E) ; h, first maxilla (E) ; i, second maxilla (E) ; ;, maxilliped (D) . Scale: B,

0.1 mm; c, D, 0.05 mm; E, 0.02 mm; F, 0.2 mm.
NUMBER 127 79

smooth. Two setae arise near each genital opening.

The egg sacs are elongated elliptical in outline,

measure 451 x 129 n, and contain numerous eggs.

The caudal ramus (Figure 44c) is 58 x 14 u, thus

about four times as long as wide, much longer

than the anal segment (40 u). The lateral seta

arises somewhat distally to the middle of the outer

margin of the ramus. All setae are smooth and not

transformed.

The rostral fold (Figure 44d) is a weakly chitinized, broad lobe, not beak-shaped or bifid as in

many other sabelliphilid species.

The first antenna (Figure 44c) is composed of

seven distinct segments; the third segment, however, bears a trace of a subdivision which is not a

functional articulation. The number of setae on

the segments is 4, 12, 5, 3, 4, 2, and 7 respectively.

Moreover, each of segments 5-7 carries an aesthete.

The second antenna (Figure 44/) roughly resembles that of S. tenax. The second segment bears

one seta and three chitinous wrinkles or folds. The

third segment is small, rectangular, and carries

three normal setae and a longer, flattened seta. The

fourth segment has three lateral setae and a termiFIGURE 45.—Serpuliphilus duplus, new species. Female: a, leg 1 (c) ; b, third segment of endopod

of leg 2 (D) ; c, leg 3 (c); d, leg 4, with detail of basis on opposite side of body (c); e, leg 5

(E) ; /, caudal ramus in a specimen from Puerto Rico (c). Scale: c, D, 0.05 mm; E, 0.02 mm.
80 SMITHSONIAN CONTRIBUTIONS 1O ZOOLOGY

nal armature consisting of one long outer seta, one

long and one very short inner seta, and one robust

and one feebler claw.

The mandible (Figure 44g) and first maxilla

(Figure 44h) resemble those of S. tenax. The second maxilla (Figure 44*) bears two projections:

one on the anterior distal corner of the first segment and one on the adjacent corner of the second

segment. A setule arises near the projection on

the second segment. The terminal lash is armed

with regular, slender teeth.

The maxilliped (Figure 44;) consists of two segments and a terminal claw. The second segment

bears two spines and a row of spinules; the claw

bears one short spine. The entire appendage is in

a more reduced condition than in S. tenax.

The spine and setal formula of legs 1-4 (Figure

45a-d) is identical with that of 5. tenax. There

exist, however, certain differences in proportions,

which are most clearly visible in the greater elongation of the third segment and in the much longer

terminal spine on this segment in S. duplus.

The fifth leg (Figure 45?) is similar to that of

S. tenax, the free segment being 20 x 8 ji.

The female from Puerto Rico is slightly longer

than the Curacao specimen, viz., 998 n. It has much

slenderer caudal rami (Figure 45/), 83 x 14 n,

whereas the anal segment is 50 n long. In its other

characters, however, it resembles fully the Curacao

specimen.

MALE.—Unknown.

COLOR.—The body is colorless; the eye is bright

red; the gut is pink; the two egg sacs are salmonpink.

AFFINITIES.—Like S. tenax, the present species is

chiefly characterized by the presence of two strong

claws on the fourth segment of the second antenna.

It differs from tenax, however, in a number of characters, the most readily visible being the more reduced female maxilliped, the much greater

elongation of the caudal rami, and the greater development of the terminal spine of the third endopod segments.

ETYMOLOGY.—The specific name duplus (Eatin,

the double number) refers to the presence of two

terminal claws on the second antenna.

Genus Synapticola Voigt, 1892

DIAGNOSIS.—Body transformed, elongated. Urosome in the female 5-segmented; in the male

6-segmented. Caudal ramus with three setae. Rostrum not developed. First antenna 7-segmented.

Second antenna 3-segmented, terminally with a

claw and a seta.

Labrum a thin plate notched in the middle.

Mandible sickle-shaped and with the convex side

of the broad blade having fine teeth; tip of the

blade with a curved and rounded thickening. First

maxilla with three elements. Second maxilla of

the usual lichomolgid form. Maxilliped in the

female 3-segmented and with a pointed tip; in the

male 4-segmemed (assuming the proximal part of

the claw to be a fourth segment).

Legs 1-4 in both sexes with 3-segmented exopods

and 2-segmented endopods. Leg 5 in both sexes a

short papilla bearing two setae, with a third seta

nearby.

Endoparasitic in holothurians.

TYPE-SPECIES Synapticola teres Voigt.

Synapticola teres Voigt, 1892

KM.IR K 4 6

Synapticola teres Voigt, 1892, pp. 32-40, pi. 5, figt 1-25

[from the body cavity of the holothurian Synapta kefersteini Selenka, Amboina].—Changeux, 1960, p. 55.

REMARKS.—Voigt's description, based on two

females and three males, is far from complete. 5.

FIGURE 46.—Synapticola teres Voigt. Female: a, outline of

body, dorsal; b, lateral; c, mandible; d, maxilliped. (Redrawn

after Voigt, 1892, pi. 5, figs. 6, 11 (part), 15, 22.) Length of

female 3 mm, of male 1J> mm.
NUMBER 127 81

teres has not been found since, and the genus remains monotypic. Hence, the characterization of

the genus is at present deficient and its completion must await the finding of new specimens. On

the basis of Voigt's information, however, we consider it to be a valid genus.

Genus Thamnomolgus Humes, 1969

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six relatively short setae. Rostrum with a broadly truncated posteroventral margin. First antenna 7-segmented, the armature in

the female being 3, 13, 6, 3, 4 -f- 1 aesthete, 2 -j1 aesthete, and 7 -f- 1 aesthete; in the male three

aesthetes added, so that the formula is 3, 13 -|- 2

aesthetes, 6, 3 + 1 aesthete, 4 -|- 1 aesthete, 2

-f 1 aesthete, and 7 -J- 1 aesthete. Second antenna

4-segmented, with the formula 0, 1, 3, I -f- a

minute setuliform element, there being one terminal claw.

I..iIniim sii!xju.it11.He, its posteroventral margin

only slightly indented and without the usual deep

median cleft. Mandible a simple blade without a

proximal constriction. Paragnath a hairy lobe.

First maxilla with two setae. Second maxilla 2-

segmented, the small second segment with a slender

blade corresponding to the lash seen in related

genera. Maxilliped in the female 3-segmented; in

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami and with the

same formula for the armature in both sexes. Leg

3 endopod with the formula 0-l;0-2;II,2. Leg 4

endopod with 0—1 ;0—1;II. Strong sexual dimorphism on the third segment of leg 1 endopod. Leg

5 in the female with a free segment bearing two

terminal setae; in the male this segment reduced

to a small lobe fused with the body but bearing

two terminal setae. Leg 6 in the female represented by a single small seta near the area of attachment of each egg sac, in the male by a posteroventral flap on the genital segment extended as a

subconical process bearing two small setae.

Other features as in the species below.

Associated primarily with antipatharians.

TYPE-SPECIES.—Thamnomolgus robustus Humes.

Thamnomolgus robustus Humes, 1969

FICURE 47

Thamnomolgus robustus Humes, 1969b, pp. 19-28, figs.

64-97 [from the antipatharians Antipathes myriophylla

Pallas, Antipathes sp. cf. spinescens Gray, Antipathes abies

(Linnaeus), and Antipathes ericoides Pallas, region of

Nosy lie, northwestern Madagascar].

NEW RECORD.—1 tf from the gorgonian Acanthogorgia aspera Pourtales, in 24 m, north of Ankazoberavina, near Nosy Be", Madagascar, 13°27.6' S,

47°58.2' E, 25 August 1967, collected by AGH. This

specimen, which occurred along with 390 adults

and copepodids of Acanthomolgus astrictus, probably is an accidental rather than a regular associate

of the gorgonian.

Sabclliphilidae Insufficiently Described or of

Uncertain Position

I) or id kola antheae Ridley, 1879

Doridicola antheae Ridley, 1879, p. 458 [from the tentacles

of the actiniarian Anthea cereus, var., in a pool at low

water, Ilfrecombe, North Devon, England].

Lichomolgus antheae.—De Zulueta, 1912, p. 11. [Anemonia

sulcata (Pennant) = Anthea cereus.]

REMARKS.—Ridley's species may be Paranthessius

anemoniae Claus, 1889. The male is unknown.

Herrmannella dubia G. O. Sars, 1921

Hermannella dubia G. O. Sars, 1921, pp. 110, 111, pi. 73,

fig. 1 [from Norway].

REMARKS.—Similar in some ways to Paranthessius

anemoniae. Probably belongs to Paranthessius according to Sewell, 1949, p. 81. The female is unknown.

Paclabius tumidus Kossmann, 1877

Paclabius tumidus Kossmann, 1877, pp. 23, 24, pi. 6, figs.

1-10 [from the pericardium of the bivalve Tridacna, Bohol, Philippine Islands; belongs in Lichomolgidae according to unpublished work of H. E. H.].—Aurivillius, 1883,

p. 3.

Paranthessius sp.

Paranthessius sp.-K6, 1961, pp. 130, 131, 138, fig. lc [from

the bivalves Mactra sulcataria Reeve and Tapes japonica

Deshayes, Japan].
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 47.—Thamnomolgus robustus Humes. Female: a, dorsal; b, mandible; c, leg 4 and intcrcoxal plate; d, leg 5. Male: e, dorsal; /, leg 5. (From Humes, 1969b, figs. 64, 74, 85-85, 96.)

Length of female 1.21 mm, of male 1.05 mm.

Sabelliphilus (?) bispirae Mclntosh, 1905

Sabelliphilus (?) bispirae Mclntosh, 1905, pp. 86-91, pi. 9,

figs. 51-56 [from the polychaete Bispira volutacarnis Montagu, South Africa].—Barnard, 1955, p. 233, fig. 5a-c [from

the polychaete Bispira volutacornis, Cape, South Africa].

Sabelliphilus leuckarti Kossmann, 1877

Sabelliphilus leuckarti Kossmann, 1877, pp. 17, 18, pi. 2,

figs. 2, 3 [from the polychaete Sabella, Massaua, Red Sea].

Family LICHOMOLGIDAE Kossmann, 1877

Sapphirinidae Thorell, 1860, p. 64.—Brady, 1880, p. 39.—

Cams, 1885, p. 350 (Section b, Lichomolgina).

Lichomolgidae Kossmann, 1877, p. lO.-Canu, 1891a, p. 477.—

G. O. Sars, 1917a, pp. 149, 150.

Lichomolgiden Claus, 1889, pp. 1, 19.

Lichomolginae Gurney, 1927, p. 463-Sewell, 1949, p. 91.

Legs 1-4 usually with all rami 3-segmented except

the endopod of leg 4 which is 1- or 2-segmented.

Reduction occurs in a posterior to anterior series,

as in Rakotoa with a vestigial leg 4 endopod, Ravahina with endopods of legs 3 and 4 vestigial, and

Amarda with endopods of the first two legs 2- or

3-segmented, endopods of legs 3 and 4 absent (in

this genus leg 3 exopod is 2-segmented and leg 4

exopod is absent). Leg 5 present in both sexes

and with a free segment (in a few genera—for

example, Octopicola—not clearly delimited from

the body).

REMARKS.—Two genera, Philoconcha and Paraphiloconcha, have been omitted from the following

key due to lack of information in the existing

descriptions.
NUMBER 127 8 3

Key to the Described Genera of the Family Lichomolgidae

1. Second antenna 3-segmented 2

Second antenna 4-segmented 11

2. Legs 1 and 2 with 3-segmented exopods and 2- or 3-segmented endopods; leg 3 with

2-segmented exopod, endopod absent; leg 4 absent Amarda

Legs 1-3 with 3-segmented rami; leg 4 with 3-segmented exopod and 1- or 2-segmented

endopod 3

3. Rostrum with slender, needle-like process; claws on second antenna pectinate Macrochhon

Rostrum rounded, triangular, or broadly truncated but lacking a needle-like process; claws

on second antenna not pectinate 4

4. Leg 4 with endopod 1-segmented 5

Leg 4 with endopod 2-segmented 6

5. With broad shield-shaped prosome; second antenna with two terminal claws; leg 4 with

endopod armed with 11,1 Aspidomolgus

Prosome not unusually broad; second antenna with one terminal claw; leg 4 with endopod

armed with II Sewellochiron

6. Second antenna with two or three terminal claws Aitericola

Second antenna with one terminal claw 7

7. Leg 4 with endopod having formula O-1;II 8

Leg 4 with endopod not thus armed 9

8. Mandible with basal area distal to indentation having on its convex side four slender

digitiform processes; first segment of first antenna with four setae Schedomolgus

Mandible with basal area distal to indentation having on its convex side a large hyaline

expansion; first segment of first antenna with one seta Spaniomolgus

9. Leg 4 with endopod having formula 0-1 ;I; mandible with basal area distal to constriction

having on its concave side two lobes with serrated margins Prionomolgus

Leg 4 with endopod having more than one element on second segment; mandible with

basal area distal to constriction having on its concave side a row of spinules 10

10. Urosome in female 5-segmented, in male 6-segmented but with segment of leg 5 fused

with genital segment; basis of leg 4 laterally elongated Stellicola

Urosome in female 4-segmented, in male 5-segmented (though in both sexes there may

be a slight indication of division of anal segment); basis of leg 4 not laterally

elongated SynsUUicola

11. Legs 1—4 with rami 3-segmented except for endopod of leg 4 or endopods of both legs 3

and 4 which are vestigial and represented only by a small unarmed knob 12

Legs 1-4 with all rami 3-segmented except endopod of leg 4 which is either 1- or 2-

segmented 13

12. Only leg 4 with vestigial endopod; legs 1-4 with inner coxal seta; second maxilla with

long digitiform process on first segment Rakotoa

Both legs 3 and 4 with vestigial endopod; legs 1-4 lacking inner coxal seta; second maxilla

without a long process on first segment Ravahina

13. Leg 4 with endopod 1-segmented 1.4

Leg 4 with endopod 2-segmented 20

14. Fourth segment of second antenna with four terminal claws; leg 5 a minute lobe; body

elongated with slender prosome Octopicola

Fourth segment of second antenna with less than four terminal claws; leg 5 with a

distinct free segment; body cyclopiform, prosome not unusually slender 15

15. Endopod of leg 4 unarmed Haptomolgus

Endopod of leg 4 armed 16

16. Second antenna with a claw on third segment in addition to either one or two terminal

claws 17

Second antenna without a claw on third segment 18

17. Second antenna with one terminal claw; leg 4 with endopod having two spines

Lichomolgella

Second antenna with two terminal claws; leg 4 with endopod having two spines and a

seta Kelleria

18. Leg 4 with endopod armed with two spines and a seta Telestacicola

Leg 4 with endopod armed with only two spines 19
8 4 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

19. Leg 4 with endopod in female almost as long as exopod in male, much shorter than exopod;

second antenna with one terminal claw Paranuurochiron

Leg 4 with endopod in both sexes much shorter than exopod; second antenna with two

terminal daws (except one clawlike npine in P. fuckolum) Pseudomacrochiron

20. Leg 4 with second segment of endopod bearing only one element 21

Leg 4 with second segment of endopod bearing more than one element 24

21. Second antenna with one terminal claw 22

Second antenna with three terminal claws 23

22. Leg 4 with endopod having formula 0-1 ;I; second maxilla with long digitiform process on

first segment AndrimteUus

Leg 4 with endopod having formula 0-0;I; second maxilla without such a process on first

segment Monomo^pu

23. Body modified with prosome in female pointed anteriorly; leg 4 with endopod having formula 0-1; 1; mandible a broad blade abruptly attenuated distally; third exopod segments

of legs 1-4 with only three spines telMtomotgut

Body cyclopifonn with prosome in female rounded anteriorly; leg 4 with endopod having

formula 0-1 ;I; mandible with broad base and slender attenuated blade; third exopod

segments of legs 1-4 with four spines ttobrumm

24. Leg 4 with second segment of endopod bearing more than two elements 25

Leg 4 with second segment of endopod bearing two elements 26

25. Leg 4 with endopod having formula O-1;II,I; mandible with slender base merging into long

attenuated lash; leg 4 with third exopod segment having armature 11,13 Atddioxynut

Leg 4 with endopod having formula 0-1:11,3, or 0~l;II.2. or 0-1:1.5; mandible with large

basal area indented; leg 4 with third exopod segment having armature 111,1,4, or III,

1,5, or 1,1X5 Indomotgui

26. Leg 4 with endopod having first segment unarmed Xenomolgui

Leg 4 with endopod having an inner element on first segment 27

27. Labrum with a pair of prominent, ventrally directed anterolatcral setae; leg 4 with endopod

having formula 0-1 ;2; maxilliped in female slender, with greatly elongated third segment

Ntuomolgui

Labrum without such setae; leg 4 with endopod having different formula; maxilliped in

female not elongated 28

28. Leg 4 with endopod having formula 0-1; 1,1 Anhomolgus

Leg 4 with endopod having different formula 29

29. Leg 4 with endopod having formula 0-I;II SO

Leg 4 with endopod having formula 0-1 ;II 31

30. Second antenna with one terminal claw; second maxilla in male with large proximally

directed seta Meringomolgus

Second antenna with two terminal claws; second maxilla in male without such a seta

Aconthomolgus

31. Second antenna with one claw on third segment and three or four terminal claws 32

Second antenna without claw on third segment but with 1-4 terminal claws 34

32. Second antenna with three terminal claws; mandible with toothlike process on convex

edge Acaenomolgus

Second antenna with four terminal claws; mandible without toothlike process on convex

edge 33

33. Female with prosome strongly inflated, and with a median ventrally produced poetoral

structure terminating in two small hooks Lichomolgidts

Female with prosome not inflated and without such a hooked structure Zygomolgus

34. Second antenna terminally with one claw and one clawlike spine Epbnolgus

Second antenna terminally with different armature 35

35. Mandible of simple type, with slender base merging gradually into long, attenuated lash. .

Mchomolgus

Mandible of more complex type, with large base often indented on convex side and variously ornamented; lash variable, very short to long 36

36. Second antenna with one terminal claw 37

Second antenna with two terminal claws (or in Doridicola fishclsoni with one claw and

one long spine) 43
NUMBER 127 85

37. Mandible with a scalelike area on convex side of base 38

Mandible without a scalelike area but with digitiform processes or tooth on convex side ...40

38. Mandible with lash reduced, and represented only by a small pointed process

Colobomolgus

Mandible with a long pectinate lash 39

39. Leg 4 with third exopod segment 11,13 Paramolgus

Leg 4 with third exopod segment 111,1,5 Paradoridicola

40. Mandible with convex side of base bearing a proximally directed tooth Odontomolgus

Mandible with convex side of base bearing one or more small digitiform lobes 41

41. Second maxilla with a large digitiform process on first segment Panjakus

Second maxilla without such a process 42

42. Mandible with convex side of base bearing two small digitiform lobes; leg 4 with third

exopod segment 11,13 AncMmolgiu

Mandible with convex side of base having a roughened area followed by a digitiform process; leg 4 with third exopod segment 111,1,5 Zamolgus

43. Mandible with very short lash 44

Mandible with long lash 45

44. Mandible with convex side of base having a scalelike area with spinules; lash of second

maxilla not "folded"; leg 4 with third exopod segment 111,1,5 Contomolgus

Mandible with convex side of base having a large hyaline area without spinules; lash of

second maxilla "folded"; leg 4 with third exopod segment H,I,5 Atcetomolgus

45. Mandible with convex side of base bearing a tooth 46

Mandible with convex side of base bearing a scalelike area with spinules 47

46. Tooth on mandible proximally directed; leg 4 with third exopod segment 11,13

PleHomolgus

Tooth on mandible distally directed; leg 4 with third exopod segment 111,13

Paredromolgus

47. Mandible with basal area beyond indentation densely spinose; free segment of leg 5 with

distal inner process Pennatuiicola

Mandible with basal area beyond indentation with a row of spinules on concave side and

scalelike area with row of spinules followed by a serrated fringe on convex side; free

segment of leg 5 without a distal inner process 48

48. Leg 4 with third exopod segment 11,13 Metaxymolgus

Leg 4 with third exopod segment 111,13 Doridicola

Genus Acaenomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform, elongated, except

the female of A. protulae where the prosome is

inflated. Urosome in the female 5-segmented, in

the male 6-segmented. Caudal ramus with six

setae. Rostrum rectangular or linguiform. First antenna 7-segmented. Second antenna 4-segmented,

slender, with the formula 1, 1, 2 or 3 -f- one

articulated claw, and several setae -j- three articulated claws (in A. protulae two of the setae are

clawlike, making five apparent claws in all).

Mandible with the base lacking a pronounced

indentation and the spinules usually seen on the

concave side but having a distally directed toothlike process on the convex edge; distal part of the

mandible an attenuated blade set at an angle to

the base and with marginal serrulations or denticulations. First maxilla with two or three elements.

Second maxilla of the usual lichomolgid type.

Maxilliped in the female 3-segmented with a

pointed tip, in the male 4-segmented (considering

the proximal part of the claw to represent a fourth

segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod with the formula 0-1 ;II, the seta being

feathered. Leg 1 endopod with the same formula

in both sexes in A. protulae (male unknown in

A. serpulae). Leg 5 with a free segment bearing

two terminal elements. Egg sacs long, reaching

well beyond the caudal rami.

Other features as in the species below.

Associated with polychaete annelids.

TYPE SPECIES.—Acaenomolgus protulae (Stock).

ETYMOLOGY.—The name is a combination of

the Greek words cuaiva (a thorn), alluding to the

toothlike process on the mandible, and \io\ya<; (a

sac made of leather), being part of the name of

the type-genus, Lichomolgus. Gender masculine.
86 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Species of the Genus Acaenomolgus

FEMALES

Prosome inflated; second antenna terminally with three articulated claws and two dawlike

setae; leg 4 endopod as long as the exopod A. protulae

Prosome not inflated; second antenna terminally with three claws and three setae; leg 4 endopod

distinctly shorter than the exopod A terpulae

Acaenomolgus protulae (Stock, 1959)

FICURE 48

tLichomolgus (Epimolgus) protulae Stock, 1959b, pp. 70-74,

figs. 7, 8 [from the plume if the polychaete Protula intestinum (Lamarck), Gulf of Naples, Italy].

Lichomolgus sp. (cf. protulae Stock) .-Stock. 1960c. p. 246

[copcpodids; from the plume of Protula tubularia (Mont.) ,

Banyuls, Mediterranean coast of France].

Lichomolgus protulae.-Gotto, 1961a. pp. 265-267 [from the

branchial filaments of the polychaete Protula tubularia,

Strangford Lough, northern Ireland]; 1962. p. 104; 1966.

p. 194.

a

FIGURE 48.—Acaenomolgus protulae (Stock). Female: a, dorsal; b, mandible; c, leg 4; d, leg 5.

Male: e, dorsal. (From Stock, 1959b, figs. 7b,c,h, 8a.b.) Length of female 1.5-2.0 mm. of male

1.0-1.4 mm.
NUMBER 127

Acaenomolgus serpulae (Stock, 1960)

Lichomolgus serpulae Stock, 1960c, pp. 242-246, figs. 13, 14

[from the plume of the polychaete Serpula vermicularis

Linnaeus, between Cerbere and Banyuls, Mediterranean

coast of France] -Gotto. 1961a, p. 267.

REMARKS.—The male is unknown.

Genus Acanthotnolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, in the

female with the formula 4, 13, 6, 3, 4 -|- 1 aesthete,

2 -f 1 aesthete, and 7 -J- 1 aesthete; in the male

with 4, 13 4- 2 aesthetes, 6, 3 + 1 aesthete,

4 -j- 1 aesthete, 2 -f- 1 aesthete, and 7+ 1 aesthete. Second antenna 4-segmented with the armature 1, 1, 3, II -\- several small elements (usually

five), there being two terminal claws, often long.

Labrum with the two lobes separated by a deep

cleft. Mandible with the basal area distal to the

indentation, having on its convex side a scalelike

area with spinules followed by a serrated fringe

and on its concave side a row of spinules; lash

long. Paragnath a small hairy lobe. First maxilla

87

with four elements (except three in A. varirostratus

and A. bilobipes). Second maxilla of the usual

lichomolgid type. Maxilliped of the female 3 -

segmented, with a pointed tip; in the male 4 -

segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Leg 4 exopod

with third segment 11,1,5. Leg 4 endopod with the

formula 0—I;II, the first segment bearing a spine

rather than a seta. Armature similar in both sexes

except for leg 1 endopod in the male where the

last segment is 1,1,4 instead of 1,5 as in the female.

Leg 5 with a free segment bearing two terminal

elements. Leg 6 in the female represented by the

two setae and the process near the area of attachment of each egg sac, in the male by a posteroventral flap on the genital segment bearing two

setae.

Other features as in the species below.

Associated with octocorals.

TYPE-SPECIES.—Acanthomolgus exilipes (Humes

and Ho).

ETYMOLOGY.—The name is formed from the

Greek words axav*a (a spine), referring to the spine

on the first segment of leg 4 endopod, and

Key to Species of the Genus Acanthomolgus

FEMALES

1. Free segment of leg 5 with inner margin bilobed A. bilobipes

Free segment of leg 5 not bilobed, either straight or with a proximal inner expansion

or toothlike process 2

2. Two setae on second segment of maxilliped about equal in length 3

One of the two setae on second segment of maxilliped much longer than the other, at

least 2:1 6

3. Two long median terminal setae on caudal ramus naked; free segment of leg 5 with a

prominent inner basal pointed expansion directed distally A. cuneipes

Two long median terminal setae on caudal ramus with long spinules; free segment of leg

5 with a small inner basal rounded expansion 4

4. Sides of expanded part of genital segment subparallel rather than rounded; mandible with

small blunt process on basal area near serrated fringe A. exilipes

Sides of expanded part of genital segment rounded; mandible without such a process 5

5. Second segment of leg 4 endopod with inner hairs A. gentilh

Second segment of leg 4 endopod with small inner spinules A. fissisetiger

6. Inner margin of basis of leg 4 with a row of hairs; rostrum sexually dimorphic; two long

median terminal setae on caudal ramus naked A. varirostratus

Inner margin of basis of leg 4 smooth; rostrum alike in both sexes; two long median

terminal setae on caudal ramus with lateral spinules (sometimes sparse and minute)... .7

7. Free segment of leg 5 with an inner proximal distally directed process separated sharply

from the more distal margin of the segment 8

Free segment of leg 5 with at most a rounded inner expansion not sharply set off from the

rest of the margin 10
g g SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

8. Genital segment broadest anterior to middle; free segment of leg 5 shorter than genital

segment A. versevtldH

Genital segment broadest posterior to middle; free segment of leg 5 as long as genital

segment 9

9. Second segment of first antenna six times longer than third; first segment of leg 1 exopod

with spine unusually long (twice the length of the spine on second segment)

A. longispinifer

Second segment of first antenna four times longer than third; first segment of leg 1 exopod

with spine about as long as that on second segment A. astrictus

10. Two long median terminal setae on caudal ramus with a few minute spinules visible only

under high magnification 11

Two long median terminal setae on caudal ramus with many long spinules 12

11. Distalmost element on third segment of second antenna a naked spine; second segment of

leg 4 endopod with inner margin smooth; free segment of leg 5 about five times longer

than wide . A. hales

Distalmost element on third segment of second antenna minute (2 ft) and hyaline; second

segment of leg 4 endopod with inner margin having small spinules; free segment of leg

5 about two times longer than wide A. plantei

12. Maxilliped with two terminal elements of about equal length and highly divergent, and

one seta on second segment about twice the length of the other; second segment of leg

4 endopod with a few hairs on inner margin A. Mans

Maxilliped with two terminal elements unequal and not conspicuously divergent, and one

seta on second segment about four times the length of the other; second segment of leg

4 endopod with a smooth inner margin d. telritophilut

MALES

1. Two long median terminal setae on caudal ramus naked 2

Two long median terminal setae on caudal ramus with spinules, either long or very short

and visible only under high magnification 3

2. Rostrum with a minute median posteroventral knob; free segment of leg 5, S3 x 9 fi

A. variroUratus

Rostrum rounded as in female; free segment of leg 5, 51 x 8 fx A. cuneipes

3. Two long median terminal setae on caudal ramus with a few very short spinules visible

only under high magnification 4

Two long median terminal setae on caudal ramus with long lateral spinules 5

4. Second segment of second antenna with smooth inner surface (no sexual dimorphism) ;

two claws on this appendage relatively short and stout, the longer claw distinctly shorter

than the greatest length of the fourth segment A. hates

Second segment of second antenna with small spines on inner surface; two claws on this

appendage relatively long and slender, the longer almost as long as the greatest length

of the fourth segment A. plantei

5. One of the two setae on the second segment of the maxilliped terminating in several pointed

spiniform elements A. fissisetiger

Both setae on the second segment of the maxilliped attenuated (but one seta obtuse in

A. verseveldti) 6

6. First segment of leg 1 exopod with spine unusually long (twice the length of the spine

on the second segment) A. longispinifer

First segment of leg 1 exopod with spine about as long as that on second segment 7

7. Second segment of leg 4 endopod with smooth inner margin 8

Second segment of leg 4 endopod with hairs or small spinules on inner margin 10

8. Setae on first and second segments of second antenna fringed on one side A. bilobipes

Setae on first and second segments of second antenna smooth 9

9. One of the three elements on third segment of second antenna angularly bent; two terminal claws on this appendage distinctly shorter than the fourth segment A. teUstophitus

All three elements on third segment of second antenna straight; two terminal claws on

this appendage nearly as long as fourth segment A. astrictus

10. Two claws on second antenna about as long as fourth segment 11

Two claws on second antenna distinctly shorter than fourth segment 12
NUMBER 127 89

11. Mandible with a small blunt process on basal area near senated fringe A. exilipes

Mandible without such a process on basal area near serrated fringe A. gentilis

12. Third segment of leg 1 endopod with outer spine recurved and process between the two

spines blunt A. verseveldti

Third segment of leg 1 endopod with outer spine straight and process between the two

spines pointed A. hians

Acanthomolgus exilipes (Humes and Ho, 1968)

Lichomolgus exilipes Humes and Ho, 1968b, pp. 7-10, figs.

32-55 [from the alcyonaceans Dendronephthya mucronata

(Putter), Dendronephthya regia Verseveldt, Dendronephthya stocki Verseveldt, and Dendronephthya koellikeri

Kiikenthal, region of Nosy Be, northwestern Madagascar].

NEW RECORDS (all from Madagascar, collected by

AGH).—From Dendronephthya mucronata: 1 9.

from single colony, in 3 m, Antsmantsara, Nosy

ht, 9 June 1967; several specimens, from four

colonies, in 10 m, Rocher du N.E., Nosy Be, 19

August 1967; many specimens, from two colonies,

in 10 m, Rocher du N.E., Nosy Be, 19 August 1967;

many specimens, from single colony, in 25 m,

north of Ankazoberavina, near Nosy Be\ 13°27.6' S,

47°58.2' E, 24 August 1967; and several specimens,

from single colony, in 24 m, north of Ankazoberavina, near Nosy Be, 13°27.6' S, 47°58.2' E, 25 August

1967.

From Dendronephthya regia: 78 9 $, 96 tf tf,

and 24 copepodids from single colony, in trawl,

26-23 m, near Nosy Be, 13°15'50" S, 48°08'35" E,

29 August 1967.

From Dendronephthya stocki: many specimens,

from single colony, in 25 m, north of Ankazoberavina, near Nosy Be, 13°27.6/

 S, 47°58.2' E, 24

August 1967.

From Dendronephthya sp.: many specimens,

from part of single colony, in 27 m, Bane des

Freres, Isles Mitsio, near Nosy B6, 12°58' S,

48°28' E, 17 August 1967.

NEW HOSTS.—From Dendronephthya speciosa Kiikenthal: 2 9 9» from single colony, in 17 m, near

black buoy, north of Pte. Ambarionaomby, Nosy

Komba, near Nosy Be\ Madagascar, 5 August 1967;

1

 9» 5 cf cf, and 2 copepodids, from single colony,

in 24 m, north of Ankazoberavina, near Nosy B£,

13°27.6' S, 47°58.2' E, 25 August 1967; both collections by AGH.

From Stereonephthya cordylophora Verseveldt:

many specimens, from single colony, in 24 m,

north of Ankazoberavina, near Nosy B£, Madagascar, 13°27.6' S, 48°58.2' E, 25 August 1967.

Acanthomolgus astrictus, new species

FICURES 49-52

TYPE MATERIAL.—384 99 , 302 cT cf, and 48

copepodids from one colony of a gorgonian, Acanthogorgia aspera Pourtales, in 23 m, Tany Kely, a

small island south of Nosy Bd, Madagascar, 30 July

1967, collected by AGH. Holotype 9, allotype,

and 580 paratypes (330 9 9, 250 cf cf) deposited

in USNM; 60 paratypes (30 99 , 30 rfcf) in

ZMA; and the remaining paratypes in the collection

of AGH.

OTHER MATERIAL EXAMINED (all from Acanthogorgia aspera).—290 9 9, 109 cf cf, and 22 copepodids from single colony, in 20 m, Bane de Cinq

Metres, west of Nosy Be\ 6 August 1967; and 150

9 9, 140 cf d\ and 100 copepodids from single

colony, in 24 m, north of Ankazoberavina, 13°27.6'

S, 47°58.2' E, west of Nosy Be, 25 August 1967. All

collected by AGH.

FEMALE.—The body (Figure 49a) resembles in

general form that of A. cuneipes (Humes and Ho).

The length is 0.89 mm (0.80-0.99 mm) and the

greatest width 0.44 mm (0.41-0.46 mm), based on

ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.65:1. The

ratio of the length of the prosome to that of the

urosome is 2.75:1. The body is rather weakly

sclerotized.

The segment of leg 5 (Figure 496) is 75 x 125 v-.

Between this segment and the genital segment there

is a very short ventral intersegmental sclerite (Figure 49c). The genital segment is as wide as long,

114 x 114 |i, in dorsal view with rounded lateral

margins which abruptly join a short posterior

region with parallel sides. The areas of attachment

of the egg sacs are located dorsolaterally near the

middle of the segment. Each area (Figure 49d)

bears two naked spiniform setae, 9 n and 13 n in

length, and a small spiniform process. The three

postgenital segments are 32 x 67 n, 23 x 66 \x, and

26 x 65 n from anterior to posterior.

The caudal ramus (Figure 49e) is shorter than

wide, 24 x 27.5 n in greatest dimensions. The outer
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 49.—Acanthornolgtis astrictus, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; segment of leg 5 and genital segment, lateral (B) ; d, area of attachment of egg sac, dorsal

(c); e, caudal ramus, dorsal (D) ; /, rostrum, ventral (B) ; g, rostrum, lateral (B) . Scale: A, 02

mm; B, 0.1 mm; c, 0.02 mm; D, 0.05 mm.
NUMBER 127 91

lateral seta is 96 n, the dorsal seta 25 n, and the

outermost terminal seta 122 n, all three of them

naked. The innermost distal seta is 195 n with

lateral hairs, and the two long median terminal

setae 440 n (outer) and 570 n (inner), both inserted

between dorsal (unornamented) and ventral (with

a marginal row of minute spinules) flaps and both

with prominent coarse lateral spinules in their

proximal halves.

The fine ornamentation of the body consists

of a few hairs (sensilla). The anal segment has a

row of minute spinules along its posteroventral

margin on both sides.

The egg sacs are elongated, reaching far beyond

ilie caudal rami and resembling those of A.

cuneipes. In nearly all ovigerous females, however,

the sacs were broken. The only complete sac seen

measured 475 x 200 n, with each of the many eggs

about 49 n in diameter.

The rostrum (Figure 49/) in ventral view is

broadly rounded posteroventrally, and in lateral

view (Figure 49g) projects ventrally.

The first antenna (Figure 50a) is about 400 \i

long. The lengths of the seven segments (measured

along their posterior nonsetiferous margins) are

31 (65>x along the anterior margin), 112, 28, 57, 60,

43, and 31 n respectively. The formula for the

armature is 4, 13, 6, 3, 4 -|- 1 aesthete, 2 —f- 1

aesthete, and 7 -f 1 aesthete. All the setae are

naked.

The second antenna (Figure 506) is 4-segmented

and rather slender. Each of the first two segments

bears an inner seta, and the inner margin of the

second segment has a few minute hyaline spinules.

The third segment bears three setae. The moderately elongated fourth segment, 79 n along its outer

edge, 50 n along its inner edge, and 21 n wide,

bears two unequal terminal claws, the slender claw

57 n along its axis, the stout claw 53 n, and five

setae. All the setae are naked.

The labrum (Figure 50c) has two broad posteroventral lobes.

The mandible (Figure 50d) resembles that of

A. longispinifer (Humes and Ho) but the scale-like

protrusion on the convex side distal to the basal

constriction is rounded rather than pointed. The

paragnath (Figure 50c) is a small hairy lobe. The

first maxilla (Figure 50e) bears four setae. The

second maxilla (Figure 50/) resembles that of A.

fissisetiger (Humes and Ho). The maxilliped (Figure 50g) has groups of spinules on its first segment.

The second segment bears two setae somewhat

unequal in length. The third segment has a small

naked seta and two nearly equal terminal elements, widely divergent as in A. hians (Humes

and Ho).

The area between the maxillipeds and the first

pair of legs (Figure 50/») is not protuberant; a fine

sclerotized line connects the bases of the maxillipeds.

Legs 1-4 (Figure 5\a-d) have the same segmentation and formula for the armature as in A. varirostratus (Humes and Ho) and all other species

in the genus. The inner seta on the coxa of leg 4

is naked and minute, only 6 n long. The setae on

the third segment of each exopod have distally

somewhat longer hairs along the inner side than

on the outer side. Leg 3 is similar to leg 2 except

for the formula of the third segment of the endopod

(Figure 51c). The endopod of leg 4 (Figure 5Id)

is 113 |i long (without the spinous processes), a

little shorter than the exopod, which is 156 n.

The first segment of the endopod is 37 x 26 n (including the terminal spinous processes) and bears

an inner distal spine that is 27 n and very finely

barbed. The second segment is 80 v. long (including the processes), 22 v- in greatest proximal width,

and 15.5 v in least distal width. Its two unequal

terminal spines are 28 n (outer, finely barbed) and

67 n (inner, with spinulose fringes). The outer

margins of both segments are haired,* their inner

margins naked, and the second segment has a row

of minute spinules near the insertions of the

terminal spines.

Leg 5 (Figure b\e) has an elongated free segment

112 x 26 n in greatest dimensions and of peculiar

form. Proximally the segment is constricted (width

here 13 v), the region proximal to the constriction

with a distomedially directed beaklike process.

Distal to the constriction the segment is rather

broadened with a curved outer margin but fairly

straight inner margin. The outer terminal seta is

88 n and naked. The inner terminal seta is 120 i*

and naked, and arises between two small lobes. The

seta on the body near the insertion of the segment

is 50 n and lightly feathered. The free segment is

ornamented outwardly on both dorsal and ventral

surfaces with small flattened spines.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 49d).
FIGURE 50.—AcarUhomolgus astrictus, new species. Female: a, first antenna, with arrows indicating positions of additional aesthetes in male, ventral (B) ; b, second antenna, postero-inner

(E) ; c, labrum and paragnaths, ventral (D); d, mandible, posterior (D) ; e, first maxilla, posterior (c); /, second maxilla, posterior (D) ; g, maxilliped, postero-inner (D) ; h, area between

maxillipeds and first pair of legs, ventral (E). Scale: B, 0.1 mm; c, 0.02 mm; D, E, 0.05 mm.
NUMBER 127 93

FIGURE 51.—Acanthomologus as trie l us, new species. Female: a, leg 1 and intercoxal plate,

anterior; b, leg 2, anterior; c, third segment of endopod of leg 3, anterior; d, leg 4 and intercoxal plate, anterior; e, kg 5, dorsal. Scale: E, 0.05 mm, applies to all drawings.
94

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The color in life in transmitted light is slightly

reddish (because of red globules within), the eye

red, the egg sacs light reddish brown or gray. (The

reddish color imitates the dark red color of the

host gorgonian).

MALE.—The body (Figure 52a) resembles in

general aspect that of the female. The length is

0.72 mm (0.67-0.78 mm) and the greatest width

0.27 mm (0.25-0.31 mm), based on ten specimens

in lactic acid. The ratio of the length to the width

of the prosome is 1.54:1. The ratio of the length

of the prosome to that of the urosome is 1.67:1.

The segment of leg 5 (Figure 526) is 39 x 70 n.

There is no ventral intersegmental sclerite between

this segment and the genital segment. The genital

segment is about as long as wide, 153 x 151 n, in

dorsal view with its lateral borders rounded. The

four postgenital segments are 23 x 44 \i, 18 x 44 n,

11 x 43 n, and 16 x 44 n from anterior to posterior.

The caudal ramus resembles that of the female,

but is smaller, 19 x 20 n.

The fine ornamentation of the body is like that

of the female.

The rostrum is similar to that of the female.

The first antenna (Figure 52a) also is like that of

the female but has three additional aesthetes (their

positions indicated by arrows in Figure 50a), so

that the formula is 4, 13 -f 2 aesthetes, 6, 3 -|- 1

aesthete, 4 -f 1 aesthete, 2 -j- 1 aesthete, and

7 —|— 1 aesthete. The second antenna (Figure 52c)

resembles that of the female, but the minute

spinules on the inner edge of the second segment

are more numerous and the stouter of the two

claws is relatively less stout than in that sex.

The labrum, mandible, paragnath, first maxilla,

and second maxilla resemble those of the female.

The maxilliped (Figure 52d) is slender and 4-

segmented (assuming that the proximal part of the

claw represents a fourth segment). In general form

it resembles that of A. hians (Humes and Ho). The

claw is 159 f* along its axis (including the terminal

lamella). One of the two unequal slender setae on

the second segment is very finely barbed.

The area between the maxillipeds and the first

pair of legs is like that of the female.

Legs 1-4 are segmented as in the female and

have the same spinal and setal formula except for

the last segment of the endopod of leg 1 (Figure

52e) where the formula is 1,1,4. Between the two

conspicuously barbed spines on this segment there

is a recurved process minutely spinulose along its

outer edge. Legs 2 and 3 are similar to those in

the female. The endopod of leg 4 (Figure 52/)

has slightly different proportions and the inner

spine on the first segment is relatively longer than

in the opposite sex. The first segment is 27 x 19 n

(including the terminal spinous processes) and its

spine is 31 n. The second segment is 52 n long (including the processes), 15.5 n in greatest width, and

11.5 ji in least width. The outer terminal spine is

22 n, the inner 53 n.

Leg 5 (Figure 52g) has a slender subrectangular

free segment, 41 x 9 n, ornamented with fewer and

smaller outer spinules than in the female, and

bearing two terminal naked setae 57 n (outer) and

23 »i (inner).

Leg 6 (Figure 52/i) consists of the usual posteroventral flap on the genital segment bearing two

naked setae, 39 n and 52 n.

The spermatophore (Figure 52i), attached to the

female in pairs, is 133 x 73 n (not including the

neck).

The color in life resembles that of the female,

but tends to be less reddish.

ETYMOLOGY.—The specific name astrictus (Latin,

bound closely or drawn together) alludes to the

proximal constriction of leg 5 in the female.

COMPARISON WITH RELATED SPECIES.—One of the

chief characters of the genus Acanthomolgus is the

possession of the formula 0—I;II on the endopod of

leg 4. In seven species the inner margin of leg 5

in the female is not deeply indented proximally

and the inner process, if present, is broadly rounded

rather than toothlike (A. exilipes, A. gentilis, A.

fissise tiger, and A. varirostratus, all described as

Lichomolgus by Humes and Ho, 1968b; and A.

hians, A. telestophilus, and A. verseveldli, all

described as Lichomolgus by Humes and Ho,

1968d).

The two remaining species, A. cuneipes (Humes

and Ho) and A. longispinifer (Humes and Ho),

have an inner proximal toothlike process on the

free segment of leg 5 in the female, a feature which

suggests affinity with A. astrictus. In the female of

A. astrictus, however, the shape of the free segment of leg 5, with its proximal constriction and

distal expansion, is unlike that in those two species.

A. cuneipes further differs from the new species

ii\ the absence of lateral spinules on the two long

median terminal setae on the caudal ramus and
NUMBER 127 95

FIGURE 52.—Acanthomolgus astrictus, new species. Male: ;, dorsal (A) ; b, urosome, dorsal (B);

c, second antenna, postero-inner (E) ; d, maxilliped, outer (E) ; e,e, thir thirdd segmen segmentt o off endopo endopodd o off

leg 1, anterior (D) ; /, endopod of leg 4, anterior (D) ; g, leg 5, dorsal (D) ; h, leg 6, ventral (E) ;

i, spermatophore, attached to female, dorsal (B) . Scale: A, 02 mm; B, 0.1 mm; D, E, 0.05 mm.
96 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

in the 4 or 5 central teeth on the lash of the second

maxilla being stouter than the other teeth; and

A. longispinifer differs in the outer spine on the

first segment of the exopod of leg 1 being unusually

long.

Acanthomolgns bilobipes, new species

FIGURES 53-55

TYPE MATERIAL.—11 9 9 , 26 cf cf txom a gorgonian, Antillogorgia acerosa (Pallas), in 3 m, Lime

Cay, near Kingston, Jamaica, 30 August 1959, collected by AGH and RUG. Holotype $, allotype,

and 31 paratypes (8 9 $, 23 cf cf) deposited in

USNM, the remaining paratypes (dissected) in

the collection of AGH.

OTHER MATERIAL EXAMINED.—From Antillogorgia

acerosa: 9 9$ » 4 cfcf. in 7 m, Oistin's Bay,

Christchurch, Barbados, 15 July 1959. From Antillogorgia elastica Bieschowsky: 7 9 ?» 2 cf cf» in

4 m, lee side of Terremoto Reef (Cayo Terremoto),

southeast of La Parguera, southwestern Puerto Rico,

6 August 1959; and 10 $ 9 , 3 cf cf. in 3 m,

Terremoto Reef, 9 August 1959. All collected by

AGH and RUG.

All of the gorgonians had basket stars, Astrophyton muricatum Lamarck, entwined about them.

Acanthomolgus bilobipes is believed to be associated with the gorgonian rather than with the basket

star, since it always occurred in washings of gorgonians and basket stars together, never in washings

of basket stars alone. Unfortunately, gorgonians

without basket stars were not examined in the

field. Doridicola astrophyticus, described elsewhere

in this work, is apparently an associate of basket

stars, since it was found in washings of these echinoderms living apart from gorgonians.

FEMALE.—The body (Figure 53a) has a moderately slender prosome. The length (without the

setae on the caudal rami) is 0.94 mm (0.86-0.98

mm) and the greatest width 0.45 mm (0.41-0.47

mm) based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome

is 1.54:1. The ratio of the length of the prosome

to that of the urosome is 2.41:1. The segment of

leg 1 is set off dorsally from the cephalosome by a

transverse furrow. The epimeral areas of the pedigerous segments are as indicated in the figure.

The segment of leg 5 (Figure 53&) is 64 x 122 I*.

There is no ventral intersegmental sclerite between

this segment and the genital segment. The genital

segment is 114 x 140 n in greatest dimensions, in

dorsal view of characteristic form (with slight

wings at the maximum width terminating in a

small knob). The areas of attachment of the egg

sac are located dorsolaterally near the middle of

the segment. Each area (Figure 53c) bears two long

naked setae, 28 n. The three postgenital segments

are 39 x 75 n, 32 x 69 n, and 30 x 68 M from anterior

to posterior.

The caudal ramus (Figure 53d) is 23 x 28 n,

wider than long. The outer lateral seta is 68 n and

naked. The dorsal seta is 30 n and naked. The

outermost terminal seta is 91 M with proximal inner

spinules. The innermost terminal seta is 133 n

with bilateral spinules. The two long median

terminal setae are 345 n (outer) and 480 n (inner),

both with spinules and inserted between dorsal

(unornamented) and ventral (with a row of

spinules) flaps.

The body surface has a few hairs (sensilla) as

shown in Figure 53a,6.

The egg sac is unknown.

The rostrum (Figure 5ie) has a broadly rounded

posteroventral margin.

The first antenna (Figure 53/) is slender and

356 »i long. The lengths of its seven segments

(measured along their posterior nonsetiferous

margins) are 24 (60 n along the anterior margin),

99, 22, 50, 50, 41, and 34 » respectively. The armature is 4, 13, 6, 3, 4 -f 1 aesthete, 2 -f 1 aesthete,

and 7 -f 1 aesthete. All the setae are naked.

The second antenna (Figure 53g) has a small

seta and a few minute spinules on both first and

second segments. The third segment has three

setae. The fourth segment, 67 n along its outer

edge, 40 n along its inner edge, and 22 i* wide, has

terminally two somewhat unequal claws, the stouter

one 35 n along its axis, the more slender one 33 n,

and five slender setae. All the setae are naked.

The labrum (Figure 53/i) has two posteroventral

lobes.

The mandible (Figure 53i) resembles that in

other species of the genus. The paragnath, indicated

by broken lines in Figure 53c, is a small hairy lobe.

The first maxilla (Figure 53;) has three elements.

The second maxilla (Figure 53A) has the usual

form for the genus. The maxilliped (Figure 54a)

has a few spinules on the first segment. The second

segment has two unequal setae. The third segment
FIGURE 53.—Acanthomolgus bilobipes, new species. Female: a, dorsal (A) ; b, urosomc, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum,

ventral (B) ; /, first antenna, dorsal (E) ; g, second antenna, anterior (F) ; h, labrum, with

paragnaths indicated by broken lines, ventral (c); i, mandible, posterior (c); ;, first maxilla,

posterior (c); k, second maxilla, posterior (c). Scale: A, 05 mm; B, 02 mm; c, D, 0.05 mm; E, F,

0.1 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE bA.—Acanthomolgus bilobipes, new species. Female: a, maxilliped, posterior (c); b, area

between maxillipeds and first pair of legs, ventral (F.) ; c, leg 1 and intercoxal plate, anterior

(E) ; d, leg 2 and intercoxal plate, anterior (E) ; e, third segment of endopod of leg 3, anterior

(E) ; /, leg 4 and intercoxal plate, anterior (F.) ; g, free segment of leg 5, inner (F) . Male: h,

dorsal (c). Scale: c, 0.05 mm; E, F, 0.1 mm; c, 0.5 mm.
NUMBER 127 99

has a barbed spine and a small setule and terminates in a barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 546) is not protuberant.

A line connects the bases of the maxillipeds.

Legs 1-4 (Figures 54c-/) are segmented and

armed as in other species of the genus. Leg 1

(Figure 54c) has a small posterior protrusion on

the outer side of the coxa. The inner coxal seta in

legs I-S is long and plumose, but in leg 4 it is

short (14 n) and naked. The inner margin of the

basis of legs 1-3 is haired, but in leg 4 it is smooth.

The exopod of leg 4 (Figure 54/) is 156 n long.

The first segment of the endopod is 36 x 24 n (not

including the spiniform processes), with the inner

distal spine 27 n and naked. The second segment

is 91 ji long (with the processes) and 15 n in least

width near the middle; the two terminal fringed

spines are 33 u (outer) and 63 n (inner). Both

segments have the outer margins haired.

Leg 5 (Figure 54g) has a free segment, 83 x 43 n,

its outer surface finely spinulose, its inner margin

bilobed. The two naked terminal setae are 57 n

and 95 n.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 53c).

The color in life in transmitted light is translucid

to very light tan, the eye red.

MALE.—The body (Figure 54/J) has a prosome

more slender than in the female. The length (excluding the ramal setae) is 0.76 mm (0.72-0.79

mm) and the greatest width 0.28 mm (0.26-0.30

mm), based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.81:1. The ratio of the length of the prosome to

that of the urosome is 1.93:1.

The segment of leg 5 (Figure 55a) is 33 x 76 n.

There is no ventral intersegmental sclerite. The

genital segment is 135 x 135 n. The four postgenital segments are 23 x 52 n, 20 x 50 n, 14 x 48 /*,

and 20 x 50 M from anterior to posterior.

The caudal ramus is similar to that of the female.

The body surface is ornamented with hairs as in

the female.

The rostrum is like that of the female.

The first antenna resembles that of the female

but has three additional aesthetes (Figure 54/i), two

on the second segment and one on the fourth seg-
100

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ment, so that the formula is 4, 13 -f- 2 aesthetes,

6, 3 -f- 1 aesthete, 4+ 1 aesthete, 2 + 1 aesthete,

and 7+ 1 aesthete. The second antenna (Figure

55b) is much like that of the female, but the setae

on the first and second segments and one on the

third segment are finely pectinate, and segments 1,

2, and 4 have minute spinules on their inner surfaces.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 55c) has an unarmed first

segment. The second segment has two setae and

two rows of spines. The third segment is unarmed.

The claw is 185 n along its axis, with an unusually

large terminal lamella, weakly divided midway,

and with two very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that in the female.

Legs 1-4 are segmented as in the female. Leg 1

is armed as in the female except for the last segment of the endopod (Figure 55d) which has the

formula 1,1,4. Leg 2 is similar to that of the female

but the last segment of the endopod (Figure 55*)

has the middle spine weaker than the other two

and the inner spinous process much longer. Legs

3 and 4 are like those of the female.

Leg 5 (Figure 55a) has a slender free segment,

33 x 8 n, not bilobed as in the female. The two

terminal elements are 14 n and 44 n. The outer

surface of the free segment has only a few spinules.

Leg 6 (Figure 55a) is a posteroventral flap on the

genital segment bearing two slender naked setae

25 »i and 36 n.

The spermatophore is unknown.

The color in life is like that of the female.

ETYMOLOGY.—The specific name bilobipes (from

the Latin bis, twice; lobus, a lobe; and pes, a foot)

alludes to the bilobed inner margin of the free

segment of leg 5 in the female.

REMARKS.—Acanthomolgus bilobipes may be

separated from all other members of the genus by

the bilobed inner margin of the free segment of

leg 5 in the female and by the unusually large

terminal lamella on the claw of the male maxilliped.

Acanthomolgus cuneipes (Humes and Ho, 1968)

Lichomolgus cuneipes Humes and Ho, 1968b, pp. 17-20, figs.

84-96 [from the alcyonacean Stereonephthya acaulis Verseveldt, Nosy Bl, Madagascar].

NEW HOST.—Several specimens from six colonies

of Dendronephthya mucronata (Putter), in 1 m,

off Ampombilava, Nosy Be, 26 September 1964,

collected by AGH.

Acanthomolgus fissisetiger (Humes and Ho, 1968)

Lichomolgus fissisetiger Humes and Ho, 1968b, pp. 14-17,

figs. 70-83 [from the alcyonaceans Stereonephthya acaulis

Versevcldt, Stereonephthya papyracea Kiikenthal, and

Lemnalia elegans (May), region of Nosy Be". Madagascar;

Dr. Versevcldt recently changed the host identification of

Stereonephthya papyracea to S. acaulis Vcrscveldt].

NEW RECORD.—22 9 9 , 42 cf cf»

 an d 4 4

podids from three colonies of Stereonephthya

acaulis, in 15 m, Bane du Touareg, near Nosy Be\

Madagascar, 1 September 1967, collected by AGH.

NEW HOST.—8 99 , 5 cf cf.

 ant^ 8 copepodids

from the alcyonacean Lemnalia humesi Verseveldt,

in 10 m, Pte. Lokobe, Nosy Be\ Madagascar, 18

July 1967, collected by AGH.

Acanthomolgus gentilis (Humes and Ho, 1968)

Lichomolgus gentilis Humes and Ho, 1968b, pp. 11-14, figs.

56-69 [from the alcyonaceans Dendronephthya mucronata

(Putter), Dendronephthya koellikeri Kiikenthal. Dendronephthya stocki Verseveldt. Stereonephthya acaulis Versevcldt.

and Stereonephthya papyracea Kiikenthal, region of Nosy

Be", Madagascar; Dr. Verseveldt recently changed the host

identification of Stereonephthya papyracea to S. acaulis

Verseveldt].

NEW RECORDS (all from Madagascar, collected by

AGH).—Fro m Dendronephthya mucronata: 3 9$ .

6 cf cf» and 13 copepodids, from single colony, in

3 m, Antsamantsara, Nosy B£, 9 June 1967; 22 9 9 ,

23 cf cf.

 an d 14 copepodids, from single colony, in

4 m, Antsamantsara, Nosy B£, 9 June 1967; many

specimens, from two colonies, in 10 m, Rocher du

N.E., Nosy Be\ 19 August 1967; and several specimens, from single colony, in 24 m, north of Ankazoberavina, near Nosy Be\ 13°27.6' S, 47°58.2/

 E,

25 August 1967.

From Dendronephthya sp.: many specimens, from

part of single colony, in 27 m, Bane des Freres,

Isles Mitsio, near Nosy Be\ 12°58' S, 48°28' E, 17

August 1967.

From Stereonephthya acaulis: 43 9 ?» '* cf cf.

and 76 copepodids, from single colony, in 4 m,

Antsamantsara, Nosy Be", 9 June 1967; 6 99 , 12

cf cf»

 ai

»d 7 copepodids, from single colony, in 2
NUMBER 127 101

m, Ambatoloaka, Nosy Be, 23 June 1967; 12 $ 9 ,

24 cf d > an d 27 copepodids, from single colony,

in 8 m, Pte. Lokobe, Nosy Be, 25 July 1967; and

26 9 9» 38 cf cf, and 45 copepodids, from single

colony, in 17 m, near black, buoy, north of Pte.

Ambarionaomby, Nosy Komba, near Nosy Be, 5

August 1967.

NEW ALCYONACEAN HOSTS (all from Madagascar,

collected by AGH).—From Umbellulifera striata

(Thompson and Henderson): 2 9 9» 4 cf cf, from

four colonies, in 17 m, in pass at Pte. Lokobe, Nosy

B*. 25 August 1967.

From Dendronephthya lokobeensis Verseveldt:

3 9 ?» 6 cf cf. from single colony, in 4 m, Antsamantsara, Nosy Be, 9 June 1967.

From Dendronephthya speciosa Kiikenthal: 5

9 V . 10 cf d. and 1 copepodid, from single colony,

in 22 m, Tany Kely, near Nosy Be\ 17 June 1967;

20 9 9 ' '-' cf cf» and 4 copepodids, from single

colony, in 17 m, near black buoy, north of Pte.

Ambarionaomby, Nosy Komba, near Nosy Be, 5

August 1967.

From Stereonephthya cordylophora Verseveldt:

many specimens, from single colony, in 24 m, north

of Ankazoberavina, near Nosy Be, 12°27.6' S,

47°58.2' E, 25 August 1967.

Acanthomolgus hales, new species

FIGURES 56-58

TYPE MATERIAL.—404 99 , 581 cf cf» and 646

copepodids from single colony of the gorgonian

Solenocaulon tortuosum Gray, on sand in 15 m,

in pass between Nosy Komba and Nosy Be, northwestern Madagascar, 13 April 1966, collected by

R. Plante. Holotype 9 , allotype, and 730 paratypes (280 99 , 450 cf cf) deposited in USNM;

200 paratypes (100 99 , 100 cf cf) in ZMA; and

the remaining paratypes and copepodids in the

collection of AGH.

OTHER MATERIAL EXAMINED (all from Solenocaulon tortuosum).—214 99 , 55 cfcf» and 61

copepodids from single colony, in 15 m, in pass at

Pte. Lokobe, Nosy Be, 13 June 1967; and 42 99 ,

29 cf cf» and 46 copepodids from four colonies,

in 18 m, in pass at Pte. Lokobe, Nosy Be, 3 July

1967. All collected by AGH.

FEMALE.—The body (Figure 56a) has a moderately expanded prosome. The length is 0.81 mm

(0.75-0.84 mm) and the greatest width 0.38 mm

(0.34-0.39 mm), based on ten specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.39:1. The ratio of the length of the

prosome to that of the urosome is 1.79:1.

The segment of leg 5 (Figure 566) is 60 x 112 n.

Between that segment and the genital segment there

is a well-developed ventral intersegmental sclerite.

The genital segment is 138 x 120 n, a little longer

than wide and not greatly expanded, with two

pairs of hyaline lateral membranes. The areas of

attachment of the egg sacs are situated dorsolaterally

near the middle of the segment. Each area (Figure

56c) bears two small naked spiniform elements,

7 n and 9 n in length, with a pointed process between them. Medial to the larger element the

cuticular fold is produced as a small thornlike

prominence. The three postgenital segments are

35 x 58 n, 31 x 56 1*, and 36 x 55 M from anterior

to posterior.

The caudal ramus (Figure 56d) is a little longer

than wide, 33 x 25 n. The outer lateral seta is 59 n

and the outermost terminal seta is 67 m both naked.

The dorsal seta is 32 n and very lightly feathered.

The innermost terminal seta is 135 n with its inner

margin haired proximally and with a few minute

spinules distally. The two long median terminal

setae are 273 n (outer) and 330 11 (inner), both

with a few widely spaced minute inner spinules

and both inserted between dorsal (unornamented)

and ventral (with a marginal row of spinules)

flaps.

The body surface is ornamented with a few

hairs (sensilla) as shown in Figure 56a,b. The

posteroventral margin of the anal segment bears

a row of spinules on both sides.

The egg sac (Figure 56a) is elongated, oval, 300

x 140 n, reaches beyond the caudal rami, and contains many eggs about 40 n in diameter.

The rostrum (Figure 56a) is broadly rounded

posteroventrally.

The first antenna (Figure 56/) is 284 n long. The

lengths of the seven segments (measured along their

posterior nonsetiferous margins) are 25 (51 n along

the anterior margin), 85, 18, 37, 39, 28, and 26 n

respectively. The formula for the armature is 4,

13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7

-|- 1 aesthete. All the setae are naked.

The second antenna (Figure 56g) is 4-segmented.

Each of the first two segments bears a naked seta.
FIGURE 56.—Acanthomolgus hales, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum, ventral

(E) ; /, first antenna, ventral (E) ; g, second antenna, anterior (F) ; h, labrum, with paragnaths

indicated by broken lines, ventral (D) ; i, mandible, posterior (D) ; ;, first maxilla, anterior (D) ;

k, second maxilla, posterior (o); I, maxilliped, postero-inner (D) . Scale: A, B, 02 mm; c, D,

0.05 mm; E, F, 0.1 mm.
NUMBER 127 103

The third segment bears a naked spine and two

naked setae. The last segment, 75 n along its outer

edge, 47 n along its inner edge, and 20 n wide,

bears two terminal unequal claws. The stouter

claw is 43 t» along its axis and the more slender

claw 32 n. Near the insertions of the claws are

five small hyaline elements.

The labrum (Figure 56h) has two broad posteroventral lobes.

The mandible (Figure 56i), paragnath (Figure

5(>/J). first maxilla (Figure 56;), and second maxilla

(Figure 56A) resemble those of A. exilipes (Humes

and Ho). The maxilliped (Figure 56/) has two very

unequal setae on the second segment, the longer

seta barbed along one edge, the shorter seta naked.

The last segment bears a naked spine, a small

naked setule, and an incompletely articulated spinifoi in process with minute barbules along one edge.

The first segment is ornamented with slender

spinules.

The ventral area between the maxillipeds and

the first pair of legs (Figure 57a) is not protuberant.

A sclerotized line connects the bases of the maxillipeds.

Legs 1-4 (Figure 57be) have the same spine and

setal formula as in other species of the genus. The

inner coxal element of leg 4 is reduced to a small

naked setule 5 n long. The inner margin of the

basis of leg 4 is smooth instead of having a row of

hairs as in legs 1-3. The outer exopod spines of

leg 1 have relatively small numbers of barbs on

their proximal edges. The inner margins of the

second and third segments of legs 1-3 lack hairs.

The exopod of leg 4 is 146 n. The endopod of leg

4 has outer hairs on both segments. The first

segment is 30 x 27 n (not including the spinous

processes) and its inner distal naked spine is 32 n.

The second segment is 70 n long (including the

terminal spinifonn processes), 25 p in greatest

width proximally, and 20 n in least width distally;

the outer terminal fringed spine is 21 M, the inner

spine is 53 n, inwardly fringed but outwardly

barbed.

Leg 5 (Figure 57/) has an elongated free segment 120 x 23 n (the width taken at the middle);

the width at the proximal inner expansion is 32 n.

The outer surface of the segment bears short, broad

spines. The two terminal setae are 57 n and naked

and 78 n with a narrow unilateral fringe. The seta

on the body near the free segment is 33 n and

lightly feathered.

Leg 6 is probably represented by the two spiniform elements near the area of attachment of the

egg sacs (Figure 56c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs grayish black.

MALE.—The body (Figure 57g) is more slender

than in the female. The length is 0.64 mm (0.58-

0.67 mm) and the greatest width is 0.25 mm (0.24-

0.25 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome is 1.65:1. The ratio of the length of the

prosome to that of the urosome is 1.59:1.

The segment of leg 5 (Figure 57h) is 32 x 57 n.

There is no ventral intersegmental sclerite. The

genital segment is 138 x 122 I*. The four postgenital

segments are 19 x 40 n, 19 x 39 n, 13 x 37 n, and

21 x 39 >i from anterior to posterior.

The caudal ramus is like that of the female,

but smaller, 23 x 19 I*.

The surface of the body is ornamented with

hairs as in the female.

The rostrum is similar to that of the female.

The first antenna is segmented and armed as in

the female but has three additional aesthetes as in

A. astrictus. The second antenna is the same as in

the female, without sexual dimorphism.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 57i) resembles in general

form that of A. varirostratus (Humes and Ho) but

the claw is shorter, 108 n along its axis (including

the terminal lamella), and more strongly recurved.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female except for the last segment of the endopod

of leg 1 (Figure 58a) which has 1,1,4 instead of 1,5

as in the opposite sex. On the last segment of the

endopod of leg 2 (Figure 586) the three spines

from outer to inner are 17.5, 15.5, and 22 H (in the

female 17, 14, and 22 v) and more strongly barbed

than in the opposite sex; and on the last segment

of the endopod of leg 3 (Figure 58c) the spines are

20, 20, and 24 n (in the female 21, 18, and 26 v)

and also are more strongly barbed. The endopod

of leg 4 (Figure 58d) has a slightly more slender

second segment than in the female. The dimensions

of this endopod are as follows: first segment 22 x
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 57.—Acanthomolgus hales, new species. Female: a, area between maxillipeds and first pair

of legs, ventral (F) ; b, leg I and intercoxal plate, anterior (F) ; c, leg 2, anterior (F) ; d, third

segment of endopod of leg 3, anterior (F) ; e, leg 4, anterior (F) ; /, leg 5, dorsal (F). Male: g,

dorsal (A) ; h, urosome, dorsal (E) ; i, maxilliped, postero-inner (F) . Scale: A, 02 mm; E, F,

0.1 mm.
NUMBER 127 105

e

FIGURE W.—Acanthomolgus hales, new specie*. Male: a, endopod of leg 1, anterior (D) ; b, third

segment of endopod of leg 2, anterior (D) ; c, third segment of endopod of leg 3, anterior (D) :

d, endopod of leg 4, anterior (D) ; e, spermatophores, attached to female, dorsal (E) . Scale: D,

0.05 mm; e, 0.1 mm.

17.5 n (without the spiniform processes) and its

spine 24 n; second segment 46 n long (including

the terminal spiniform processes), 14 M^ in greatest

width proximally, and 11 n in least width distally;

the outer terminal spine is 18 n, the inner spine

42 n.

Leg 5 (Figure 57/») has an elongated, slender

free segment, 30 x 7.5 \i, without a proximal expansion. The outer surface bears a few small

spinules and the two terminal elements are 14 n

and 39 p.

Leg 6 (Figure 51 h) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 21 n and 30 |».

The spermatophore (Figure 58e), attached to

the female in pairs, is oval, 104 x 55 n (without the

neck).

The color in life in transmitted light resembles

that of the female.

ETYMOLOGY.—The specific name hales (Greek

ahrc, in abundance) alludes to the great numbers

of this copepod (more than 1,600 specimens) found

on a single host colony (type collection).

REMARKS.—By the nature of the ornamentation

on the two long median setae on the caudal ramus,

A. hales may be distinguished from eight species

that have long spinules on these setae—A. exilipes

(Humes and Ho), A. fissisetiger (Humes and Ho),

A. gentilis (Humes and Ho), A. hians (Humes and

Ho), A. longispinifer (Humes and Ho), A. telestophilus (Humes and Ho), A. verseveldti (Humes

and Ho), and A. astrictus and A. bilobipes, described above—and from two species that have

these setae naked—A. cuneipes (Humes and Ho)

and A. varirostratus (Humes and Ho). Extremely

high magnification is needed in order to determine

whether these setae are naked. There are, however, other characters more easily seen by which

the last two species may be separated from A. hales.

A. cuneipes differs from the new species by the

claws on the second antenna being shorter than

the greater length of the fourth segment and in

having a wedgelike inner expansion on leg 5 in the

female. A. varirostratus is unlike A. hales in showing sexual dimorphism in the rostrum and in having hairs on the inner margin of the basis of leg 4.

Acanthomolgus hians (Humes and Ho, 1968)

Lkhomolgus hians Humes and Ho, 1968d, pp. 719-725, figs.

90-108 [from the alcyonacean Siphonogorgia pichoni Verseveldt, region of Nosy B£, Madagascar; host originally

reported as Siphonogorgia pendula Studer].
106

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NEW RECORDS (species mixed with Acanthomolgus longispinifer and all from Siphonogorgia pichoni, collected by AGH).—1,266 9 9 , 713 tfo",

and 260 copepodids, in 17 m, near black buoy between Pte. Ambarionaomby, Nosy Komba, and Pte.

de Tafondro, Nosy B£, Madagascar, 5 August 1967;

491 $ $ , 199 0*0*. and 46 copepodids, in 25 m,

Tany Kely, a small island south of Nosy Be, Madagascar, 14 August 1967.

Acanthomolgus longispinifer (Humes and Ho,

1968)

Lichomolgus longispinifer Humes and Ho, 1968d, pp. 715—

719, figs. 69-89 [from the alcyonacean Siphonogorgia pichoni Verseveldt, Nosy Be*. Madagascar; host originally reported as Siphonogorgia pendula Studer].

NEW RECORDS (species mixed with Acanthomolgus hians and all from Siphonogorgia pichoni, collected by AGH).—1,266 ? $ . 713 o" d, and 260

copepodids, in 17 m, near black buoy between Pte.

Ambarionaomby, Nosy Komba, and Pte. de Tafondro, Nosy Be, Madagascar, 5 August 1967; 491 9 9 ,

199 o* o*» and 46 copepodids, in 25 m, Tany Kely,

a small island south of Nosy Be, Madagascar, 14

August 1967.

Acanthomolgus plantei, new species

Ficuus 59-61

TYPE MATERIAL.—418 9 9 , 401 tfo*. and 219

copepodids from a single colony of the alcyonacean

Umbellulifera striata (Thompson and Henderson),

in 47 m, 13°29/ S, 48°06' E, southwest of Nosy Be,

Madagascar, 24 July 1967, collected by R. Plante.

Holotype $, allotype, and 770 paratypes (390

$ 9 , 380 o* cf) deposited in USNM, the remaining

paratypes and the copepodids in the collection of

AGH.

OTHER MATERIAL EXAMINED (all from Umbellulifera striata).—75 9 9 , 31 tf Q\ and 9 copepodids

from single colony, in 17 m, pass at Pte. Lokobe,

Nosy Be, 23 August, 1967; 1 9 , 3 tf tf from single

colony, in 17 m, pass at Pte. Lokobe, 23 August

1967; 4 9 9 , 22 copepodids from four colonies, in

17 m, pass at Pte. Lokobe, 25 August 1967; and

97 9 9 , 32 o* o*» and 9 copepodids from two colonies, in 17 m, pass at Pte. Lokobe, 27 August 1967.

All collected by AGH.

FEMALE.—The body (Figure 59a) has a moderately broad prosome. The length is 0.91 mm (0.81-

0.99 mm) and the greatest width 0.43 mm

(0.42-0.45 mm), based on ten specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.44:1. The ratio of the length of the

prosome to that of the urosorae is 2.48:1.

The segment of leg 5 (Figure 596) is 78 x 127 n.

There is no clearly identifiable ventral intersegmental sclerite between this segment and the genital segment. The genital segment is 109 x 117 n,

slightly wider than long, its anterior three-fourths

with rounded lateral margins in dorsal view. The

areas of attachment of the egg sacs are located

dorsolaterally at about the middle of the segment.

Each area (Figure 59c) bears two naked setae, 6 n

and 10 n, with a small pointed process between

them. The cuticular fold medial to the larger seta

is enlarged to form a beaklike process, much larger

than in A. hales. The three postgenital segments

are 33 x 72 i», 26 x 68 n, and 33 x 67 n from anterior to posterior.

The caudal ramus (Figure 59d) is only a little

longer than wide, 26 x 24 n. The dorsal seta is

22 u> the outer lateral seta 35 n, the outermost

terminal seta 44 i», and the innermost terminal

seta 88 n, all of them naked. The two long median

terminal setae are 166 i* (outer) and 255 t» (inner),

both with extremely short lateral spinules and both

inserted between dorsal (unornamented) and ventral (with a marginal row of spinules) flaps.

The body surface is ornamented with hairs (sensilla) as indicated in Figure 59a,b. The posteroventral margin of the anal segment bears a row of

spinules on both sides.

The egg sac (Figure 59a) is elongated oval, 418

x 176 K, reaches far beyond the caudal rami, and

contains many eggs 50 n in diameter. (Egg sacs

transferred from 70 percent ethyl alcohol to lactic

acid disintegrated slowly, releasing the individual

eggs.)

The rostrum resembles that of A. hales.

The first antenna (Figure 59*) is 312 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 26 (55 n

along the anterior margin), 94, 20, 36, 46, 35, and

26 n respectively. The formula for the armature is

the same as in A. hales. All the setae are naked.

The second antenna (Figure 59/) is 4-segmented.

The first two segments are relatively short and ro-
NUMBER 127

FIGURE 59.—Acanthomolgus plantti, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (c); e, first antenna,

anterodorsal (D) ; /, second antenna, anterior (E) ; g, labrura, ventral (F) ; h, mandible, posterior

(F) ; i, first maxilla, anterior (c); /, terminal segment of second maxilla, anterior (c). Scale:

A, 05 mm; B, 0.02 mm; c, F, 0.05 mm; D, E, 0.1 mm.
108 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

bust, and their setae are very short; the first segment is ornamented with a few small spines near

the seta. The third segment bears two setae and a

vestige of a spiniform element which is hyaline and

only 2 n long. The fourth segment is 90 n along

the outer edge, 58 n along the inner edge, and

22.5 n wide, bearing two long unequal terminal

claws, the stouter claw 70 n along its axis, the more

slender claw 89 n. Near the insertion of the claws

are five small hyaline elements as in A. hales.

The labrum (Figure 59g) has two broad posteroventral lobes, less truncated than in A. hales.

The mandible (Figure 59/i) is similar to that of

A. varirostratus (Humes and Ho). The paragnath

is like that of A. hales. The first maxilla (Figure

59i) resembles that of A. exilipes (Humes and Ho).

The second maxilla also is much like that of A.

exilipes, but the spinules on the inner distal spine

are longer (Figure 59;). The maxilliped (Figure

60a) resembles that of A. hales but is a little more

elongated.

The ventral area between the maxillipeds and

the first pair of legs is like that in A. hales.

Legs 1-4 (Figure 606-^) have the same spinal and

setal formula as in other species of the genus. The

inner coxal element of leg 4 is reduced to a small

naked setule 6 n long. The inner margin of the

basis of leg 4 is smooth instead of bearing a row

of hairs as in legs 1-3. The exopod of leg 4 is

136 »* long. The endopod of leg 4 has hairs along

the outer margins of both segments and a few short

spinules on the inner margin of the second segment. The first segment is 36 x 28 ]i (not including

the processes) and its inner distal spine is 44 n and

finely barbed. The second segment is 68 n long

(including the terminal spiniform processes), 25 n

in greatest width proximally, and 16.5 n in least

width distally; the outer spine is 28 n, the inner

spine 55 n, both barbed.

Leg 5 (Figure 60/) has a free segment about

twice as long as wide in greatest dimensions. The

length is 79 n and the greatest width at the proximal inner expansion is 37 I*. The outer surface

of the segment bears relatively long spines. The

two terminal elements are 39 ji and naked, and

66 n with a unilateral fringe. The seta on the

body near the insertion of the free segment is 34 n

and naked.

Leg 6 is probably represented by the two setae

near the areas of attachment of the egg sacs (Figure

59c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.

MALE.—The body (Figure 60g) is more slender

than in the female. The length is 0.79 mm (0.72-

0.86 mm) and the greatest width 0.28 mm (0.26-

0.30 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome

is 1.78:1. The ratio of the length of the prosome

to that of the urosome is 1.73:1.

The segment of leg 5 (Figure 60A) is 39 x 70 »i.

There is no ventral intersegmental sclerite. The

genital segment is longer than wide, 162 x 138 I*.

The four postgenital segments are 28 x 53 n, 21

x 51 n, 11 x 50 n, and 23 x 51 n from anterior to

posterior.

The caudal ramus is similar to that of the female, but smaller, 23 x 21 n.

The surface of the body is ornamented with hairs

as in the female.

The rostrum is like that of the female.

The first antenna has the same armature as in

the male of A. astrictus but with three additional

aesthetes. The second antenna (Figure 60i) shows

sexual dimorphism. The setae on the first two segments are longer than in the female and both segments bear a few short spines. The spiniform

element on the third segment is well formed (1 2 M

long) and has a few minute barbules on one edge.

The fourth segment is 88 n along the outer edge,

63 n along the inner edge, and 23 n wide. The

terminal claws are 59 n and 76 n.

The labrum (Figure 60;) has a form slightly different from that in the female.

The mandible, paragnath, first maxilla, and second maxilla are like those in the female. The

maxilliped (Figure 61a) resembles that of A. hales.

The claw is 172 \i along its axis and has on its

proximal area a few striations in addition to the

two unequal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are like those of the female, except for

the last segment of the endopod of leg 1 (Figure

616), which is 1,1,4. Slight sexual dimorphism occurs on the last segment of the endopod of leg 2

(Figure 61c) where the three spines from outer to

inner are 19, 26, and 26 n (in the female 20, 15,

and 20 n); and on the last segment of the endopod
NUMBER 127 109

FIGURE 60.—Acanthomolgus plantei, new species. Female: a, maxilliped, postero-inner (F) ; b, leg

1 and intercoxal plate, anterior (D) ; c, endopod of leg 2, anterior (D) ; d, endopod of leg 3,

anterior (D) ; e, leg 4, anterior (D) ; /, leg 5, dorsal (F) . Male: g, dorsal (A) ; h, urosome, dorsal

(B) ; i, second antenna, anterior (E) ; ;, labrum, ventral (F) . Scale: A, 0.5 mm; B, 0.2 mm; D, E,

0.1 mm; r, 0.05 mm.
no

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 61.—Acanthomolgus plantei, new species. Male: a, maxilliped, ventro-outer (D); b,

endopod of leg 1, anterior (F) ; c, third segment of endopod of leg 2, anterior (e); d, third

segment of endopod of leg 3, anterior (E) ; e, leg 5. dorsal (c). Scale: c, F, 0.05 mm; o, l, 0.1 mm.

of leg 3 (Figure 6Id) where the spines are 24, 23,

and 34 u (in the female 26, 19, and 27.5 v). The

endopod of leg 4 is like that in the female.

Leg 5 (Figure 61^) has a moderately short free

segment, 22 x 8 u, without a proximal expansion.

The dorsal surface bears a few broad spines. The

two terminal elements are 19 M. and 31 n.

Leg 6 (Figure 60A) is like that in A. hales, its

two setae being 23 n and 31 n.

The spermatophore, attached to the female in

pairs and resembling that of A. hales, is 117 x 55 n

(without the neck).

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—This species is named for Dr.

Raphael Plante, Centre ORSTOM de Nosy Be",

Madagascar, who, by SCUBA diving, collected the

colony of Umbellulifera striata on which the type

specimens were found.

REMARKS.—Acanthomolgus plantei may be separated from all other species in the genus except

A. hales on the basis of the ornamentation of the

two long ramal setae, consisting of extremely short

lateral spinules. (As mentioned above, eight species have long spinules on these setae, and in two

species the setae are naked.) The new species may

be distinguished from A. hales by the shape of the

genital segment and leg 5 in the female, and by

the nature of the ornamentation of leg 5 and the

degree of sexual dimorphism in the second antenna

of the male.

The vestigial nature of the spiniform element on

the third segment of the second antenna of the

female of A. plantei is a feature found in no other

species of Acanthomolgus.

Acanthomolgus telestophilus (Humes and Ho,

1968)

Lichomolgus telestophilus Humes and Ho, 1968d, pp. 725-

730, figs. 109-127 [from the telestacean Telesto arborea

Wright and Studer, Antany Mora, Isles Radama, Madagascar, 14°06'10" S, 47°45'1O" E.

Acanthomolgus varirostratus (Humes and Ho,

1968)

Lichomolgus varirostratus Humes and Ho, 1968b, pp. 2-7,

figs. 1-21 [from the alcyonaceans Dendronephthya mucronata (Putter), Dendronephthya regia Verseveldt, Dendronephthya stocki Verseveldt, and Dendronephthya koellikeri

Kiikenthal, region of Nosy B<\ Madagascar],

NEW RECORDS (all from Madagascar, collected by

AGH).—From Dendronephthya mucronata: 8 $ $,
NUMBER 127 111

7 d <$ > from single colony, in 4 m, Antsamantsara,

Nosy Be\ 9 June 1967; 6 9 9, 2 o* c?» and 2 copepodids, from single colony, in 2 m, off Ampombilava,

Nosy Be\ 7 July 1967; 2 $ $, 4 tf <f, and 3 copepodids, from single colony, in 24 m, Bane des Freres,

Isles Mitsio, near Nosy Be, 12°58' S, 48°28' E, 18

August 1967; many specimens, from four colonies,

in 10 m, Rocher du N.E., Nosy Be\ 19 August 1967;

many specimens, from two colonies, in 10 m,

Rocher du N.E., Nosy Be", 19 August 1967; many

specimens, from single colony, in 25 m, north of

Ankazoberavina, near Nosy Be, 13°27.6' S, 47°58.2'

E, 24 August 1967; and several specimens, from

single color.,, in 24 m, north of Ankazoberavina,

near Nosy Be\ 25 August 1967.

From Dendronephthya regia: 28 99 , 11 $ £,

and 52 copepodids, from single colony, in 25 in,

north of Ankazoberavina, near Nosy Be", 13°27.6' S,

47°58.2' E, 24 August 1967.

From Dendronephthya stocki: many specimens,

from single colony, in 25 m, north of Ankazoberavina, near Nosy Be\ 13°27.6' S, 47°58.2' E, 24

August 1967.

From Dendronephthya sp.: many specimens, from

part of single colony, in 27 m, Bane des Freres,

Isles Mitsio, near Nosy Be, 12°58' S, 48°28' E, 17

August 1967.

NEW HOSTS (all from Madagascar, collected by

AGH).—From Dendronephthya cirsium Kiikenthal: 7 9 9, 13 cfcf, and 9 copepodids, from

single colony, in 35 m, near Nosy Be\ 13°15'5O" S,

48°08'35" E, 29 August 1967.

From Dendronephthya lokobeensis Verseveldt:

many specimens, from single colony, in 15 m, Pte.

Lokobe, Nosy Be, 13 June 1967; 35 9 9, 14 tf o*.

from seven colonies, in 18 m, in pass at Pte. Lokobe, Nosy Be, 30 August 1967.

From Dendronephthya speciosa Kukenthal: 10

9 9» 5 cf 0* t and 7 copepodids, from two colonies,

in 10 m, Rocher du N.E., Nosy Be\ 19 August 1967.

From Stereonephthya cordylophora Verseveldt:

many specimens, from single colony, in 24 m, north

of Ankazoberavina, near Nosy Be", 13°27.6' S,

47°58.2' E, 25 August 1967.

Acanthomolgus verseveldii (Humes and Ho, 1968)

Lichomolgus verseveldti Humes and Ho, 1968d, pp. 694-702,

figs. 1-26 [from the alcyonacean Heteroxenia elisabethae

Kdlliker, region of Nosy Be, Madagascar].

NEW ALCYONACEAN HOSTS (collected by AGH).—

28 9 9, 26 ^ cf, and 68 copepodids from Xenia

lepida Verseveldt, in 10 m, Pte. Lokobe, Nosy Be\

Madagascar, 18 July 1967; 3 9 9, 3 tf d\ and 4

copepodids from Heteroxenia fuscescens (Hemprich and Ehrenberg), in 20 m, Bane de Cinq Metres, near Nosy Be, Madagascar, 6 August 1967.

Genus Amarda Humes and Stock, 1972

DIAGNOSIS.—Body transformed. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six short setae. Rostrum subtriangular but rounded posteroventrally. First antenna 7-segmented, in the female with the formula

3 or 4, 12, 6, 3, 4 -j- 1 aesthete, 2 + 1 aesthete,

and 7 + 1 aesthete; in the male 3 or 4, 12 + 2

aesthetes, 6, 3 -f- 1 aesthete, 4 -\- 1 aesthete,

2 -f 1 aesthete, and 7 -\- 1 aesthete. Second antenna 3-segmented, with the armature 1, 1, 3 -jI, there being a single terminal claw.

Labrum with a median cleft separating the two

lobes. Mandible with its basal area having on its

convex side a slight rounded swelling followed

by a prominent winglike process and then by

a serrated fringe; its concave side beyond the indentation having two lobes, one with small serrations, the other with large serrations; lash relatively

short and smooth. First maxilla with four elements.

Second maxilla with the armature on the second

segment much reduced; lash short and reflexed.

Maxilliped in the female 3-segmented with a somewhat pointed tip; in the male 4-segmented (assuming the proximal part of the claw to be a fourth

segment) and unusual in lacking rows of spinules

on the second segment.

Legs 1-3 similar in both sexes with reduced armature. Legs 1, 2 with 3-segmented exopods and

either 2- or 3-segmented endopods. Leg 3 with a

2-segmented exopod (0-0;I,II orO-0;II) but lacking an endopod. Leg 4 absent in both sexes. Leg

5 in both sexes with a small free segment not clearly

delimited from the body bearing two terminal

setae. Leg 6 in the female represented by the two

setae and the spiniform process near the area of

attachment of each egg sac; in the male by a

posteroventral flap on the genital segment bearing

two setae.

Other features as in the species described below.

Associated with madreporarian corals.
112

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

TVPE-SPECIES.—Amarda cultrata, new species.

ETYMOLOGY.—The generic name is an anagram

of Radama I, a king of Madagascar who reigned

1810-1828. Gender feminine.

Key to Species of Amarda

Endopods of legs 1 and 2 two-segmented; leg 3 exopod with the formula 0-0-;I.II; caudal ramus

with the two median terminal setae broad and bladelike A. cultrata

Endopods of legs 1 and 2 three-segmented; leg 3 exopod with the formula 0-0;II; caudal

ramus with the two median terminal setae slender A. compta

Amarda cultrata, new species

FIGURES 62-64

TYPE MATERIAL.—2 $ $ , 5 rf o* from single

colony of Favia sp., in 2 in, Ambariotelo, a small

island almost between Nosy Komba and Nosy Be,

northwestern Madagascar, 23 May 1964, collected

by AGH. Holotype °., allotype, and 5 paratypes

(1 9.4 <td) deposited in USNM.

OTHER MATERIAL EXAMINED (all from Favia

sp.).—1 <j> from several colonies, in 2 m, Pte. a

la Fievre, Nosy Be\ Madagascar, 26 December 1963;

1 $, 1 c? from single colony, in 2 m, Ambariotelo,

23 May 1964; 2 ? $, 5 <$ <?, and 1 copepodid from

single colony, in 2 m, off Ampombilava, Nosy Be,

26 September 1964; and 1 ? , 1 tf from single

colony, in 2 m, east of Ambariobe, a small island

almost between Nosy Komba and Nosy Bi, 4 October 1964. All collected by AGH.

FEMALE.—The body (Figure 62a,b) has a moderately broad and flattened prosome. The length

(excluding the setae on the caudal rami) is 1.59 mm

(1.49-1.68 mm) and the greatest width 0.75 mm

(0.68-0.79 mm) based on four specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.29:1. The ratio of the length of the

prosome to that of the urosome is 1.51:1. The

segment of leg 1 is distinctly set off dorsally from

the cephalosome by a transverse furrow. The

epimeral areas of the segments of legs 1-3 are

prominent, but the segment which ordinarily

would bear leg 4 (absent in this species) lacks

such epimera.

The segment of leg 5 (Figure 62c) is 86 x 285 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 177 x 280 n in greatest dimensions in

dorsal view, broadest in its anterior half. The areas

of attachment of the egg sacs are situated dorsolaterally at the middle of the segment. Each area

(Figure 62d) bears two minute naked setae, 8 n

and 7 n, and a small spinous process. The three

postgenital segments are 109 x 169 n, 86 x 139 |i,

and 133 x 127 M^ from anterior to posterior. The

posteroventral border of the anal segment is

smooth.

The caudal ramus (Figure 62*) is 85 x 32 n, or

2.66 times longer than wide. The outer lateral

seta is 30 n, the dorsal seta 28 n, the outermost

terminal seta 34 n, and the innermost terminal

seta 36 n, all of them naked. The two median

terminal setae are relatively short and broad, the

outer 53 n, the inner 82 n, both naked.

The body surface has numerous minute hairs

(sensilla) and refractile points.

The egg sac is unknown, but a few eggs were

attached to one female. The eggs are 115-130 »i

in diameter.

The rostrum (Figure 62/) has a subtriangular

but rounded posteroventral margin. Between the

rostrum and the labrum there is a knoblike area

protruding ventrally.

The first antenna (Figure 62g) is 297 » long. It

is considered as 7-segmented, but the last two segments appear to be incompletely separated. The

lengths of the segments (measured along their posterior nonsetiferous margins) are 33 (73 n along

the anterior margin), 96, 29, 36, 31, 21, and 11 |i

respectively. The formula for the armature is 3,

12, 6, 3, 4 -f 1 aesthete, 2 -f 1 aesthete, and

7 -f- 1 aesthete. All the setae are naked.

The second antenna (Figure 62h) is 3-segmented.

The first two segments bear a short inner seta. The

third segment (representing a fusion of two segments) bears on its inner side two small setae and

the suggestion (vaguely defined) of a very minute

third element. Terminally the segment has a

digitiform extension with a few minute spinules

and bears three small setae and a sinuous terminal

claw 39 n along its axis (Figure 62i).
NUMBER 127

FIGURE 62.—A mar da cultrata, new species. Female: a, dorsal (A) ; b, ventral (A) ; c, urosome,

dorsal (B) ; d, area of attachment of egg sac, dorsal (c); e, caudal ramus, dorsal (c); /, rostrum,

ventral (D) ; g, first antenna, with arrows indicating positions of additional aesthetes in male,

anterior (E) ; h, second antenna, posterior (t); i, tip of second antenna, anterior (F); ;', labrura,

ventral (c). Scale: A, 05 mm; B, D, 02 mm; c, E, 0.1 mm; F, 0.05 mm.
114 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The labrum (Figure 62;) has two, weakly cleft,

posteroventral lobes.

The mandible (Figure 63a) has on the concave

side of the base a deep indentation followed by

two transverse lobes, one with a finely serrated

margin, the other with a coarsely serrated margin;

on the convex side there is a slight rounded swelling and distally a very prominent winglike process

followed by a serrated hyaline fringe. The lash is

relatively short and smooth. Paragnaths could not

be identified with certainty. The first maxilla (Figure 63b) consists of a single segment bearing four

elements. The second maxilla (Figure 63c) is 2-

segmented. The first segment is large and unornamented. The second segment is much modified,

with only one minute seta on its posterior surface

and with its stout short terminal lash reflexetl and

bearing a row of prominent teeth. The maxilliped

(Figure 63d) is 3-segmented. The first segment is

unarmed. The second segment bears two very unequal minute inner setae and a row of extremely

small spines distally. The third segment bears a

blunt seta and a row of small spines near the

pointed tip.

The ventral area between the maxillipeds and

the first pair of legs (Figure 63e) is not protuberant.

A fine line connects the bases of the maxillipeds.

The region immediately in front of the intercoxal

plate of leg 1 lacks the elongated sclerite seen in

most lichomolgids.

Leg 1 (Figure 63/) and leg 2 (Figure 63g) have

3-segmented exopods and 2-segmented endopods.

Leg 3 (Figure 63/i) has a 2-segmented exopod but

lacks an endopod entirely. Leg 4 is absent.

The armature of the legs may be expressed as

follows:

Px coxa 0-0 basis 1-0 exp 1-0; 1-0; IIIJ.l

enp 0-0; 1,2

P, coxa 0-0 basis 1-0 exp 1-0; 1-0; III.U

enp 0-0; IJI

P, coxa 0-0 basis 1-0 exp 0-0; 1,11

enp absent

Pt absent

The coxa in all three legs lacks an inner seta.

Leg 4 is absent (Figure 626), but there is a small

internal sclerotization indicating its former position.

Leg 5 (Figure 63») has a small unornamented

segment, 15.5 x 11 n, which is not clearly separated

from the body. It bears two terminal naked setae,

40 it and 45 n. The seta on the body near the segment is 50 n and naked.

Leg 6 is represented by the two setae on the area

of attachment of each egg sac (Figure 62d).

The color in life in transmitted light is opaque,

the ventrally placed eye red, the intestine light

brown, the ovary gray, the extruded eggs black.

MALE.—The body (Figure 64a) resembles in general form that of the female. The length (not including the ramal setae) is 1.31 mm (1.18-1.37

mm) and the greatest width 0.59 mm (0.54-0.63

mm), based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.19:1. The ratio of the length of the prosome to

that of the urosome is 1.14:1.

The segment of leg 5 (Figure 646) is 65 x 280 n.

There is no ventral intersegmental sclerite. The

genital segment is 146 x 264 n, much wider than

long. The four postgenital segments are 83 x 149 i*,

88 x 133 |i, 75 x 115 n, and 112 x 114 n from anterior to posterior.

The caudal ramus resembles that of the female.

The rostrum is like that of the female.

The first antenna is similar to that of the female

but has three additional aesthetes so that the formula is 3, 12 + 2 aesthetes, 6, 3 -f- 1 aesthete,

4 + 1 aesthete, 2 -f 1 aesthete, and 7 + 1 aesthete.

The second antenna, labrum, mandible, paragnath (?), first maxilla, and second maxilla are like

those of the female.

The maxilliped (Figure 64c) is slender and 4—

segmented (assuming that the proximal part of the

claw represents a fourth segment). The first segment is unarmed. The second bears two inner setae

and a few retractile areas but lacks the rows of

spinules commonly seen in other lichomolgids. The

small third segment is unarmed. The claw is 120 n

along its axis, lacks a terminal lamella and any indication of division, and bears two very unequal

proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-3 have the same segmentation and armature as in the female. The outer margin of the

second segment of the endopods of leg 1 (Figure

64d) and leg 2 may be smooth or may have 1-3

spiniform processes which are larger than those in

the female. Leg 4 is absent as in the female.
NUMBER 127

FIGURE 63.—Amarda cultrata, new species. Female: a, mandible, posterior (F) , b, first maxilla,

posterior (F) ; c, second maxilla, anterior (F) ; d, maxilliped, anterior (F) ; e, area between

maxillipeds and first pair of legs, ventral (E) ; /. leg 1 and intercoxal plate, anterior (c); g, leg

2 and intercoxal plate, anterior (c); h, leg 3 and intercoxal plate, anterior (c); i, leg 5, dorsal

(F) . Scale: c, E, 0.1 mm; F, 0.05 mm.
116

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

a

FIGURE 64.—Amarda cultrata, new species. Male: a, dorsal (A) ; b, urosomc, dorsal (B); c,

maxilliped, inner (c); it. endopod of leg 1, anterior (c). Scale: A, 0.5 mm; B, 0.2 mm; c, 0.1 mm.

Leg 5 is similar to that of the female, though the

segment is somewhat smaller and less clearly set

off from the body.

Leg 6 (Figure 646) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 34 \i and 39 n.

The spermatophore was not observed.

The color in life in transmitted light is opaque,

the eye red, the intestine brown.

ETYMOLOGY.—The specific name cultrata (Latin,

shaped like a knife) alludes to the two broad,

bladelike setae on the caudal ramus.

Amarda co.ipta, new species

FIGURES 65, 66

TYPE MATERIAL.—4 99 , 4 tf tf from single

colony of Faxria sp. (the same colony in which A.

cultrata was found), in 2 m, off Ampombilava,

Nosy Be, northwestern Madagascar, 26 September

1964, collected by AGH. Holotype $, allotype,

and 2 paratypes (1 9, 1 <?) deposited in USNM,

the remaining paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED (also from Faxria

sp.).—2 9?, 4 <3<f, and 1 copepodid from single

colony (the same colony in which A. cultrata was

found), in 2 m, east of Ambariobe, a small island

nearly between Nosy Komba and Nosy B£, Madagascar, 4 October 1964, collected by AGH.

FEMALE.—The body (Figure 65a,b) has a large,

broad, thickened prosome. The length (without

the ramal setae) is 1.12 mm (1.06-1.20 mm) and

the greatest width 0.60 mm (0.57-0.62 mm), based

on four specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.17:1. The

ratio of the length of the prosome to that of the

urosome is 2.46:1. The segment of leg 1 is distinctly delimited from the cephalosome by a dorsal

transverse furrow. The segment which ordinarily

would bear leg 4 (absent in this species) lacks

projecting epimera.

The segment of leg 5 (Figure 65c) is 65 x 229 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 117 x 205 n. In dorsal view the lateral

borders of the segment are rounded; in lateral

view (Figure 656) the dorsal surface is somewhat
NUMBER 127 11'

FIGURE 65.—Amarda cotnpta, new species. Female: a, dorsal (A) ; b, lateral (A) ; c, urosome,

dorsal (B) ; d, area of attachment of egg sac, dorsal (c); e, caudal ramus, dorsal (c); /, first

antenna, posterior (D) ; g, leg 1, anterior (D) ; h, leg 2, anterior (D) ; i, leg 3, anterior (c);

j , leg 5, dorsal (E) . Scale: A, 05 mm; B, 02 mm; c, E, 0.05 mm; D, 0.1 mm.
118 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

protruded. The areas of attachment of the egg sacs

are located dorsally behind the middle of the segment. Each area (Figure 65d) bears two small

naked setae, 6 n and 9 n, and a small spinous process. The three postgenital segments are 47 x 120 n,

44 x 104 n, and 67 x 99 n from anterior to posterior.

The posteroventral border of the anal segment is

smooth.

The caudal ramus (Figure 65e) is 44 x 23 n, or

about 1.9 times longer than wide. It bears six

naked setae, all of them slender and shorter than

the ramus itself.

The body surface bears many small round refractile areas (Figure 65a,b).

The egg sac is unknown.

The rostrum resembles that of the male, shown

in Figure 66c.

The first antenna (Figure 65/) is 233 n long. The

sixth and seventh segments are incompletely separated as in the preceding species. The lengths of

the segments (measured along their posterior nonsetiferous margins) are 23 (61 n along the anterior

margin), 72, 17, 27, 27, 20, and 9 n respectively.

The formula for the armature is 4, 12, 6, 3, 4 -f- 1

aesthete, 2 + 1 aesthete, and 7 + 1 aesthete. All

the setae are naked.

The second antenna, labrum, mandible, paragnath (?), first maxilla, second maxilla, maxilliped,

and ventral area between the maxillipeds and the

first pair of legs are similar to those in A. cultrata.

Leg 1 (Figure 65g) and leg 2 (Figure 65h) have

3-segmented exopods and endopods. Leg 3 (Figure 65i) has a 2-segmented exopod but lacks an

endopod. Leg 4 is absent.

The armature of the legs is as follows:

Px coxa 0-0 basis 1-0 exp 1-0; 1-0; 111,1,2

enp 0-0; 0-0; 1,2

P, coxa 0-0 basis 1-0 exp 1-0; 1-0; 111,1,2

enp 0-0; 0-0; 1,11

P, coxa 0-0 basis 1-0 exp 0-0; II

enp absent

P« absent

As in A. cultrata, the coxa in all three legs lacks

an inner seta and leg 4 is absent.

Leg 5 (Figure 65/) has a small unornamented

segment only 8 n long which is not clearly delimited from the body.

Leg 6 is represented by the two setae on the area

of attachment of each egg sac (Figure 65d).

The color in life resembles that of A. cultrata.

MALE.—The body (Figure 66a) resembles in outline that of the female. The length (not including

the setae on the caudal rami) is 0.85 mm (0.81-0.92

mm) and the greatest width 0.42 mm (0.41-0.43

mm), based on four specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.22:1. The ratio of the length of the prosome to

that of the urosome is 2.49:1.

"The segment of leg 5 (Figure 66b) is 55 x 213 >i.

There is no ventral intersegmental sclerite. The

genital segment is 91 x 208 n, much wider than

long. The four postgenital segments are 42 x 107 n,

42 x 94 \i, 34 x 87 n, and 62 x 78 n from anterior

to posterior.

The caudal ramus is like that of the female.

The rostrum (Figure 66c), like that of the female, has a broad trilobate posteroventral margin.

In the midline between the rostrum and the labrum there is a large ventrally protruding knob.

The first antenna has three additional aesthetes,

as in the male of A. cultrata.

The second antenna, labrum, mandible, paragnath (?), first maxilla, and second maxilla resemble

those of the female. The maxilliped, its claw 108 n

along its axis, is similar to that in the male of A.

cultrata. The ventral area between the maxillipeds

and the first pair of legs, legs 1-3, and leg 5 (with

its segment even smaller than in the female) resemble those of the female. Leg 4 is absent as in

the female.

Leg 6 (Figure 66b) is similar to that of the male

of A. cultrata.

The spermatophore was not observed.

The color in life resembles that of A. cultrata.

ETYMOLOGY.—The specific name compta (Latin,

ornamented) refers to the small refractile areas ornamenting the body surface.

REMARKS.—Amarda compta may be readily distinguished from A. cultrata by its small size, the

refractile areas scattered over the body surface, the

nature of the two median terminal setae on the

caudal rami (slender rather than broad), the number of setae on the first segment of the first antenna (four rather than three as in A. cultrata),

the segmentation of the endopod in legs 1 and 2

(3-segmented instead of 2-segmented as in A. cultrata), and the number of spines on the second segment of the exopod in leg 3 (two rather than three

as in A. cultrata).
NUMBER 1X7 119

FIGURE 66.- A ma rda compta, new species. Male: a, dorsal (A); b, urosome, dorsal (B) ; c,

rostrum, ventral (D) . Scale: A, 05 mm; B, 02 mm; D, 0.1 mm.

Amur da compta is similar to A. cultrata in important ways, such as the transformed body, the

segmentation of the first antenna (with the last

two segments incompletely separated), the 3-segmented second antenna, the nature of the labrum

and mouthparts, the absence of the endopod in leg

3, the absence of leg 4, and the reduction of leg 5.

The host of the two species is the same, Favia sp.

Genus Anchimolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ram us with six moderately short setae.

Rostrum weakly developed. First antenna 7-segmented, in the female with the formula 4, 13, 6,

3, 4 + 1 aesthete, 2 -f 1 aesthete, and 7 + 1

aesthete; in the male with 4, 13 + 2 aesthetes, 6,

3 + 1 aesthete, 4+ 1 aesthete, 2+ 1 aesthete,

and 7+ 1 aesthete. Second antenna 4—segmented,

with the formula 1, 1, 3, and I + a few minute

elements, there being only a single short terminal

claw.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal area beyond the

indentation having on its concave side two small

digitiform lobes followed by a serrated fringe and

on its concave edge two lobes with marginal spinules; lash long. Paragnath a small hairy lobe.

First maxilla with three or four elements. Second

maxilla of the usual lichomolgid type. Maxilliped

in the female 3-segmented with a pointed tip; in

the male 4-segmented (regarding the proximal part

of the claw as a fourth segment).

Legs 1-^1 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

Key to Species of the Genus Anchimolgus

FEMALES

Free segment of leg 5 small, 36 x 17 pi, much shorter than genital segment; caudal ramus about

6.1:1 A. digitatus

Free segment of leg 5 elongated, 203 x 30 \i, longer than genital segment; caudal ramus 2.65:1....

A. protixipes

MALES

Genital segment a little wider than long; caudal ramus about 4.3:1 A. digitatut

Genital segment longer than wide; caudal ramus 2:1 A. pnlixipes
120

the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1;II, the seta being feathered.

Leg 1 endopod of the male with the third segment

1,1,4 instead of 1,5 as in the female. Leg 5 with a

free segment bearing two setae. Leg 6 in the female represented by the two setae and the process

near the area of attachment of each egg sac; in the

male by the ventral posterolateral flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with madreporarians.

TYPE-SPECIES.—Anchimolgus digitatus (Humes

and Ho).

ETYMOLOGY.—The name is a combination of the

Greek words avxi (near), suggesting the affinity of

this genus with other lichomolgids, and

Gender masculine.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Anchimolgus digitatus (Humes and Ho, 1968)

FIC.URE 67

l.ichomolgus digitatus Humes and Ho, 1968a, pp. 357-359,

figs. 25-52 [from the hard coral Goniopora sp.. Nosy BeMadagascar].

NEW HOST.—1 ° , 1 d1 from Favia sp., in 5 m,

Nosy Taolankena, near Nosy Be, Madagascar, 24

July 1964, collected by AGH.

NEW RECORD.—8 $9. 7 cf o*» and 3 copepodids

from Goniopora sp., in 17 m, near black buoy

north of Pte. Ambarionaomby, Nosy Komba, near

Nosy Be, Madagascar, 5 August 1967, collected by

AGH.

FIGURE 67.—Anchimolgus digitatus (Humes and Ho). Female: a, dorsal; b, second antenna; c,

mandible; d, second maxilla; e, maxilliped; /, leg 4 and intercoxal plate. Male: g, dorsal. (From

Humes and Ho, 1968a, figs. 25, 31, 33, 36, 42, 44.) Length of female 1.89 mm, of male 155
NUMBER 127 121

Anchimolgus prolixipes (Humes and Ho, 1968)

Lichomolgus prolixipes Humes and Ho, 1968a, pp. 359-362,

figs. 53-78 [from the hard corals Porites sp. cf. andrewsi

Vaughan, Porites sp. cf. nigrescens Dana, and Porites (Synaraea) sp., region of Nosy Be, Madagascar].

NEW RECORD.—28 $ c», 14 cf cf. and 16 copepodids from Porites sp., in 2 m, off Ampombilava,

Nosy B£, Madagascar, 5 June 1967, collected by

AGH.

Genus Andrianellus Humes and Stock, 1972

DIAGNOSIS.—Body modified. Urosome in the female 5-segmented, in the male 6-segmented. Caudal raimis with six relatively short setae. Rostrum

with a weakly defined posteroventral margin. First

antenna 7-segmented, in the female with the formula 4, 13, 6, 3, 4 4- 1 aesthete, 2 -f 1 aesthete,

and 7 -f 1 aesthete, in the male with 4, 13 -f- 2

aesthetes, 6, 3 -f 1 aesthete, 4 -f- 1 aesthete, 2 -f1 aesthete, and 7 4- 1 aesthete. Second antenna

4-segmented, with the formula 1,1,3,1, there being

a single small terminal claw.

Labrum with an indentation separating the two

lobes. Mandible with the base divided by an indentation into an unornamented proximal part

and a distal area having on its convex side a slender spiniform process followed by a weakly serrated fringe and on its concave side two lobes, one

with a single row of spinules, the other with two

rows; lash stout and moderately long. First maxilla

with four elements. Second maxilla of the usual

lichomolgid type, but the first segment bearing a

long digitiform process. Maxilliped in the female

3-segmented with a pointed tip, in the male 4 -

segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Formula for the

armature the same in both sexes. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod 0-1 ;I. Leg 5 in both sexes with a free segment

(not clearly delimited from the body) with two

terminal setae. Leg 6 in the female represented by

the two seta near the area of attachment of each egg

sac; in the male by a posteroventral flap on the

genital segment bearing two setae.

Odier features as in the species described below.

Associated with madreporarian corals.

TYPE-SPECIES.—Andrianellus exsertidens, new

species.

ETYMOLOGY.—The generic name is a combination of "Andrian-," from Andrianampouinimerina,

the first king of Madagascar who reigned 1787—

1810, and the Latin diminutive suffix "-ellus." Gender masculine.

Andrianellus exsertidens, new species

FIGURES 68-70

TYPE MATERIAL.—18 $ <J>, 10 cf cf fr°ni single

colony of the madreporarian Favia sp., in 2 m, Ambariotelo, a small island nearly between Nosy

Komba and Nosy Be, northwestern Madagascar, 20

July 1967, collected by AGH. Holotype $, allotype, and 20 paratypes (14 $ $, 6 cf cf) deposited

in USNM; 2 paratypes (1 9 , 1 <?) in ZMA; and

the remaining paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED.—2 $ $, 2 cf cf» and

1 copepodid from single colony of the madreporarian Platygyra daedala (Ellis and Solander), in

2 m, Pte. a la Fievre, Nosy Be, 26 December 1963.

FEMALE.—The body (Figure 68a,6) has a somewhat thickened prosome. The length (not including the setae on the caudal rami) is 1.28 mm (1.26-

1.33 mm) and the greatest width 0.4? mm (0.44-

0.49 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome is 1.4:1. The ratio of the length of the prosome to that of the urosome is 1.13:1. The segment

of leg 1 is fused with the cephalosome. The epimera of the segments of legs 1-4 are as illustrated

in Figure 68a.

The segment of leg 5 (Figure 68c) is 62 x 220 \i.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 210 x 234 n, broadest anteriorly and tapered posteriorly. The areas of attachment of the

egg sacs are dorsal in position, situated near the

middle of the segment. Each area (Figure 68d) has

two small naked setae, 8 n and 4 n, and a small

spinous process. The three postgenital segments

are 75 x 120 n, 57 x 96 n, and 68 x 93 n from anterior to posterior. The posteroventral border of

the anal segment bears a row of small spinules on

each side.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

FIGURE 68.—Andrianellus exsertidem, new species. Female: a, dorsal (A) ; b, lateral (A); c,

urosome, dorsal (B) ; d, area of attachment of egg sac, dorsal (c); e, caudal ramus, dorsal (D) ;

/, rostrum, ventral (E) ; g, first antenna, anterodorsal (»); /;, second antenna, posterior (D) ;

i, labrum, ventral (D) ; ;, mandible, posterior (F) . Scale: A, 0J5 mm; B, 02 mm; c, F, 0.05 mm;

D, £, 0.1 mm.
NUMBER 127 123

The caudal ramus (Figure 68?) is elongated, 120

x 31 n, or 3.87 times longer than wide. The outer

lateral seta is 39 n, the dorsal seta 22 n, the outermost terminal seta 42 n, the innermost terminal

seta 52 n, and the two median terminal setae 88 ii

and 120 n (outer and inner respectively), all six

setae being naked. The tips of the two median

terminal setae are blunt.

The body surface has small hairs (sensilla) and

refractile points as in Figure 68a.

The egg sac is unknown.

The rostrum (Figure 68/) is a moderately elongated lobe with a weakly defined posteroventral

margin. Between the rostrum and labrum there is

an oval protrusion.

The first antenna (Figure 68g) is 7-segmented

and slender, 278 jt long. The lengths of the segments (measured along their posterior nonsetiferous margins) are 22 (44 v along the anterior

edge), 75, 30, 41, 40, 26, and 22 n respectively.

The formula for the armature is 4, 13, 6, 3, 4 -j- 1

aesthete, 2 -f- 1 aesthete, and 7 + 1 aesthete. All

the setae are naked.

The second antenna (Figure 68/J) is 4-segmented.

Each of the first two segments bears a small inner

seta. The third segment has three such setae. The

fourth segment bears only a small terminal claw

20 » in length.

The labrum (Figure 68i) has two rounded

posteroventral lobes with somewhat lobate margins.

The mandible (Figure 68;) has on its concave

edge an indentation followed by two transverse

lobes, one with two rows of slender spinules and

the other with a single row of coarser spinules; on

its convex edge there is a slender spiniform process

followed by a weakly serrated hyaline fringe. The

lash is moderately long, stout, and sparsely barbed.

Paragnaths could not be identified with certainty.

The first maxilla (Figure 69a) has four elements.

The second maxilla (Figure 69b) is 2-segmented.

The large first segment has no armature, but bears

a long (60 n), slender, blunt, digitiform process

which may be seen in the whole animal extending

ventrally and anteriorly in lateral view (Figure

686). The second segment bears two setae, one

barbed, the other naked, and terminates in a moderately long lash with a row of spinules. The

maxilliped (Figure 69c) is 3-segmented. The first

segment is unarmed. The second segment bears

two minute setae. The third segment bears two

small setae and terminates in a finely barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 69d) protrudes ventrally. A line connects the bases of the maxillipeds.

Legs 1-4 (Figure 69e-h) have 3-segmented rami

except for the endopod of leg 4 which is 2-segmented. The armature of these legs is as follows:

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; IIU.4

enp 0-1; 0-1; 13

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,5

enp 0-1; 0.2; 1,11,3

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,13

enp 0-1; 0-2; 1,11,2

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; IIJJ5

enp 0-1; I

The coxa of legs 1-3 bears a long plumose inner

seta, but in leg 4 this seta is minute (6 n) and

naked. The inner margin of the basis is haired in

legs 1-3 but smooth in leg 4. The spines on all

four legs have smooth rather than barbed lamellae.

The exopod of leg 4 is 112 n long. The first segment of the endopod is 22 x 16.5 n, with its distal

inner plumose seta 30 n; the second segment, rather

almond-shaped, is 35 x 16.5 n (the length not including the small spiniform process) and its terminal spine 33 n. Both segments of the endopod have

outer marginal hairs.

Leg 5 (Figure 70a) has a small unornamented

segment 11 n long, not clearly set off from the body

and armed with two setae.

Leg 6 is represented by the two setae on the area

of attachment of each egg sac (Figure 68d).

The color in life in transmitted light is opaque,

the eye red.

MALE.—The body (Figure 70&) resembles in its

general form that of the female. The length (without the ramal setae) is 1.19 mm (1.17-1.22 mm)

and the greatest width 0.43 mm (0.42-0.44 mm),

based on ten specimens in lactic acid. The ratio

of the length to the width of the prosome is 1.39:1.

The ratio of the length of the prosome to that of

the urosome is 1.04:1. The segmentation and the

development of the epimera resemble the female.

The segment of leg 5 (Figure 70c) is 44 x 195 n.

There is no ventral intersegmental sderite. The

genital segment is 230 x 234 n, about as long as

wide. The four postgenital segments are 65 x 99 n,

56 x 91 n, 42 x 82 n, and 60 x 83 n from anterior

to posterior.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 69.—Andrianellus exsertidens, new species. Female: a, first maxilla, posterior (F) ; b,

second maxilla, anterior (c); c, maxilliped, inner (c); d, area between maxillipeds and first

pair of legs, ventral (E) ; e, leg 1 and intercoxal plate, anterior (D) ; /, leg 2, anterior (D) ; g,

endopod of leg 3, anterior (o); h, leg 4 and intercoxal plate, anterior (D) . Scale: c, r, 0.05 mm;

D, E, 0.1 mm.
NUMBER 127 125

FIGURE 70.—Andrianellus exsertidens, new species. Female: a, leg 5, dorsal (F) . Male: b, dorsal

(A) ; c, urosome, dorsal (B) ; d, maxilliped, inner (E) . Scale: A, 0.5 mm; B, 02 mm; E, 0.1 mm;

F, 0.05 mm.

The caudal ramus is similar to that of the female,

but a little shorter, 107 x 30 n, or 3.57 times longer

than wide.

The body surface has hairs and retractile points

as in the female.

The rostrum resembles that of the female.

The first antenna is like that of the female, but

there are three additional, long aesthetes so that

the formula is 4, 13 -+- 2 aesthetes, 6, 3 -j- 1 aesthete, 4 -f- 1 aesthete, 2 -f- 1 aesthete, and 7

+ 1 aesthete.

The second antenna is similar to that of the female except that there is an additional row of small

spinules along the inner margin of the second segment proximal to the seta.

The labrum, mandible, paragnath (?), first

maxilla, and second maxilla are like those in the

female. The maxilliped (Figure 70d) is slender

and 4-segmented (assuming that the proximal part

of the claw represents a fourth segment). The first

segment is unarmed. The second segment bears a

seta with a bulbous base, a recurved seta, and a

row of spinules. The small third segment is unarmed. The slender, slightly recurved claw is 185 Malong its axis including the terminal lamella, is

divided about midway, and bears proximally two

very unequal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-5 are like those of the female.

Leg 6 is a posteroventral flap on the genital

segment bearing two naked setae, 22 n and 30 n.

The spermatophore was not observed.

The color in life resembles that of the female.

ETYMOLOGY.—The specific name exsertidens is

derived from the Latin words exsertus (protruded)

and dens (tooth or tusk), alluding to the extraordinary toothlike process on the first segment of

the second maxilla.
126

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Anisomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, in the

female with the formula 4, IS, 6, 3, 4 + 1 aesthete,

2 + 1 aesthete, and 7+ 1 aesthete; in the male

with 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete, 4

-f 1 aesthete, 2 + 1 aesthete, and 7+ 1 aesthete.

Second antenna 4-segmented, with the armature 1,

1, 3, I + several small elements, there being a

single terminal claw.

Labrum with the two lobes separated by a deep

cleft. Mandible with the basal area distal to the

indentation having on its convex side a scalelike

area with spinules followed by a serrated fringe

and on its concave side a row of spinules; lash long.

Paragnath a small hairy lobe. First maxilla with

three elements. Second maxilla of the usual lichomolgid form. Maxilliped of the female 3-segmented with a pointed tip; in the male 4-segmented

(assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Third segment

of the exopod of legs 3 and 4 either 11,1,5 or

111,1,5. Leg 4 endopod with the formula 0-1; 1,1,

the seta on the second segment being naked. Armature similar in both sexes except for leg 1 endopod in the male where the last segment is 1,1,4

instead of 1,5 as in the female. Leg 5 with a free

segment bearing two setae. Leg 6 in the female

represented by the two setae and the process near

the area of attachment of each egg sac; in the male

by a ventral posterolateral flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with alcyonaceans.

TVPE-SPECIES.—Anisomolgus protentus (Humes

and Frost).

ETYMOLOGY.—The name is a combination of the

Greek cmoo; (unequal), referring to the spine and

the seta on the second segment of leg 4 endopod,

and POXYOC. Gender masculine.

REMARKS.—The male of A. spinipes is unknown.

Key to Species of the Genus Anisomolgus

FEMALES

1. Maxilliped of female with outer side of third segment swollen and membranous

A. incisus

Maxilliped of female with outer side of third segment not swollen and membranous 2

2. Third segment of leg 3 exopod with 11,13; free segment of leg 5 unornamented A. insolent

Third segment of leg 3 exopod with 111,13; free segment of leg 5 ornamented with

spinules 3

3. Free segment of leg 5 relatively short (53 p) and straight; genital segment expanded

laterally near the middle; caudal ramus about 2:1 A. protentus

Free segment of leg 5 long and slightly arcuate, about two-thirds as long as genital

segment; caudal ramus wider than long A. spMpes

KNOWN MALES

1. Inner side of second segment of second antenna with a notched lamella A. incisus

Inner side of second segment of second antenna with a striated membrane or spinules 2

2. This segment with a long striated membrane; third segment of leg 3 exopod with 11,13

A. insolens

This segment with rows of spinules; third segment of leg 3 exopod with 111,13 . A. protentus

Anisomolgus protentus (Humes and Frost, 1964) NEW HOST.—The alcyonacean Sarcophyton glauFIGURE 71 cum (Quoy and Gaimard), collected by AGH: 10

??, 6 o* c?» and 11 copepodids from three coloLichomolgus protentus Humes and Frost 1964 DD 148-150 • • * * » T -m. •»«• J —-.

«M 9ft*_o«« r«. k , „

 p r 1*°^1W' rues, in 4 m, Antsamantsara, Nosy B6, Madagascar,

figs. 203-236 [from the alcyonacean Sarcophyton sp., n T ,«- - «* « «/. J. «» J* J

Ankify. on the mainland of Madaga«ar. near Nosy £ ;

 9 J u n e 1967 ' 24 $ $ , 26 o" d". and 27 copepodids

the host later being described as Sarcophyton gtobosum fro m single colony, in 17 m, near black buoy north

Tixier-Durivauit. 1966]_Bouligand, 1966, p. 269. of Pte. Ambarionaomby, Nosy Komba, near Nosy
NUMBER 127 127

a

FIGURE 11.—Anisomolgus protentus (Humes and Frost). Female: a, dorsal; ft, mandible; c, leg

4 and intercoxal plate. Male: d, dorsal. (From Humes and Frost, 1964, figs. 203, 214, 224, 226.)

Length of female 1.67 mm, of male 1.14 mm.

B£; 14 9 9« 3 o*cf, and 13 copepodids from two

colonies, in 1 m, Ankify, near Nosy Be, 23 August

1967.

Anisomolgus incisus (Humes and Ho, 1968)

Lichomolgus incisus Humes and Ho, 1968c, pp. 680-685,

figs. 148-167 [from the alcyonacean Sarcophyton ehrenbergi Marenzeller, Nosy Be, Madagascar].

NEW RECORD.—21 9 9, 16 o*o". and 14 copepodids from single colony of Sarcophyton ehrenbergi,

in 4 m, Antsamantsara, Nosy Be\ 9 June 1967.

Anisomolgus insolens (Humes and Ho, 1968)

Lichomolgus insolens Humes and Ho, 1968c, pp. 668-674,

figs. 107-127 [from the alcyonacean Lobophytum crassum

Marenzeller, Nosy Be, Madagascar].

Anisomolgus spinipes (Sewell, 1949)

Macrochiron (Macrochiron) spinipes Sewell, 1949, pp. 106-

108, fig. 27a-i [in weed washings, Nicobar Islands].

Macrochiron spinipes.—Stock, 1957, p. 382.

Lichomolgus spinipes.-Stock, 1957, p. 382.

REMARKS.—The male is unknown.

Genus Ascetomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cydopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum broadly

rounded posteroventrally. First antenna 7-segmented, in the female with the formula 4, 13, 6,

3, 4 + 1 aesthete, 2+ 1 aesthete, and 7 -f 1 aesthete. Second antenna 4-segmented with the armature 1, 1, 3, II -(- 5 setae, there being two terminal

claws.

Labrum with the two lobes separated by a deep

cleft. Mandible with the basal area distal to the

indentation having on its convex side a large

hyaline area followed by a serrated fringe and on

its concave side a row of spinules; lash very short.

Paragnath a small hairy lobe. First maxilla with
128

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

four elements. Second maxilla with its general

structure as in other lichomolgids, but the lash

of peculiar form and set off from the segment by

a break in the sclerotization. Maxilliped of the

female 3-segmented, with a pointed tip; in the

male 4-segmented (assuming the proximal part of

the claw to represent a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Leg 4 exopod

with 11,1,5. Leg 4 endopod with 0-1 ;II, the seta

on the first segment being naked. Armature similar

in both sexes except for leg 1 endopod in the male

where the formula of the third segment is 1,1,4

instead of 1,5 as in the female. Leg 5 with a free

segment bearing two terminal setae. Leg 6 in the

female represented by the two setae and spiniform

process near the area of attachment of each egg

sac; in the male by a posteroventral flap on the

genital segment bearing two setae.

Other features as in the species described below.

Associated with octocorals.

TYPE-SPECIES.—Ascetomolgus plicatus, new species.

ETYMOLOGY.—The name is formed from the

Greek words aoKntog (curiously made), alluding to

the form of the lash on the second maxilla, and

noXyog. Gender masculine.

REMARKS.—The genus Ascetomolgus resembles

the genera Colobomo!gus and Contomolgus in

having a large protruding hyaline area (instead

of a scale with spinules) on the convex edge of the

base of the mandible and in having the formula

11,1,5 (instead of 111,1,5) on the third segment

of the exopod of leg 4. In addition, it is unlike

Colobomolgus in having two claws (instead of one

claw) on the last segment of the second antenna.

Ascetomolgus plicatus, new species

FIGURES 72-74

TYPE MATERIAL.—6 99 , 6 0*0" bom three

colonies of the alcyonacean Studeriotes semperi

(Studer), in 17 m, on sandy bottom in pass between Nosy Komba and Nosy B£, northwestern

Madagascar, 23 August 1967, collected by AGH.

Holotype $, allotype, and 6 paratypes (3 $ 9, 3

cfcf) deposited in USNM and the remaining

paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED.—6 99 , 4

from single colony of Studeriotes semperi, in the

same locality as type material, 25 August 1967.

FEMALE.—The body (Figure 72a) has a moderately slender prosome. The length (excluding the

ramal setae) is 0.85 mm (0.78-0.90 mm) and the

greatest width is 0.42 mm (0.40-0.45 mm), based on

five specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.38:1. The

ratio of the length of the prosome to that of the

urosome is 2.21:1. The segment of leg 1 is distinctly separated dorsally from the head by a transverse furrow. The epimeral areas of the segments

of leg 1-4 are more or less rounded.

The segment of leg 5 (Figure 726) is 52 x 128 1*.

There is a short ventral intersegmental sclerite

between that segment and the genital segment. The

genital segment is 122 x 135 n in greatest dimensions, a little wider than long, in dorsal view distinctly divided transversely into two parts, the

smaller posterior part being 113 M in width; in ventral view the segment is entire, without a transverse division. The areas of attachment of the egg

sacs are located dorsolaterally near the middle of

the anterior part of the genital segment. Each area

(Figure 72r) bears two small naked setae, 8 1* and

9 \i, with a small spiniform process between them.

The three postgenital segments are 31 x 74 1*,

19 x 57 n, and 23 x 61 n from anterior to posterior.

The anal segment has on its posteroventral margin

on both sides a row of very small spinules.

The caudal ramus (Figure 72d) is quadrate,

24 x 24 i*. The outer lateral seta is 96 (*, the dorsal

seta 30 ft, and the outermost terminal seta 140 n,

all three of these setae being naked. The innermost terminal seta is 208 1* and usually naked,

though in one female there were a few inner hairs

proximally. The two long median terminal setae

are 340 i* (outer) and 450 »i (inner), both with

prominent lateral spinules along their midregions

and both inserted between dorsal (unornamented)

and ventral (with a marginal row of minute

spinules) flaps.

The surface of the body is ornamented with a

few small hairs (sensilla) as in Figure 72a,b.

The egg sac (Figure 72a) is moderately elongated,

410 x 198 ix, reaches far beyond the caudal rami,

and contains many eggs rather loosely bound together, each egg about 49 n in diameter.
NUMBER 127 129

FIGURE 12,.—Ascetomolgus plicatus, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (c); e, rostrum, ventral

(B) ; /, first antenna, ventral (B) ; g, second antenna, anterior (D) ; h, labrum, ventral (E) ; i,

mandible, posterior (E) ; /, paragnath, ventral (c); k, first maxilla, posterior (E) . Scale: A, 05

mm; B, D, 0.1 mm; c, E, 0.05 mm.
ISO SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The rostrum (Figure 72<?) has a well-defined and

broadly rounded posteroventral margin.

The first antenna (Figure 72/) is 305 n long.

The lengths of the seven segments (measured

along their posterior nonsetiferous margins) are

as follows: 18 (45 n along the anterior margin),

80, 23, 47, 44, 35, and 29 n respectively. The formula

for the armature is 4, 13, 6, 3, 4 + 1 aesthete,

2 -f 1 aesthete, and 7 -f 1 aesthete. All the setae

are naked.

The second antenna (Figure 72g) is 4-segmented.

The first and second segments bear an inner seta.

The third segment has three inner setae, one of

them bent. The fourth segment, 65 n along its

outer edge, 44 n along its inner edge, and 17 n

wide, bears two unequal terminal claws, 45 n and

35 »i, along their axes and five hyaline setae. All the

setae are naked.

The labrum (Figure 72h) has two rather narrow

posteroventral lobes.

The mandible (Figure 72s) has on the concave

side of its basal region a moderately deep indentation followed by a row of spinules; on the

convex margin there is a large, protruding, very

hyaline area followed by a serrated fringe. The

lash is very short and finely barbed. The paragnath

(Figure 72;) is a small hairy lobe. The first maxilla

(Figure 72A) has four elements. The second maxilla

(Figure 73a) has a large unornamented first segment. The slender second segment bears a minute

outer (ventral) setule, a surficial posterior seta

finely barbed along one edge, and an inner (dorsal)

seta similarly barbed along one edge; the segment

terminates in a long lash of peculiar form delimited

from the segment by a break in the sclerotization.

The proximal spinose margin of the lash is protruded ventrally, thus giving the lash the appearance of being folded. The maxilliped (Figure

736) is 3-segmented. The first segment is unarmed.

The second segment bears two very unequal setae,

the longer one barbed. The third segment, bearing

a small naked seta and a bilaterally barbed spine,

terminates in a bilaterally barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 73c) is not protuberant.

A sclerotized line connects the bases of the maxillipeds.

Legs 1-4 (Figures 7$d-g) have trimerous rami

except for the endopod of leg 4 which is 2-segmented. The armature is like that of species of

Metaxymolgus—for example, Af. cuspis (Humes).

The inner seta on the coxa of legs 1-3 is long and

plumose, but in leg 4 it is minute (7 n) and naked.

The inner margin of the basis is haired in legs

1-3 but naked in leg 4. The exopod of leg 4 is

138 n long, its last segment having the formula

11,1,5. The first segment of the endopod is 28 x 24

n (not including the spiniform processes), with its

inner distal seta 21 n and naked. The second segment is 61 n long, 22 ii in greatest width, and

15.5 n in least width (including the spiniform

processes), and its two terminal fringed spines are

26 ix (outer) and 48 n (inner). Both segments have

outer hairs and, in addition, the second segment

bears minute terminal spinules.

Leg 5 (Figure 73/<) has a free segment, 80 x 33 n

(greatest dimensions including the ventral terminal

pointed process), broadest proximally and tapering

distally. Its outer surface bears a few very small

spinules. The two terminal elements consist of a

slender naked seta, 67 n, and a stouter somewhat

spiniform seta, 72 n, with a unilateral fringe. The

seta on the body near the free segment is 55 i*

and naked; near its insertion there are a few minute spinules.

Leg 6 is probably represented by the two setae

and the spiniform process near the attachment of

each egg sac (Figure 72c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.

MALE.—The body (Figure 73s) resembles that

of the female in general form. The length (not

including the setae on the caudal rami) is 0.72 mm

(0.68-0.74 mm) and the greatest width 0.31 mm

(0.29-0.32 mm), based on six specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.45:1. The ratio of the length of the

prosome to that of the urosome is 1.60:1.

The segment of leg 5 (Figure 74a) is 42 x 81 it.

There is no ventral intersegmental sclerite between

that segment and the genital segment The genital

segment is 159 x 155 n. The four postgenital segments are 26 x 49 n, 26 x 47 n, 15.5 x 42 n, and

18 x 47 n from anterior to posterior.

The caudal ramus is like that of the female, but

smaller, 21 x 21 n. The proximal inner hairs on

the innermost terminal seta are present more often

than in the female.
NUMBER 127 131

FIGURE 73.—Ascetomolgus plicatus, new species. Female: a, second maxilla, posterior (E) , b,

maxilliped, anterior (E) , c, area between maxillipeds and first pair of legs, ventral (B) ; d, leg 1

and intercoxal plate, anterior (B) ; e, leg 2 and intercoxal plate, anterior (B) ; /, third segment

of endopod of leg 3, anterior (B) ; g, leg 4 and intercoxal plate, anterior (B) ; h, leg 5, dorsal

(D) . Male: i, dorsal (A) . Scale: A, 0.5 mm; B, D, 0.1 mm; E, 0.05 mm.
132 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The surface of the body has a few small hairs

(sensilla) as in the female.

The rostrum is like that of the female.

The first antenna resembles that of the female

but has three additional aesthetes so that the

formula is 4, 13 -f- 2 aesthetes, 6,3 -f 1 aesthete,

4 -f 1 aesthete, 2 + 1 aesthete, and 7 -f 1 aesthete.

The second antenna also is similar to that of the

female but it shows sexual dimorphism in having

very small spinules on the inner surfaces of the

first, second, and fourth segments.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those in the female.

The maxilliped (Figure 746) is 4-segmented (assuming that the proximal part of the claw represents a fourth segment). The first segment is

unornamented. The second segment bears two

naked setae and a row of spines. The small third

segment is unornamented. The claw is 139 M. along

its axis (including the prominent terminal lamella)

and shows a weak division midway. Its concave

margin has a few small serrations proximal to the

division. The claw bears two very unequal proximal setae, the longer one finely barbed.

The ventral area between the maxillipeds and

the first pair of legs resembles that of the female.

Legs 1-4 are segmented as in the female, with

the spine and setal formula as in that sex except

for the last segment of the endopod of leg 1

(Figure 74r) which has the formula 1,1,4 (the

outer spine being 21 n, the inner 20 n) instead of

1,5. Sexual dimorphism is expressed also in the

last segment of the endopod of leg 2 (Figure lid)

where the three spines, from outer to inner, are

13 n, 7 n, and 17 n (instead of 20 n, 16.5 n, and

25 n in the female). Legs 3 and 4 are like those of

the female.

I^g 5 (Figure 74<r) has an elongated free segment, 38.5 x 8.5 n, not expanded proximally and

bearing a few outer small spinules. It carries

terminally a long naked seta, 46 M, and a short

spiniform element, 21 n, with a serrated fringe along

one edge. The adjacent naked seta on the body is

40 n.

FICURE 74.—Ascetomolgus plicatus, new species. Male: a, urosome, dorsal (F); b, maxilliped, inner

(E) ; c, endopod of leg 1, anterior (E) ; d, third segment of endopod of leg 2, anterior (E) ; e,

leg 5, dorsal (c); /, leg 6, ventral (E) . Scale: c, E, 0.05 mm; F, 0.2 mm.
NUMBER 127 133

Leg 6 (Figure 74/) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 41 u and 44 v, and a small spiniform process.

The spermatophore was not observed.

The color in life is like that of the female.

ETYMOLOGY.—The specific name plicatus (Latin,

folded) refers to the folded appearance of the lash

of the second maxilla.

Ascidioxynus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum triangular

or pointed. First antenna 7-segmented, in the female with the armature 4, 13, 6, 3, 4 -|- 1 aesthete,

2 -f- 1 aesthete, and 7 -f- 1 aesthete; in the male

1, 13 -f 2 aesthetes, f>, 3 -f 1 aesthete, 4 -j- 1 aesthete, 2 -j- 1 aesthete, and 7 -f 1 aesthete. Second

antenna 4-segmented, with the formula 1, 1, 3, and

2 terminal claws -f 5 setae.

Labrum with two lobes separated by a median

indentation. Mandible of simple form, with a

slender base merging into a long bipectinate blade

and lash. Paragnath a small lobe. First maxilla

with three elements. Second maxilla of the usual

lichomolgid type. Maxilliped in the female 3-

segmented, with very small and indistinct third

segment; in the male 4-segmented (assuming that

the proximal part of the claw represents a fourth

segment).

Legs 1-4 with 3-segmented rami, except leg 4

endopod which is 2-segmented. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

the formula 0—1;II,1. Leg 1 endopod with the same

formula in both sexes. Leg 5 with a free segment

bearing two terminal elements. Leg 6 represented

by two setae near the genital openings.

Other features as in the species described below.

Associated with ascidians.

TYPE-SPECIES.—Ascidioxynus floridanus, new species.

ETYMOLOGY.—The generic name is a combination

of agxiftiov, alluding to acidians, and twos, companion or partner. Gender masculine.

REMARKS.—Ascidioxynus resembles species of

Lichomolgus Thorell, from ascidians, in the simple

form of the mandible. The genus differs notably

from such species, however, in the armature of the

endopod of leg 4 (0-1,11 in Lichomolgus).

The formula for the endopod of leg 4 in Ascidioxynus (0—I;II,1) suggests a relationship with

Stellicola Kossmann and Synstellicola Humes and

Stock. However, in those genera (mostly associated

with echinoderms) the second antenna is 3-segmented and the mandible has a more complex

form.

Key to Species of the Genus Ascidioxynus

FEMALES

1. Caudal ramus a little wider than long (30 x 38 /£) A. jamaicensis

Caudal ramus longer than wide 2

2. Genital segment elongated in dorsal view, 195 x 138 ft, broadest near the middle; free

segment of leg 5 with an inner proximal expansion without a lamella A. bermudensis

Genital segment less elongated in dorsal view, 160 x 140 /j,, broadest behind the midregion;

free segment of leg 5 with an inner proximal expansion bearing a hyaline lamella with

a seriated margin A. floridanus

MALES

1. Leg 1 showing slight sexual dimorphism in the stronger development of the obtusely

tipped inner coxal seta; both setae on second segment of maxilliped strongly bent

A. jamaicensis

Leg 1 not showing sexual dimorphism, with inner coxal seta plumose; setae on second

segment of maxilliped not strongly bent 2

2. Second segment of maxilliped with one modified seta (Figure 76ft) and one straight

seta; genital segment with sides subparallel in dorsal view A. floridanus

Second segment of maxilliped with two straight setae; genital segment with slightly

rounded margins in dorsal view A. bermudensis
134

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ascidioxynus ftoridanus, new species

FIGURES 75, 76

TYPE MATERIAL.—3 ? $ , 3 o" o" fro**1 &*

pharynx of two large solitary ascidians (unidentified), dredged in 13-14 fathoms, 290°-340° northwest of wreck buoy on chart 1113, 15-25 miles from

Key West, Florida, in the Gulf of Mexico, August

1951. Holotype $, allotype, and 3 paratypes (2

? ? , 1 cf) deposited in USNM and the remaining

paratypic male (dissected) in the collection of

AGH.

OTHER MATERIAL EXAMINED.—1 9,10 " from a

large unidentified solitary ascidian, ± 3.5 miles

SW of Longboat Pass, Sarasota Bay, on the Gulf

coast of Florida, 9 February 1951, collected by J. B.

Knight; and 2 $ $, 1 d1 from single ascidian, in

5-6 fathoms, ± 3.5 miles SW of Longboat Pass,

Sarasota Bay, Florida, 24 March 1951, collected

by J. B. Knight. Both collections in USNM

(190466).

FEMALE.—The body (Figure 75a) is moderately

slender, and in lateral view the prosome has approximately the same relative thickness as in

Henicoxiphium redactum Illg and Humes (1971).

The length (excluding the ramal setae) is 1.13 mm

(1.01-1.22 mm) and the greatest width 0.43 mm

(0.35-0.47 mm), based on three specimens in lactic

acid. The segment of leg 1 is separated from the

cephalosome by a dorsal transverse furrow. The

epimeral areas of the pedigerous segments are as

indicated in the figure. The ratio of the length

to the width of the prosome is 1.55:1. The ratio of

the length of the prosome to that of the urosome

is 1.75:1.

The segment of leg 5 (Figure 756) is 78 x 135 it.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 169 x 140 n, in dorsal view is broadest

just behind the middle and gently contracted

posteriorly, and bears a posteroventral row of

spines. The areas of attachment of the egg sacs are

situated dorsolaterally in front of the widest part

of the segment. Each area (Figure 75c) bears two

naked setae, about 17 n, and a minute spiniform

process. The three postgenital segments are 83 x

60 H, 73 x 48 \i, and 70 x 55 (i from anterior to

posterior. The posteroventral borders of the first

two postgenital segments bear a row of spines and

the posteroventral border of the anal segment

bears on each side a row of minute spinules.

The caudal ramus (Figure 75d) is moderately

short, 49 x 34 n in greatest dimensions, or 1.44

times longer than wide. The outer lateral seta is

70 n, the dorsal seta 75 n, the outermost terminal

seta 130 n, the innermost terminal seta 143 n, and

the two long median terminal setae 200 M (outer)

and 360 M (inner), both inserted dorsally to a ventral

flap which bears a marginal row of small spinules.

All six setae are naked, but the two long median

terminal setae have a lamella along one edge (on

the inner edge of the outer seta and on the outer

edge of the inner seta).

The surface of the body bears a few hairs (sensilla) as indicated in Figure 75a.

The egg sac is unknown.

The rostrum (Figure 75r) is a triangular, bluntly

pointed beak.

The first antenna (Figure 75/) is 7-scgmented

and 289 n long. The lengths of the segments

(measured along their posterior nonsetiferous margins) are 23 (58 n along the anterior margin), 69,

23, 50, 43, 28, and 18 n respectively. The formula

for the armature is 4, 13, 6, 3, 4 -f 1 aesthete, 2 -f- 1

aesthete, and 7 -f 1 aesthete. All the setae are

naked.

The second antenna (Figure 75g) is 4-segmented,

with the separation between the third and fourth

segments strong on the anterior (inner) surface

but weak on the posterior (outer) surface. The first

and second segments bear a small inner seta. The

third segment bears three setae, one much shorter

than the other two. The fourth segment, 78 n

along its outer edge, 50 n along its inner edge, and

24 ix wide, bears terminally two claws, 35 i* along

their axes, and five slender setae. All the setae are

naked.

The labrum (Figure 75/i) has a short nipplelike protusion on the margin of both posteroventral

lobes.

The mandible (Figure 75i) is slender, its concave

side without a pronounced indentation or a transverse row of spinules, its convex side smooth and

lacking a serrated fringe. The lash merges gradually

with the base and bears a row of spinules on both

sides. Paragnaths could not be identified with certainty but probably are represented by two small

lobes dorsal to the labrum (Figure 75h). The first

maxilla (Figure 75;) has three elements. The second

maxilla (Figure 76a) is 2-segmented, with the
e

FIGURE 75.—Ascidioxynus floridanus, new species. Female: a, dorsal (A); b, urosome, ventral

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (c); e, rostrum,

ventral (D) ; /, first antenna, with arrows indicating positions of additional aesthetes in male,

ventral (D) ; g, second antenna, posterior (outer) (D) ; h, labrum, with paragnaths (?) indicated

by broken lines, ventral (F) ; i, mandible, posterior (c); ;, first maxilla, anterior (E) . Scale:

A, 02 mm; B, D, E, 0.1 mm; c, 0.05 mm.
136 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

second segment armed with the usual number of

elements seen in lichomolgids; the lash has a row

of long slender spines. The maxilliped (Figure

766) is 3-segmented. The first segment is unarmed.

The second segment bears two naked setae and its

posterior and inner surfaces are finely spinulose.

The minute third segment bears a very small

setule and terminates in a minuscule spiniform

process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 76c) is protuberant as

in Henicoxiphium redactum. A line connects the

bases of the maxillipeds.

Legs 1-4 (Figure 76d-g) have the same segmentation and armature as in Henicoxiphium redactum

except for the endopod of leg 4 which here is 2-

segmented with the formula O-1;II,1.

The inner coxal seta of legs 1-4 is plumose, that

of leg 4 about half as long as in the preceding legs.

The inner margin of the basis bears a row of hairs

in all four legs. The exopod of leg 4 is 164 n long,

the armature of the third segment being 11,1,5. The

endopod of leg 4 (Figure 76g) is about 120 j* long.

The first segment is 33 x 32 n (not including the

spiniform processes), with its plumose inner seta

58 n. The second segment is 88 n long (including

the terminal spiniform processes), its greatest width

34 i* proximal to the outer spinous process, its

least width distally 19 n. The inner plumose seta

is 66 n and the two terminal fringed spines are 38

n (outer) and 65 n (inner). The outer margin of

the first segment has a row of hairlike spinules;

that of the second segment has two rows of such

spinules and a third distal row of very short spinules. There is a row of extremely small spinules

at the insertions of the terminal spines. Although

the outer spinous process and the inner seta suggest

a division of the segment, no line of articulation

or break in the sclerotization was observed.

Leg 5 (Figure 76/i) has a broad unornamented

free segment 69 x 50 n in greatest dimensions (the

length including the terminal spiniform process of

3 n). The inner margin of the segment is expanded

to form a hyaline lamella with a serrated margin.

The segment bears terminally a bilaterally lamellose

spine, 55 n, and a naked seta, 66 n. The adjacent

seta on the body is 57 I* and naked.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 75c).

The color in life is unknown.

MALE.—The body is slender, resembling in general form that of the female. (Each of the few

males studied had the urosome strongly flexed,

making it impossible to prepare an accurate figure

of the entire animal.) The length (estimated, and

not including the setae on the caudal rami) is

0.65 mm (0.63-0.70 mm) and the greatest width

0.21 mm (0.21-0.22 mm), based on three specimens

in lactic acid. The ratio of the length to the width

of the prosome is 2.08:1.

The segment of leg 5 (Figure 76i) is 29 x 69 n.

There is no ventral intersegmental sclerite. The

genital segment is 104 x 94 n. The four postgenital

segments are 31 x 56 n, 31 x 52 n, 23 x 52 n, and 29

x 17 n from anterior to posterior.

The caudal ramus is similar to that of the female,

but smaller, 26 x 22 n, and relatively shorter, about

1.18 times longer than wide.

The body surface has a few hairs as in the female.

The rostrum resembles that of the female.

The first antenna is similar to that of the female

but has three additional aesthetes (at the locations

indicated by arrows in Figure 75/) so that the

formula is 4, 13 -f- 2 aesthetes, fi, 3 -f 1 aesthete,

4 -f 1 aesthete, 2 -f 1 aesthete, and 7 -f 1 aesthete. The second antenna, labrum, mandible,

paragnath (?), first maxilla, and second maxilla are

like those of the female. The maxilliped (Figure

76;*) is 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

The first and third segments are unarmed. The

second segment bears two setae (one modified as

in Figure 76A) and two rows of spines. The slender

claw is 114 n along its axis, with a very small

terminal lamella, with a slight indentation about

midway on its concave edge, and with the usual

two very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are like those of the female, with the

same spinal and setal formula.

Leg 5 (Figure 76/) has a small rectangular unornamented free segment, 21 x 11 i*. The terminal

spine is 28 n and the seta 40 i*.

Leg 6 (Figure 76i) is a posteroventral flap on the

genital segment bearing two slender naked setae,

25 H.

The spermatophore was not observed.

The color in life is unknown.
FIGURE 76.—Ascidioxynus floridanus, new species. Female: a, second maxilla, posterior (c); b,

maxillipcd, postero inner (c); c. area between maxillipeds and first pair of legs, ventral (D) ;

d, leg 1 and intercoxal plate, anterior (D) ; e, leg 2, anterior (D) ; /, endopod of leg 3, anterior

(

D

) "> g> leg 4 and intercoxal plate, anterior (D) ; h, leg 5, dorsal (E) . Male: i, urosome, ventral (D) ; ;', maxillipcd, inner (E) ; k, modified seta on second segment of maxilliped, outer

(F) ; /, leg 5, dorsal (c). Scale: c, 0.05 mm; D, E, 0.1 mm; F, 0.02 mm.
138 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

ETYMOLOGY.—The specific name floridanus refers

to the known geographical range, Florida.

Ascidioxynus bermudensis, new species

FIGURES 77-79

TYPE MATERIAL.—195 9 9» 12 tf cf from several

masses of the ascidian Ecleinascidia turbinata Herdman, in 1 m, on rocks and piling, Long Bird Bridge,

between Ferry Reach and Castle Harbor, Bermuda,

24 July 1962, collected by AGH and RUG. Holotype 9, allotype, and 181 para types (172 9 9, 9

0*0*) deposited in USNM; 10 paratypic 9 9 in

ZMA; and the remaining paratypes in the collection

of AGH.

OTHER MATERIAL EXAMINED (all from Ecteinascidia turbinata).—25 9 $• 3 o* d\ and 5 copepodids, in 3 m, northwest of Long Bird Bridge,

Bermuda, 14 July 1962; and 8 9 9, 10 o* tf, and

16 copepodids, in 9 m, Murray's Anchorage, 3

miles northwest of St. George's Island, Bermuda,

9 August 1962. All collected by AGH and RUG.

FEMALE.—The body (Figure 77a) has a moderately broad prosome. The length (without the

ramal setae) is 1.29 mm (1.12-1.36 mm) and the

greatest width is 0.50 mm (0.46-0.54 mm), based

on ten specimens in lactic acid. The segment of

leg 1 is separated from the cephalosome by a dorsal

transverse furrow. The epimeral areas of the pedigerous segments are similar to those in A. floridanus. The ratio of the length to the width of the

prosome is 1.51:1. The ratio of the length of the

prosome to that of the urosome is 1.89:1.

The segment of leg 5 (Figure lib) is 73 x 148 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 195 x 138 H in greatest dimensions

dorsally, slightly expanded in its anterior half. The

areas of attachment of the egg sacs are situated

dorsolaterally near the middle of the segment.

Each area (Figure 77c) bears two naked setae,

about 25 n, and a small spiniform process. The

three postgenital segments are 73 x 94 n, 65 x 90 n,

and 78 x 96 n from anterior to posterior. The

posteroventral border of the anal segment bears a

row of minute spinules on each side. The posteroventral borders of the genital segment and the first

two postgenital segments are smooth, without

spinules.

The caudal ramus (Figure 77^) is moderately

short, 52 x 41 i> in greatest dimensions or 1.27

times longer than wide. The outer lateral seta is

40 \a with outer hairs. The dorsal seta is 65 n with

a few short hairs. The outermost terminal seta is

110 n with outer hairs and the innermost terminal

seta is 190 n with inner hairs. The two long median

terminal setae are 100 n (outer) and 500 n (inner),

both naked; the inner and longer of these two

setae has a lamella along its outer edge. Both

terminal setae are inserted dorsally to a terminal

ventral flap bearing a marginal row of minute

spinules.

The surface of die Ixxly has very few hairs

(sensilla).

The egg sac (Figure 77*) is elongated, 860 x

230 \i, extends beyond the tips of the longest ramal

setae, and contains numerous eggs about 65 n in

diameter.

The rostrum (Figure 77/) is less pointed than in

A. floridanus.

The first antenna (Figure 77g) is 322 n long. The

lengths of its seven segments (measured along their

posterior nonsetiferous margins) are 24 (53 n along

the anterior margin), 88, S3, 50, 45, 32 and 21 n

respectively. The formula for the armature is as in

A. floridanus. All the setae are naked.

The second antenna (Figure llh) resembles that

of A. floridanus. The fourth segment, 66 n along

its outer edge, 45 n along its inner edge, and 23 n

wide, bears two small claws, the stouter claw 25 m

the more slender claw 21 n, and five setae, the

longest slightly unguiform. A few small spinules

occur along the outer margin of the segment All

the setae are naked.

The labrum (Figure lit) resembles that of A.

floridanus.

The mandible (Figure 77;) and first maxilla

(Figure Ilk) are very similar to those of A. floridanus. The paragnath (Figure Hi) is a small lobe

with a few hairs. The second maxilla (Figure 78a)

resembles that of A. floridanus, but the lash is long

and reflexed, with fewer long spines. The maxilliped (Figure 786) is 3-segmented. The second segment has two naked setae. The third segment is

weakly delimited, bears a slender naked setule, and

terminates in a spiniform process with a few small

barbules.

The ventral area between the maxillipeds and

the first pair of legs (Figure 78c) is slightly pro-
FIGURE 77.—Ascidioxynus bermudensis, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal and slightly lateral (c); d, caudal ramus, dorsal

(D) ; e, egg sac, ventral (A); /, rostrum, ventral (E) ; g, first antenna, with arrows indicating

positions of additional aesthetes in male, dorsal (E) ; h, second antenna, posterior (outer) (E) ;

i, labrum and paragnaths, ventral (F); ;, mandible, posterior (F) ; k, first maxilla, posterior (D) .

Scale: A, 1.0 mm; B, 02 mm; c, D, 0.05 mm; E, r, 0.1 mm,
FIGURE 78.—Ascidioxynus bermudensis, rew species. Female: a, second maxilla, posterior (F) ; b,

maxilliped, postero-inner (F) ; c, area between maxillipeds and first pair of legs, ventral (B) ;

d, leg 1 and intercoxal plate, anterior (E) ; e, leg 2, anterior (E) ; /, endopod of leg 3, anterior

(E) ; g, leg 4 and intercoxal plate, anterior (E) ; h, leg 5, dorsal (D) . Scale: B, 0.2 mm; D, 0.05

mm; E, F, 0.1 mm.
NUMBER 127 141

tuberant. A line connects the bases of the maxillipeds.

Legs 1-4 (Figures 78d-g) have the same segmentation and armature as in A. floridanus. The

exopod of leg 4 is 160 n long, the armature of the

third segment being 11,1,5. The endopod of leg 4

(Figure 78g) is 111 n long, including the terminal

spiniform processes. The first segment is 33 x 29 \i

(without the spiniform processes) and its plumose

inner seta is 55 n. The second segment is 81 n

long (including the terminal spiniform processes),

its greatest width 28 n, its least width 17 n. The

inner plumose seta is 58 n and the two terminal

fringed spines are 31 n (outer) and 73 \* (inner).

The outer margin of both segments bears hairlike

spinules. The second segment has outer and inner

spinous processes suggesting division of the segment, but no line of articulation or break in the

soleroti/ation was seen. A row of very small spinules

occurs near the insertions of the two terminal

spines.

Leg 5 (Figure 78/i) has an unornamented free

segment 47 x 36 n in greatest dimensions, with

its proximal margin expanded inwardly. The two

terminal elements are 68 n (naked) and 86 n (with

very small barbules). The adjacent seta on the

body is 40 n and naked.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 77c).

The color in life in transmitted light is translucid to opaque, the eye red, the egg sacs gray or

tan.

MALE.—The body (Figure 79a) is slender. The

length (not including the setae on the caudal rami)

is 0.75 mm (0.73-0.79 mm) and the greatest width

0.24 mm (0.22-0.26 mm), based on ten specimens in

lactic acid. The ratio of the length to the width

of the prosome is 1.85:1. The ratio of the length

of the prosome to that of the urosome is 1.35:1.

The segment of leg 5 (Figure 79&) is 30 x 67 n.

There is no ventral intersegmental sclerite. The

genital segment is 110 x 94 ft. The four postgenital

FIGURE 79.—Ascidioxynus bermudensis, new species. Male: a, dorsal (c); b, urosome, ventral

(E) ; c, maxilliped, inner (D) ; d, leg 5, dorsal (D) ; e, spermatophores, attached to female,

dorsal (E) . Scale: D, 0.05 mm; E, 0.1 mm; c, 0.5 mm.
142 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

segments are 29 x 43 m 26 x 42 n, 18 x 41 v and 32

x 48 n from anterior to posterior.

The caudal ramus, 30 x 23 n, resembles that of

the female.

The body surface has few hairs as in the female.

The rostrum is like that of the female.

The first antenna is similar to that of the female

but has three additional, long aesthetes (at locations indicated by arrows in Figure 77g), so that

the formula is 4, 13 -f 2 aesthetes, 6 3 -f- 1

aesthete, 4 + 1 aesthete, 2 + 1 aesthete, and

7 -f- 1 aesthete. The second antenna, labrum,

mandible, paragnath, first maxilla, and second

maxilla are like those of the female. The maxilliped (Figure 79c) has the first and third segments

unarmed. The second segment bears two naked

setae and a V-shaped group of small spines. The

claw is 96 n along its axis with a very small terminal

lamella, with a line of division about midway, and

with two very unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that in the female.

Legs 1-4 are similar to those in the female, with

the same spine and setal formula but with very

slight sexual dimorphism in the relative length of

the spine on the third segment of the endopod of

leg 1, where, in the male, the spine is 18 n and the

segment 24 n without spiniform processes (in the

female the spine is 20 n and the segment 36 n).

Leg 5 (Figure 79d) has a free segment, 19 x 9 n,

without a proximal inner expansion.

Leg 6 (Figure 796) is a posteroventral flap on

the genital segment, bearing two naked setae about

18 ii long.

The spermatophore (Figure 79e), attached to

the female, is 86 x 43 n, not including the neck.

ETYMOLOGY.—The specific name bermudensis

refers to the known geographical range, Bermuda.

REMARKS.—This species differs from A. floridanus

in several, easily recognizable features. In the female of A. bermudensis the free segment of leg

5 is much less conspicuously expanded inwardly and

the genital segment is broadest near the middle

instead of beyond the middle. In the male both

setae on the second segment of the maxilliped

are unmodified. Other, more subtle, differences

may be seen in the nature of the setae on the

caudal rami, the shape of the rostrum, the relative

size of the two claws on the second antenna, the

form of the lash on the second maxilla, and the

details of the female maxilliped.

Ascidioxynus jamaicensis (C. B. Wilson, 1921)

Fici)R£S 80-82

Modiolicola jamaicensis C. B. Wilson, 1921, pp. 15, 16, pi. 7,

figs. 58-65 [from black ascidians on mangrove roots, Bogue

Islands, Montego Bay, Jamaica].

NEW MATERIAL EXAMINED.—8 9 9 , 9 d1 o*. from

the branchial sac of Ascidia atra (Savigny), depth

about 2 m, entrance of Fuik Bay, Curacao, 2S

October 1958, collected by JHS.

FEMALE.—The total length is 0.93-1.01 ram

(mean, based on five specimens, 0.99 ram), the

greatest width of the cephalosome 0.40-0.47 mm

(mean 0.45 mm). The first pedigerous segment is

almost completely incorporated in the cephalosome

(Figure 80a). Pedigerous segments 2-4 rapidly and

regularly diminish in width. The urosome is 5-

segmented. The genital segment is swollen in its

anterior two-thirds, narrower and regularly cylindrical in its posterior third. On the ventral surface,

the posterior margin of urosomal segments 2-4

bears coarse cuticular serrulations; urosomal segment 5 (or anal segment) is finely denticulated. The

caudal ramus (Figure 806) is 30 n long and 38 11

wide. The smooth lateral seta is implanted not far

from the distal margin. There are four terminal

setae, all of which are smooth except for a distinct

ciliation of the inner margin of the inner seta.

The dorsal seta, implanted over the base of the

innermost terminal seta, is indistinctly plumose.

The longest seta presents a curious lateral, membranous, keel-like projection at some distance from

the base.

The rostrum (Figure 80c) is a fleshy, rather

nondescript mass, tapering into an obtuse point.

The first antenna (Figure 80d) is fundamentally

7-segmented, though it may appear to be 8-«egmented (segment 3 being partially subdivided into

two segments). The armature of the segments is

as follows: 4, 13, 6, 3, 4 -f- 1 aesthete, 2 + 1

aesthete, and 7 -f- 1 aesthete.

The second antenna (Figure 80^) probably

should be considered as 4-segmented. Segment 1

is armed with one short distal seta; segment 2 has

one short subdistal seta. Segments 3 and 4 are

separated rather weakly; at the junction there is
FIGURE 80.—Ascidioxynus jamaicensis (C. B. Wilson). Female: a, dorsal (B) ; b, posterior part

of urosome and caudal ramus, ventral (c); c, rostrum (D) ; d, first antenna, with the aesthete

added in the male indicated by broken lines (D) ; e, second antenna (D) ; /, mandible (c); g,

tint maxilla (c); h, second maxilla (c); i, maxilliped (c). Scale: B, 02 mm; c, 0.05 mm;

D, 0.1 mm.
144 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

on one surface a membranaceous, curved ridge, on

the other surface there are wrinkles in the chitin.

No doubt, this ridge and these wrinkles are rudiments of an articulation line, but no functional

articulation is present. The basal portion (segment

3) of this apparent fusion complex bears two long

and two short setae; the distal portion (segment 4)

has a terminal armature consisting of two setae,

one daggerlike spine, two long clawlike setae, and

two claws.

The mandible (Figure 80/) has a somewhat

widened blade, tapering abruptly into a long lash.

Both margins of the mandible bear sharp, fine spinules, longest in the proximal portion.

The first maxilla (Figure 80g) bears two long,

annulated, terminal setae and one short medial

setule or spinule.

The second maxilla (Figure 80/i) has long, sharp,

needle-like teeth on the medial margin of its terminal lash. The auxiliary lash also has teeth (shorter

than on the terminal lash) on its medial margin.

The maxilliped (Figure 80«) is 2-segmented; the

robust basal segment is unarmed; the second segment is produced into a finger-shaped, hispid terminal lobe; the medial side bears two spines.

Legs 1-3 (Figures 8la-c) are biramous, each

ramus being 3-segmented. Leg 4 (Figure 8Id) has

a 3-segmented exopod and a 2-segmented endopod.

Intercoxal plates, lateral basipod setae (plumose),

and medial coxopod setae (reduced in leg 4) are

present in all legs. The fourth endopod has a

normal basal segment, but the second segment is

constricted, bearing spinelike protuberances on

either end of the swollen part and a plumose seta

on the medial end. The terminal armature consists of two spines.

The chaetotaxis formula of the rami of legs 1-4

is as follows:

exp

end

exp

end

exp

end

exp

end

1-0;

0-1;

1-0;

0-1;

i-O;

0-1;

1-0;

0-1;

1-1;

0-1;

I-l;

0-2;

I-l;

0-2;

I-l;

II-l

III-I-4

1-5

III-I-5

III-3

III-I-5

III-2

II-I-5

The fifth leg (Figure 81e) has a single free segment with a strong, more or less rounded-triangular

projection at the inner margin near the base. The

segment has a width of 39 n at the level of this

projection. Distally, the segment is nearly rectangular and has a width of 19 n. The segment is

59 n in total length. It is distally armed with an

annulated seta and a denticulated spine.

MALE.—The total length is 0.69-0.79 mm (mean,

based on five specimens, 0.75 mm); the width of the

cephalosome is 0.27-0.37 mm (mean 0.31 mm).

The genital segment (Figure 82b) is slightly wider

than long and bears two feathered setules on each

side posteroventrally (^rudimentary leg 6). There

are four postgenital segments (Figure 82a), denticulated as in the female. The caudal ramus (38 n

wide, 30 n long) is armed and ornamented as in

the female.

The first antenna is like that of the female but

has an additional aesthete (dashed area in Figure

SOd) on segment 5.

The maxilliped (Figure 82c) is prehensile. Segment 1 is naked. Segment 2 has two thorny patches

and one band of longer spinules; its medial margin

bears two abruptly bent spines. The terminal claw

is long, curved, and armed with a minute spinule

and a crenulated spine near its base.

Leg 1 (Figure 82d) shows slight sexual dimorphism in the stronger development of the obtusely

tipped medial coxopod seta.

Leg 5 (Figure 82<?) is without a basal projection

and measures 35 x 11 n.

The other appendages are like those in the female.

REMARKS.—Wilson's Modiolicola jamaicensis was

found in "black ascidians common upon the mangrove roots." The present specimens were recovered from the same host, Ascidia atra (=znigra).

Since Wilson's description deviates in several points

from the new material, it was thought wise to reexamine the type material. Through the kind assistance of Drs. Fenner A. Chace and Thomas E.

Bowman, a female paratype (USNM 53568) was

borrowed and dissected. From this study it became clear that Wilson's specimens (from Montego

Bay, Jamaica) were identical with the present material (from Curasao). It is rather irrelevant to

indicate the points where Wilson's description was

incomplete or erroneous (e.g., his figure of leg 5

of the female is in reality leg 5 of the male; his

figure of leg 3 is in reality leg 2, etc.).

In our opinion, this species should be attributed

to the genus Ascidioxynus, with which it shares the

structure and armature of leg 4, the nature of the
NUMBER 127 145

FIGURE 81.—Ascidioxynus jamaicensis (C. B. Wilson). Female: a, leg 1 and part of intercoxal

plate (E) ; b, leg 2 and part of intercoxal plate (E) ; c, third segment of endopod of leg 3

(E) ; d, leg 4 and intercoxal plate (E) ; e, leg 5 (c). Scale: c, E, 0.05 mm.
146

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 82.—Ascidioxynus jamaicensis (C. B. Wilson). Male: a, dorsal (A) ; b, anterior part of

urosome (F) ; c, maxilliped (c); d, leg 1 and part of intercoxal plate (E) ; e, leg 5 (c). Scale:

A, 02 mm; E, c, 0.05 mm; F, 0.1 mm.

mandible, the pointed rostrum, the very small or

totally absent third segment on the female maxilliped, and the 4-segmented nature of the second

antenna with two claws.

Genus Aspidomolgus Humes, 1969

DIAGNOSIS.—Body modified. Prosome in the female shield-shaped, expanded, and orbicular, with

the tergum of the segment of leg 4 overlapping

the urosome. Prosome in the male similar, but the

tergum of the segment of leg 4 not expanded and

the orbicular outline completed by the large broadened genital segment. Urosome 5-segmented in the

female, 6-segmented in the male. Caudal ramus

with six setae. Rostrum broadly truncated posteroventrally. First antenna 7-segmented in both sexes

with the armature being 4, 13, 6, 3, 4 -J- 1 aesthete,

2 -\- 1 aesthete, and 7+ 1 aesthete. Second antenna 3-segmented (the original third and fourth

segments apparently fused), with the formula 1, 1,

3 _|_ II _j_ 5 small setae, there being two terminal

claws. In the male the seta on the second segment

transformed to a sucker.
NUMBER 127 147

Labrum with a deep median cleft between the

two lobes. Mandible with the concave side of the

base having an indentation followed by a row of

stout spinules and the convex side having a scalelike area with a row of short spinules followed by

a serrated fringe; terminal lash long. Paragnath

a small hairy lobe. First maxilla with three setae.

Second maxilla of the usual lichomolgid type. Maxilliped of the female digitiform, apparently 2-segmented though a partial crease suggests the

separation of second and third segments; tip of

maxilliped bluntly rounded. Maxilliped of the

male 4—segmented (assuming that the fourth segment is represented by the proximal part of the

claw).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 1-segmented or with a slight

indication of division. Armature of the usual lichomolgidiform pattern, except leg 4 endopod which

is II,I. Formula of the male similar except leg 1

endopod where the third segment is 1,1,5 instead of

1,5 as in the female. In both sexes the first segment of leg 1 exopod elongated and broad, with

its outer distal area much produced to form a

rounded lobe. Leg 5 in both sexes with a free

segment bearing two terminal elements (a spine

and a seta). Leg 6 in the female represented by

two minute setae near the area of attachment of

each egg sac; in the male by a ventral flap on the

genital segment bearing two setae.

Other features as in the species below.

Associated with actiniarian coelenterates.

TYPE-SPECIES. — Aspidomolgus stoichactinus

Humes.

Aspidomolgus stoichactinus Humes, 1969

FIGURE 83

Aspidomolgus stoichactinus Humes, 1969a, pp. 226-241, figs.

1-68 [associated with the actiniarian Stoichactis helianthus

(Ellis), apparently living in the gastrovascular cavity;

from Barbados, Puerto Rico, Jamaica, and the Bahamas].

f

FIGURE 83.—Aspidomolgus stoichactinus Humes. Female: a, dorsal; b, mandible; c, maxilliped;

d, leg 1 and intercoxal plate; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes,

1969a, figs. 1, IS, 17, 19, 23, 26.) Length of female 1.84 mm, of male 1.60 mm.
148 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Aster kola Rosoll, 1889

DIAGNOSIS.—Body cyclopiform, the prosome not

unusually broad and the urosome relatively long.

Urosome in the female 5-segmented, in the male

6-segmented. Caudal ramus with six setae. Rostrum broadly truncated or rounded. First antenna

7-segmented, in both sexes with the armature 4,

13, 6, 3, 4 4- 1 aesthete, 2 + 1 aesthete, and

7 -j- 1 aesthete. Second antenna 3-segmented, the

third segment resulting from the fusion of two

original segments; formula 1, 1, 3 -f- three terminal claws -|- several small elements (two claws in

A. clausi).

Labrum deeply incised medially. Mandible with

the basal area distal to the indentation, having on

its convex side a row of teeth or serrations and on

its concave side a row of spinules; lash long. Paragnath a small hairy lobe. First maxilla with four

elements. Second maxilla with the initial few teeth

on the lash enlarged. Maxilliped in the female

3-segmented with a pointed tip; in the male 4-segmented (assuming that the proximal part of the

claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Armature of the

lichomolgidiform pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

0-l;2,l. Basis of leg 4 not laterally elongated. No

sexual dimorphism in the formula of leg 1 endopod.

Leg 5 with a free segment bearing two terminal

elements and having its inner edge swollen, irregular, or with a process. Leg 6 represented by the

two setae near the genital openings.

Other features as in the species below.

Associated with asteroid echinoderms.

TYPE-SPECIES.—Astericola clausi Rosoll.

Aster kola clausi Rosoll, 1889

Astericola clausii Rosoll. 1889. pp. 197-201. pi. 2. figf. 7, 8

[from the asteroid Asteracauthion glaciate O. F. Miiller,

Trieste, Italy|.-Gracn*c. 1900. p. 9 [from the aneroid

Astropecten aurantium, Trieste, Italy].—Clark, 1921, p. 636

[on Marthnsteriai glacialis (O. F. Miiller) ].

IAchoiuolgus nstrrinur. llocquet, 1952, pp. 498-501, figs. I-S

[from the asteroid Astrrina gibbosa (Pennant), Roicoff,

northern France).

Key to Species of the Genus Astericola

FEMALES

1. With two daws on last segment of second antenna A. cloud

With three claws on last segment of second antenna 2

2. Length about 1.0 mm; genital segment a little longer than wide 3

Length 1.3-1.6 mm; genital segment about as long as wide 4

3. Caudal ramus 48 x 36 ft, ratio 1.33:1 A. frequens

Caudal ramus 60 x 28 p, ratio 2.14:1 A. astropectims

4. Genital segment much wider in its anterior half than in its posterior half; free segment

of leg 5 with a prominent, toothlike, distally directed process on inner margin A. dentijer

Genital segment not much wider anteriorly than posteriorly; free segment of leg 5 without

such a process 5

5. Caudal ramus 61 x 38 ft, ratio 1.61:1 A. luidiae

Caudal ramus 96 x 43 u, ratio 253:1 A. tautus

MALES

1. With two claws on last segment of second antenna A. clausi

With three claws on last segment of second antenna 2

2. Free segment of leg 5 about 24-26 x 12-13 /x. 3

Free segment of leg 5 about 36-38 x 17 /t 5

3. Caudal ramus about 4.4:1 A. dentifer

Caudal ramus about 1.3-2.1:1 4

4. Caudal ramus a little longer than wide, ratio about 1.3:1 A. frequent

Caudal ramus more elongated, ratio about 2.1:1 A. astropecUnis

5. Caudal ramus somewhat elongated, ratio about 1.6:1 A. luidiae

Caudal ramus more elongated, ratio about 2.2:1 A. tautus
NUMBER 127 149

Stellicola asterinae.—Stock, 1960c, p. 247 [from Asterina

gibbosa, Banyuls, southern France].—Bocquet, Carton, and

Sroehlich, 1970, pp. 500-508, figs. 1-5 [from Asterina gibbosa, Roscoff, France]. [These authors hold that S. asterinae, though close, is distinct from S. clausi].

Stellicola rfatwi.-Bocquet and Stock, 1962a, pp. 80-90, figs.

1-4 [from Marthasterias glacialis and Asterina gibbosa,

Channel coast of France, and from Asterina gibbosa, Banyuls, Mediterranean coast of France].—Carton, 1964, p. 14

[from Marthasterias glacialis and Asterina gibbosa, region

of Roscoff, northern France].—Barel and Kramers, 1970,

table 5 [from Asterina gibbosa, Roscoff, northern

France].—Bocquet, Carton, and Sroehlich, 1970, pp. 508-

516, fig. 9 [from Marthasterias glacialis, Roscoff, France].

Astericola astropectinis (Humes and Cressey,

1958)

Lichomolgus astropectinis Humes and Cressey, 1958b, pp.

334-336, figs. 41-54 [from the astcriod Astropecten michaelseni Koehlcr, Accra, Ghana; and Freetown, Sierra

Leonc].-Humes and Ho, 1967e, p. 221.—Ummerkutty,

1968, p. 317.

Stellicola astropectinis.—Humes, I969d, p. 6.

Astericola dentifer (Humes, 1969)

Stellicola dentifer Humes, 1969d, pp. 2-6, figs. 1-30 [from

the asteroid Luidia clathrata Gray, Port Royal, Jamaica].

Astericola frequens (Humes and Cressey, 1958)

FIGURE 84

Lichomolgus frequens Humes and Cressey, 1958b, pp. 330-

334, figs. 1-40 [from the asteroid Asterina stellifer (M6-

bius), Dakar, Senegal; Accra, Ghana; and Pointe Noire,

Moyen Congo].

Stellicola frequens.—Humes and Ho, 1967e, p. 221.—Humes,

1969d, p. 6.

Astericola lauttts (Humes and Cressey, 1958)

Lichomolgus lautus Humes and Cressey, 1958b, pp. 339-341,

figs. 81-94 [from the asteroid Astropecten hupferi Koehler,

near Freetown, Sierra Leone].

Stellicola lautus.—Humes and Ho, 1967e, p. 221.—Humes,

1969d, p. 6.

Astericola luidiae (Humes and Cressey, 1958)

Lichomolgus luidiae Humes and Cressey, 1958b, pp. 336-338,

figs. 55-80 [from the asteroids Luidia altemata Say var.

numidica Koehler and Luidia atlantidea Madsen, near

Freetown, Sierra Leone].

Stellicola luidiae.-Humes and Ho, 1967e, p. 221.-Humes,

1969d, p. 6.

FIGURE 84.—Astericola frequens (Humes and Cressey). Female: a, dorsal; b, urosome; c, second

antenna; d, mandible; e, second maxilla; /, leg 4; g, endopod of leg 4; h, leg 5. Male: i, dorsal.

(From Humes and Cressey, 1958b, figs. 1, 3, 10, 12, 14, 21, 22, 24, 33.) Length of female 0.99

mm, of male 0.81 mm.
150 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Colobomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male either 5- or 6-segmented (in C. dentipes the preanal and anal segments fused, but in C. laboutei separate). Caudal

ramus with six setae. Rostrum either with a delicate rounded posteroventral margin or without a

well-defined margin. First antenna 7-segmented in

the female with the formula 4, 13, 6, 3, 4 -f- 1

aesthete, 2 -|- 1 aesthete, and 7 —|— 1 aesthete; in

the male 4, 13 -f- 2 aesthetes, 6, 3 -f 1 aesthete,

4 -f 1 aesthete, 2 + 1 aesthete, and 7 -f 1 aesthete. Second antenna 4-segmented, with the formula 1, 1, 3, I + 6, there being a single terminal

claw.

Labrum with a median cleft separating the two

lobes. Mandible with the basal area beyond the

indentation having on its convex edge a scalelike

area bearing a row of spinules and a serrate fringe

and on its concave side a row of spinules; lash extremely reduced, represented only by a small

pointed process. Paragnath a small hairy lobe. First

maxilla with two or three elements. Second maxilla of the usual lichomolgid type. Maxilliped of

the female 3-segmented, with a pointed tip; in the

male 4-segmented (taking the proximal part of

the claw to represent a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Armature of the

usual lichomolgidiform type. Leg 4 exopod with

the third segment 111,1,5. Leg 4 endopod 0-1 ;II,

the seta on the first segment being feathered. Leg

1 endopod in the male with the third segment 1,1,4

instead of 1,5 as in the female. Leg 5 in both sexes

with a free segment bearing two terminal setae.

Leg 6 in the female represented by two setae and

pointed process near the area of attachment of each

egg sac; in the male by a posteroventral flap on

the genital segment bearing two setae.

Other features as in the species below.

Associated with alcyonaceans.

TYPE-SHF.CIES.—Colobomolgus dentipes (Thompson and A. Scott).

ETYMOLOGY.—The name is a combination of the

Greek words xoXoflos (short-horned), referring to

the very reduced lash on the mandible in the form

of a small process, and noXyos. Gender masculine.

REMARKS.—The male of C. cristatus is unknown.

Colobomolgus dentipes (Thompson and A. Scott,

1903)

l.ichomolgus dentipes Thompson and A. Scott, 1903, p. 281,

pi. 16, figs. 27-30 [in washings of dredged invertebrates,

CeylonJ.-Humes and Ho, 1968c. pp. 637-644, figs. 1-28

[from IIK alcyonaccan Sinularia humesi Verseveldt, No«y

Be. northwestern Madagascar].

I.ichonwlgus (Siellicola) dentipes.—Monod and Dollfus,

1932a. p. 139.

NEW RECORDS.—78 9 9, 52 cf cf from Sinularia

humesi in 18 m, Bane du Touareg, Bay of Ampasindava, near Nosy Be, Madagascar, 11 July 1967,

collected by AGH; 23 $ 9, 19 cf cf from Sinularia

humesi in 13 m, opposite Antsiabe, southern shore

of Nosy Komba, near Nosy B£, 2 September 1967,

collected by AGH.

Colobomolgus cristatus (Humes and Ho, 1968)

Lichomolgus cristatus Humes and Ho, 1968c, pp. 644-650,

figs. 29-50 [from the alcyonacean Sinularia leptoclados

Ehrenberg, Nosy Be, Madagascar].

Key to Species of the Genus Colobomolgus

FEMALES

1. Free segment of leg 5 with a very large toothlike process C. dentipes

Free segment of leg 5 with at most only a slight expansion 2

2. Caudal ramus elongated, ratio 4.5:1 C. cristatus

Caudal ramus short, ratio 1.6:1 C. laboutei

KNOWN MALES

Third and fourth postgenital segments fused, the region of the fourth segment distinctly

broader than the other postgenital segments C. dentipes

Third and fourth postgenital segments not fused, fourth segment not distinctly broader than

the other postgenital segments C. laboutei
NUMBER 127 151

NEW RECORDS (all from Sinularia leptoclados in

Madagascar, collected by AGH).—2 9 9 in 15 m,

Nosy Iranja, southwest of Nosy Be\ 9 August 1967;

1 9 ' n 1 m ' Ankify, on the mainland near Nosy

Be, 11 August 1967; and 21 9 9 in 20 m, west of

Andilana, Nosy Be, 13°18' S, 48°07' E, 24 August

1967.

REMARKS.—The male is unknown.

Colobomolgus laboutei, new species

FIGURES 85. 86

TYPE MATERIAL.—51 $ 9» 20 <$ tf from two

colonies of the alcyonacean Sinularia leptoclados

(Hemprich and Ehrenberg), in 1 m, Ankify, on

the mainland of Madagascar opposite Nosy Komba,

near Nosy Bd, northwestern Madagascar, 23 August 1967, collected by AGH. Holotype ? , allotype, and 64 paratypes (48 9 ? , 16 <? cf) deposited

in USNM, the remaining paratypes (dissected) in

the collection of AGH.

OTHER MATERIAL EXAMINED (all from Sinularia

leptoclados).—5 9 ?» from single colony, in 1 m,

Ankify, 11 August 1967; and 24 o" tf from single

colony, in 20 m, west of Andilana, Nosy B£, 13°18'

S, 48°07' E, 24 August 1967. All collected by

AGH.

FEMALE.—The body (Figure 85a) resembles in

general form that of C. cristatus (Humes and Ho).

The length is 0.83 mm (0.77-0.96 mm) and the

greatest width 0.52 mm (0.50-0.53 mm), based on

ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.21:1. The

ratio of the length of the prosome to that of the

urosome is 2.17:1. The segment of leg 1 is separated from the head dorsally by a transverse furrow. The epimera of the segments of legs 2-4 are

truncated.

The segment of leg 5 is 86 x 161 n. Near the

base of the leg the edge of the segment is protruded laterally, forming a small bulge. Ventrally

between this segment and the genital segment is a

short very weak ventral intersegmental sclerite. The

genital segment, 82 x 134 n in greatest dimensions,

resembles that of C. cristatus. The areas of attachment of the egg sacs are like those of C. cristatus.

The three postgenital segments are 36 x 62 n, 23.5

x 61 n, and 47 x 57 v from anterior to posterior.

The caudal ramus (Figure 85b) is 34 x 22 n,

about 1.6 times longer than wide. The outer lateral seta is 68 \i and naked. The dorsal seta is 44 n

and lightly feathered. The outermost terminal seta

is 70 \i and the innermost terminal seta 153 n, both

with spinules. The two long median terminal setae

are 250 *i (outer) and 410 n (inner), both with spinules and both inserted between dorsal (unornamented) and ventral (with a row of marginal

spinules) flaps. Several small spinules occur near

the insertion of the outer lateral seta and there are

a few small surficial hairs.

The body surface is ornamented with a few hairs

(sensilla) as indicated in Figure 85a. The posteroventral margin of the anal segment is unornamented.

No complete egg sacs were seen. Each egg is

about 47 n in diameter.

The rostrum is like that of C. cristatus.

The first antenna has the same segmentation and

armature as in C. cristatus. The lengths of the

segments (measured along their posterior nonsetiferous margins) are 33 (68 n along the anterior

edge), 135, 29, 53, 24, 17, and 15 n respectively.

The plumose setae have fewer, shorter, and more

weakly developed hairs than in C. cristatus.

The second antenna resembles that of C. cristatus. The fourth segment is 75 n along its outer

edge, 45 n along its inner edge, and 19 n wide.

The terminal claw is 45 n along its axis.

The labrum, mandible (Figure 85c), paragnath,

first maxilla, and second maxilla are like those in

C. cristatus. The maxilliped (Figure 85d) resembles that of C. cristatus but is a little more ornamented.

The ventral area between the maxillipeds and

the first pair of legs is similar to that in C. cristatus.

Legs 1-4 are segmented and armed as in C. cristatus. The spine on the last segment of the endopod of leg 1 (Figure 85e) is straight rather than

recurved as in that species. The exopod of leg 4

is 80 v- long. The first segment of the endopod of

this leg is 22 x 9 n and its distal inner feathered

seta 31 n. The second segment is 45 x 7.5 n (the

length including the terminal spiniform processes

and the width taken at the middle); its two terminal unequal barbed spines being 17 n (outer) and

39 n (inner). As in C. cristatus, there is an outer

row of hairs on bodi segments.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 85.—Colobomolgus laboutei, new species. Female: a, dorsal (A) ; b, caudal ramus, dorsal

(B) ; c, mandible, posterior (B) ; d, maxilliped, posterior (c); e, third segment of endopod of

leg 1, anterior (D) ; /, leg 5, dorsal (c). Male: g, dorsal (E) ; //, urosomc, dorsal (F) . Scale:

A, 0.5 mm; B, 0.02 mm; c, 0.05 mm; t>, r, 0.1 mm; E, 02 mm.
NUMBER 127 153

Leg 5 (Figure 85/) has a moderately elongated

free segment 47 n long, 16 n wide at the slight

proximal inner expansion, and 11 n wide near the

distal end. The outer surface is ornamented with

spinules. The two terminal naked elements are

26 n and 28 n. The adjacent seta on the body is

52 n and lightly feathered.

Leg 6 is probably represented by the two setae

near the area of attachment of each egg sac.

The color in life in transmitted light is opaque,

the eye red.

MALE.—The body (Figure 85g) is a little less

expanded than in the female. The length is 0.58

mm (0.56-0.63 mm) and the greatest width 0.23

mm (0.21-0.26 mm), based on ten specimens in

lactic acid. The ratio of the length to the width

of the prosome is 1.33:1. The ratio of the length

of the prosome to that of the urosome is 1.84:1.

The segment of leg 5 (Figure 85/i) is 24 x 59 n,

without the lateral swelling seen in the female.

There is no ventral intersegmental sclerite. The

genital segment is 122 x 117 I*. The four postgenital segments are 13 x 25 n, 14 x 32 n, 10 x

31 \i, and 18 x 34 n from anterior to posterior.

The caudal ramus is 17 x 14.5 n, relatively

shorter than in the female. The armature is like

that of the female, except that the inner (and

sometimes also the outer) of the two long median

terminal setae is often naked or nearly so.

The rostrum is like that of the female.

The first antenna is segmented and armed as in

the female, but there are two additional aesthetes

on the second segment and one additional aesthete

on the fourth segment (Figure 85g) so that the

formula is 4, 13 -f 2 aesthetes, 6, 3 -f- 1 aesthete,

4 -f 1 aesthete, 2 -f 1 aesthete, and 7 + 1 aesthete.

The second antenna (Figure 86a) is segmented

and armed in the female, but the second segment

has fewer outer spinules and bears a long inner

fringe of short slender spinules. The fourth segment, more slender than in the female, is 53 n

along its outer edge, 35 n along its inner edge, and

10 |i wide, with its claw 28 |i along its axis.

B

FIGURE 86.—Colobomolgus laboutei, new species. Male: a, second antenna, posterior (c); b,

maxilliped, inner (D) ; c, endopod of leg 1, anterior (B) ; d, posterior portion of genital segment, dorsal (F) ; e, spermatophores. attached to female, dorsal (F) . Scale: B, 0.02 mm; c, 0.05

mm; D, F, 0.1 mm.
154 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 86 b) is slender and 4-

segmented (assuming that the proximal part of the

claw represents a fourth segment). The armature

and ornamentation are similar to C. dentipes. The

recurved claw is relatively short, 86 n along its

axis.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female except that the armature for the last segment of the endopod of leg 1 (Figure 86c) is 1,1,4

instead of 1,5 as in the other sex. The outer spine

is recurved, the terminal spine straight, and both

are prominently spinulose along one edge and have

very small spinules along the other edge. Between

the two spines there is a long, pointed, somewhat

sinuous process with very small spines. The endopod of leg 4 is like that of the female.

Leg 5 (Figure 85/?) has an unornamented rectangular free segment, 16.5 x 5.5 n. The terminal

elements are 23 n and 27 n, and the seta on the

body adjacent to the free segment is 33 n and

naked.

Leg 6 is a posteroventral flap on the genital segment, slightly protruded in some specimens (Figure 86d) or not protruded in others (Figure 85/i).

The two naked setae are 14 n and 17 n.

The spermatophore (Figure 86e), attached to the

female in pairs, is 120 x 49 n, not including the

neck. (In the figure one spermatophore appears

shorter than the other because of the slight angle

at which it was held in the specimen drawn.)

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—This species is named for Pierre

Laboute, the diver at the Centre ORSTOM de

Nosy Be who accompanied AGH on many SCUBA

dives during the fieldwork at Nosy B£ and thereby

made possible the collection of hosts otherwise difficult to obtain.

REMARKS.—The new species resembles closely C.

cristatus in the form of the genital segment in the

female, in the numerous feathered setae on the

first antenna, and in the crest of long setae on the

lash of the second maxilla, but it may be distinguished from that species by the shorter caudal

ramus (4.5:1 in C. cristatus) and by the spinules

on leg 5 (this leg being unornamented in C. cristatus). The male of C. cristatus is unknown.

Colobomolgus laboutei differs from C. dentipes

in three readily observable features. In C. laboutei

the first antenna in both sexes has numerous feathered setae, while in C. dentipes there are very few

such setae. In the female of C. laboutei leg 5 lacks

the large toothlike process seen in C. dentipes. In

the male of the new species the outer spine on the

last segment of the endopod of leg 1 is recurved,

while in C. dentipes this spine is straight.

Genus Contomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. Rostrum broadly rounded

posteroventrally. First antenna 7-segmented, the

armature in the female being 4, 13, 6, 3, 4 -f- 1

aesthete, 2 -(- 1 aesthete, and 7 -f- 1 aesthete; in

the male 4, 13 -|- 2 aesthetes, 6, 3 -j- 1 aesthete,

4 -f I aesthete, 2 + 1 aesthete, and 7 -f 1 aesthete. Second antenna 4-segmented, with the armature 1, I, 3, and II -j- 5 hyaline elements, there

being two terminal claws.

Labrum with the two lobes separated by a deep

cleft. Mandible with the basal area distal to the

indentation having on its convex side a scalelike

protrusion with a row of spines followed by a serrated fringe and on its concave side a row of slender spinules followed by a few stouter ones; lash

very short and barbed. Paragnath a small hairy

lobe. First maxilla with four elements. Second

maxilla of the usual lichomolgid type. Maxilliped

in the female 3-segmented, with a pointed tip; in

the male 4-segmented (the proximal part of the

claw probably representing a fourth segment).

Legs 1-4 with 3-segmented raini, except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform type, leg 4 endopod being

0-1 ;II, the seta on the first segment naked. Leg 4

exopod with the third segment having 111,1,5. In

the male, leg 1 endopod showing sexual dimorphism, with 1,1,4 instead of 1,5 as in the female.

Leg 5 in both sexes with a free segment bearing

two terminal setae. Leg 6 in the female represented by the two setae and small process near the

area of attachment of each egg sac; in the male by

a posteroventral flap on the genital segment bearing two setae.
NUMBER 127 155

Associated with alcyonaceans.

TYPE-SPECIES.—Contomolgus lokobeensis, new

species.

ETYMOLOGY.—The name is a combination of the

Greek words KOVTO; (short), referring to the very

short lash on the mandible, and noXyos. Gender

masculine.

Contomolgus lokobeensis, new species

FIGURES 87-90

TYPE MATERIAL.—14 99 , 53 <S <$, and 70

copepodids from single colony of the alycyonacean

Studeriotes sernperi (Studer), in 17 m, with its base

embedded in sand, in pass at Pte. Lokobe, Nosy

B£, northwestern Madagascar, 25 August 1967, collected by AGH. Holotype 9, allotype, and 61 paratypes (11 9 9, 50 cTcf) deposited in USNM, the

remaining paratypes (dissected) and the copepodids in the collection of AGH.

OTHER MATERIAL EXAMINED.—From Studeriotes

semperi: 7 9 9, 13 <? c?. and 92 copepodids from

three colonies, in 17 m, in pass at Pte. Lokobe, 23

August 1967; 5 9 9, 20 rfcf, and 118 copepodids

from single colony, in 18 m, in pass at Pte. Lokobe,

14 July 1967. From the alcyonacean Dendronephthya (Morchellana) stocki Verseveldt : 11 99 ,

7 0* cf, and 3 copepodids from single colony, in

25 m, north of Ankazoberavina, 13°27.6' S, 47°58.2/

E, near Nosy Be, 24 August 1967. All collected by

AGH.

FEMALE.—The body (Figure 87a) has a moderately broad prosome. The length is 1.41 mm (1.39-

1.49 mm) and the greatest width 0.70 mm (0.67-

0.72 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome is 1.42:1. The ratio of the length of the

prosome to that of the urosome is 2.46:1.

The segment of leg 5 (Figure 87b) is 99 x 250 ji.

Between this segment and the genital segment there

is a short, weakly developed ventral intersegmental

sclerite. The genital segment is 220 x 205 n, a

little longer than wide, expanded in the anterior

two-thirds where there is a pair of median, dorsally

raised areas. The areas of attachment of the egg

sac are located dorsolaterally at about the middle

of the segment. Each area (Figure 87c) bears two

small naked spiniform setae 11 n and 22 >*, with a

bluntly pointed process between them. The three

postgenital segments are 62 x 117 n, 47 x 112 \t,

and 43 x 104 n from anterior to posterior.

The caudal ramus (Figure 87d) is quadrate, 39

x 40 n. The outer lateral seta is 225 n and naked.

The dorsal seta is 47 n and lightly feathered. The

outermost terminal seta is 285 v, with spinules

along its proximal outer fourth. The innermost

terminal seta is 400 n, the spinules along its outer

edge a little more sparse than those on its inner

edge. The two long median terminal setae are 616

n (outer) and 670 n (inner), both with spinules

and both inserted between dorsal (unornamented)

and ventral (with a few very minute marginal

spinules) flaps.

The body surface bears hairs (sensilla) as shown

in Figure %la,b. The posteroventral margin of the

anal segment bears a row of extremely small spinules on both sides.

The egg sac (Figure 87a) is oval, 660 x 310 n,

reaches far beyond the caudal rami, and contains

many eggs about 48 n in diameter.

The rostrum (Figure 87e) is well denned and

rounded posteroventrally.

The first antenna (Figure 87/) is 583 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are as follows: 50 (95 n along the anterior margin), 146,

36, 81, 97, 78, and 50 n respectively. The formula

for the armature is 4, 13, 6, 3, 4 -j- 1 aesthete,

2 -\- 1 aesthete, and 7 -f 1 aesthete. All the setae

are naked except for one on segment 5, one on

segment 6, and two on segment 7.

The second antenna (Figure 87g) is 4-segmented.

Each of the first two segments bears a seta. The

third segment has three setae. The last segment,

110 M- along its outer edge, 73 n along its inner

edge, and 31 ft wide, bears two slightly unequal

terminal claws, 75 n and 64 1*, along their axes

and five hyaline elements, the longest one setiform.

All the elements are naked. The inner surface of

the fourth segment bears a row of minute spines.

The labrum (Figure 87/i) has two posteroventral

lobes, their borders at the median indentation bearing a row of small spinules.

The mandible (Figure 88a) has on the concave

side of the base a strong indentation followed by

a row of slender spinules and then a group of

stouter ones. The convex margin of the base bears

a protruding scalelike area with a row of spinules

followed by a serrated hyaline fringe. The lash is
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 87.—Contomolgus lokobeensis, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c. area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum,

ventral (E) ; /, first antenna, dorsal (r) ; g, second antenna, posterior (F); h, labrum, with

paragnaths indicated by broken lines, ventral (c). Scale: A, 0.5 mm; B, E, 0.2 mm; c, F, 0.1 mm;

D, 0.05 mm.
NUMBER 127 157

very short and bilaterally barbed (not a smooth

spike as in Colobomolgus). The paragnath (Figure

%lh) is a small hairy lobe. The first maxilla (Figure 886) bears four elements, the longest with a

few minute barbules. The second maxilla (Figure

88c) has a large unornamented first segment. The

second segment has an outer (ventral) proximal

naked setule, a more distal posterior surficial spiniform seta, and a bilaterally spinulose spine which

is much shorter than the terminal lash (with small

teeth along one edge). The maxilliped (Figure

88rf) is 3-segmented. The first segment is unornamented. The second bears two very unequal

setae, the longer one almost as long as the entire

maxilliped and bearing a row of erect slender

spines. The third segment bears a barbed spine,

a small setule, and a terminal barbed spiniform

process lacking an articulation.

The ventral area between the maxillipeds and

the first pair of legs (Figure 88*) is not protuberant. A fine line connects the bases of the maxillipeds.

Legs 1-4 (Figure 88/-1) have trimerous rami,

except for the endopod of leg 4 which is 2-segmented. The armature is as follows:

m,i,4

13

HM,5

I.H.S

11143

IMS.

niiniiii. 111c iiiiiiiiiuiL 11 .!•> IUIIUW:

P, coxa 0-1 basis 1-0 cxp 1-0; 1-1

enp 0-1; 0-1

P, coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; 0-2

P, coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; 0-2

P4 coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; II

The inner element on the coxa of legs 1-3 is a

large feathered seta, but in leg 4 it is reduced to a

minute, naked element 5 n long. The inner margin

of the basis in legs 1-3 bears a row of hairlike

spinules but in leg 4 this margin is smooth. The

exopod of leg 4 is 234 JA in length. The usual

formula for the last segment is 111,1,5, but one

female showed 11,1,5 on the right leg. The first

segment of the endopod is 57 x 44 ji (not including

the spiniform processes) and its inner distal naked

seta is 54 n. The second segment is 112 n long

(including the terminal spiniform processes), 42 n

in greatest width proximally, and 27 n in least

width distally; the outer terminal spine is 46 n

and the inner spine 106 n, both fringed and with

minutely bifurcated tips. Both segments have an

outer row of hairs.

Leg 5 (Figure 88;) has a broad free segment

226 x 101 n in greatest dimensions, finely spinulose

on its dorso-outer surface. (The free segment in

dorsal view, as in Figure 87b, appears to be much

more slender.) The two terminal naked jointed

setae are 130 n and 270 n. The adjacent seta on

the body is approximately 45 n and lightly feathered.

Leg 6 is probably represented by the two elements near the attachment of each egg sac (Figure

87c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.

MALE.—The body (Figure 89a) is a little more

slender than in the female. The length is 1.15 mm

(1.10-1.22 mm) and the greatest width 0.44 mm

(0.42-0.46 mm), based on ten specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.63:1. The ratio of the length of the

prosome to that of the urosome is 1.81:1.

The segment of leg 5 (Figure 896) is 61 x 130 n.

There is no ventral intersegmental sclerite. The

genital segment is 235 x 239 n. The four postgenital segments are 34 x 78 n, 34 x 77 n, 26 x 74

n, and 29 x 75 n from anterior to posterior.

The caudal ramus is like that of the female, but

smaller, 30 x 29 n.

The surface of the body is ornamented with

hairs as in the female.

The rostrum is similar to that in the female.

The first antenna is segmented and armed as in

the female, but there are two additional aesthetes

on segment 2 and one additional aesthete on segment 4 (Figure 89a) so that the formula is 4, 13

-(- 2 aesthetes, 6, 3 + 1 aesthete, 4 -|- 1 aesthete,

2 + 1 aesthete, and 7+ 1 aesthete. Certain setae

are feathered as in the female.

The second antenna (Figure 89c) is a little more

slender than in the female. The fourth segment

is 102 n along its outer edge, 73 n along its inner

edge, and 24 n wide; its two terminal claws are

60 n and 53 n. The inner surfaces of the first two

segments and the fourth segment bear small spines.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 89d) is slender and 4-

segmented (assuming that the proximal part of the

claw represents a fourth segment). The first segment is unornamented. The second segment bears

two setae and two rows of spines (one row short).
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 8S.—Contomolgus lokobeensis, new species. Female: a, mandible, posterior (c); b, first

maxilla, anterior (c); c, second maxilla, posterior (c); d, maxilliped, postero-inner (F) ; e,

area between maxillipeds and first pair of legs, ventral (E) ; /, leg 1 and intercoxal plate,

anterior (E) ; g, leg 2, anterior (E) ; h, third segment of endopod of leg 3, anterior (E) ; i, leg

4 and intercoxal plate, anterior (E) ; ;, leg 5, dorso-inner (E) . Scale: c, r, 0.1 mm; E, 02 mm.
NUMBER 127 159

g

FIGURE 89.—Contomolgus lokobeensis, new species. Male: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, second antenna, anterior (F) ; d, maxilliped, inner (E) ; e, endopod of leg 1, anterior

(F) ; /, third segment of endopod of leg' 2, anterior (F) ; g, third segment of endopod of leg 3,

anterior (F) ; h, leg 5, dorsal (D) ; i, spermatophores, attached to female, dorsal (B) . Scale:

A, 0.5 mm; B, E, 02 mm; D, 0.05 mm; F, 0.1 mm.
160 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The small third segment is unarmed. The claw

is 273 n along its axis (including the terminal

lamella) and only slightly recurved. It is weakly

divided about midway and is ornamented on the

proximal half with a pointed knob. There are two

very unequal proximal elements, the longer one

with unilateral barbules.

The ventral area between the maxillipeds and

the first pair of legs is like that in the female.

Legs 1-4 are segmented and aimed as in the

female, except for the last segment of the endopod

of leg 1 (Figure 89e), where the formula is 1,1,4

instead of 1,5 as in the female. Slight sexual dimorphism occurs in the last segment of the endopod

of leg 2 (Figure 89/) where the outer spine is 40 ^

and smooth, the terminal outer spine 33 n, and the

terminal inner spine 40 n (in the female these

spines being 36, 33, and 48 n respectively, all

fringed). The last segment of the endopod of leg

3 (Figure 89g) shows only very slight dimorphism

in the lengths of the spines, 45, 43, and 53 >i from

outer to inner (in the female 46, 44, and 60 n).

Leg 4 is like that of the female.

Leg 5 (Figure 89/i) has a moderately slender free

segment 52 x 19 n, spinulose on its dorso-outer surface as in the female. The terminal elements consist of a long naked seta, 74 u, and a short fringed

spine, 25 n. The adjacent seta on the body is 33 n

and naked.

Leg 6 (Figure 896) is a posteroventral flap on

the genital segment bearing two naked setae, 39 n

and 56 n.

The spemiatophore (Figure 89»), attached to the

female in pairs, is oval, 187 x 96 ^ (not including

the neck). Eight females were observed with spermatophores, seven with a pair on both sides, and

one with a pair on the right side only.

The color in life in transmitted light is like

that of the female.

ETYMOLOCY.—The specific name refers to Pte.

Lokobe, the type-locality.

REMARKS.—The distinguishing features of C.

lokobeensis are the very short barbed lash on the

mandible, the two claws on the second antenna,

the distal spine on the second segment of the second maxilla much shorter than the lash, and the

FICURE 90.-Contomolgus lokobeensis, new species. Female: a, urosome, dorsal (A); b, leg 5.

dorsal (B) . Male: c, leg 5, dorsal (c); d, spermatophores, attached to female, dorsal (B) .

Scale: A, 0.3 mm; B, 02 mm; c, 0.05 mm.
NUMBER 127 161

naked seta on the first segment of the endopod of

leg 4.

The specimens from Dendronephthya (Morchellana) stocki differ in certain minor details from

those found on Studeriotes semperi. In the specimens from the former the genital segment of the

female (Figure 90a) is 109 x 211 n and has a

slightly different form; the free segment of leg 5

in the female (Figure 906), 216 x 86 n, has a

slightly different contour (though the contour may

depend upon the precise orientation of the appendage when drawn); the free segment of leg 5

in the male (Figure 90c) is 53 x 13 \i, a little more

slender than in the material from Studeriotes, the

ratio being 4.08:1; the serrated fringe on the terminal spine is less developed; and the spermatophore

(Figure 90d) is more elongated, 208 x 94 n, not

including the neck.

These differences are so minor that we consider

the specimens from Dendronephthya to represent

C. lokobeensis. In other respects, such as body size

and the structure of the appendages, the specimens

from the two hosts are very similar.

Genus Debrwna Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. Rostrum linguiform.

First antenna 7-segmented, in the female with the

formula 4, 13, 6, 3, 4 -|- 1 aesthete, 2+ 1 aesthete,

and 7 -f- 1 aesthete; in the male 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete, 4 + 1 aesthete, 2+ 1

aesthete, and 7 + 1 aesthete. Second antenna 4—

segmented, with the armature 1, 1, 4, and III + 3,

there being three terminal claws.

Labrum with the two lobes separated by an indentation. Mandible with the base bearing a slender bipectinate blade. First maxilla with one

element. Second maxilla of the usual lichomolgid

type. Maxilliped of the female 2-segmented, the

second and third segments fused, with a rounded

tip; in the male 4-segmented (the proximal part

of the claw probably representing the fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform type, but leg 4 endopod

having the formula 0—1 ;I. Leg 4 exopod with the

third segment 11,1,5. No sexual dimorphism in leg

1 endopod. Leg 5 in both sexes with a free segment

bearing two unequal terminal elements. Leg 6

represented in the female by the two setae and

process near the area of attachment of each egg sac

and in the male by a posteroventral flap on the

genital segment bearing two setae.

Other features as in the species described below.

Associated with ascidians.

TYPE-SPECIES.—Debruma clavelinae, new species.

ETYMOLOGY.—The generic name is an anagram

of Bermuda, the type locality of D. clavelinae. Gender masculine.

REMARKS.—Debruma appears to be related to

Gelastomolgus Humes, 1968. Both genera have

four setae on the third segment and three claws on

the fourth segment of the second antenna, a bladelike mandible, a fusion of the second and third

segments of the maxilliped in the female, the formula 11,1,5 on the third segment of the exopod of

leg 4, and the endopod of leg 4 with only a single

terminal element. There are several important differences, however. In Debruma the first and second

segments of the second antenna are clearly separated; the mandible has a base wider than the

blade (which lacks a distal spinous process); the

third segments of the exopods in legs 1-3 have

three outer spines instead of two; and legs 1-4

lack hyaline flanges.

Debruma clavelinae, new species

FIGURES 91-93

TYPE MATERIAL.—3 $ $, 1 c? > an d 6 copepodids

from the pharynx of the ascidian Clavelina picta

(Verrill), in 1 m, Swing Bridge, Ferry Reach, Bermuda, 15 July 1962, collected by AGH and RUG.

Holotype 9, allotype, one paratypic female, and

the copepodids deposited in USNM, the remaining

paratypic female (dissected) in the collection of

AGH.

FEMALE.—The body (figure 91a) is cyclopiform.

The length (not including the setae on the caudal

rami) is 1.14 mm (1.12-1.15 mm) and the greatest

width 0.45 mm (0.44-0.48 mm), based on three

specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.49:1. The ratio

of the length of the prosome to that of the urosome

is 1.33:1.

The segment of leg 5 (Figure 916) is 78 x 120 n.

There is no ventral, intersegmental sclerite between
FIGURE 91.—Debru ma clavelinae, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ; c,

area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum, ventral

(D) ; /, first antenna, dorsal (E) ; g, second antenna, anterior (D) ; h, labrum, ventral (F) ; i,

mandible, anterior (c) ; ;", first maxilla, anterior (c). Scale: A, 03 mm; B, 0.1 mm; c, 0.02

mm; D, F, 0.05 mm; E, 0.1 mm.
NUMBER 127 163

that segment and the genital segment. The genital

segment is elongated, 151 x 122 n, broadest in its

anterior half. The areas of attachment of the egg

sacs are located dorsolaterally near the middle of

the segment. Each area (Figure 91c) bears two

naked setae, 19 n and 23 n, with a minute jointed

process between them. The three postgenital segments are 52 x 70 n, 48 x 62 n, and 75 x 65

\t from anterior to posterior.

The caudal ramus (Figure 9Id) is moderately

elongated, 96 x 31 n, or 3.1 times longer than wide.

The outer lateral seta is 73 n and naked. The

short, stout, naked dorsal seta is 20 n. The outermost terminal seta is 96 n, the innermost 114 n,

both with lateral hairs proximally. The two median terminal setae (broken in most specimens) are

210 M^ (outer) and 260 n (inner), with inner and

outer lamellae respectively, but otherwise naked;

both of these setae inserted between dorsal (unornamented) and ventral (with a row of spinules)

flaps.

The body surface has very few hairs (sensilla).

The posteroventral margin of the anal segment

bears a row of minute spinules.

The egg sac is unknown.

The rostrum (Figure 91e) is linguiform, but

slender, with a narrow rounded tip.

The first antenna (Figure 91/) is 225 \i long. The

lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 19 (39 n

along the anterior margin), 62, 22, 39, 43, 31, and

19 ji respectively. The formula for the armature is

4, 13, 6, 3, 4 -f- 1 aesthete, 2 + 1 aesthete, and

7 + 1 aesthete. All the setae are naked.

The second antenna (Figure 91g) is 4-segmented.

Both the first and second segments bear an inner

naked seta. The third segment carries four setae,

the longest with minute barbs along the proximal

edge. The fourth segment, 64 n along its outer

edge, 44 n along its inner edge, and 19 n wide,

bears a strong recurved claw 25 n along its axis,

two weaker, less recurved claws, 36 n and 40 n,

and three naked setae, one of them weakly jointed.

The second segment is ornamented along its outer

margin with small spinules.

The labrum (Figure 9lh) has two rather narrow

posteroventral lobes, the indentation between them

not acute and having a small rounded median lobe.

The mandible (Figure 91i) has a moderately

broad base which carries a slender attenuated blade

having short dentiform spines on its convex side

and longer spinules on its concave side. Paragnaths were not identified in the specimens available. The first maxilla (Figure 91;) bears a single

naked seta. The second maxilla (Figure 92a) is

2-segmented. The first segment is unornamented.

The second segment bears two naked setae and

terminates in a long lash with short spinules along

one edge. The maxilliped (Figure 92&) is 2-segmented, the original third and fourth segments

having fused. The compound second segment bears

about midway two naked setae and distally two

small spiniform elements, the tip of the segment

being finely spinulose.

The ventral area between the maxillipeds and

the first pair of legs (Figure 92c) is not protuberant. There is a sclerotized line (incomplete medially) between the bases of the maxillipeds.

Legs 1-4 (Figure 92d-g) are segmented and

armed as in Contomolgus lokobeensis, but the third

segment of the exopod of leg 4 is 11,1,5 and the

endopod of leg 4 is 0-1 ;I. The inner seta on the

coxa of legs 1-3 is large and feathered, but in leg 4

this seta is shorter (24 n) and naked. The inner

margin of the basis of all four legs is haired. The

exopod of leg 4 is 148 ji long. The first segment

of the endopod is 24 x 23 n, with its inner distal

plumose seta 50 n. The second segment is 55 n

long, including the terminal spiniform processes,

and 22 n in greatest width. The outer margin of

this segment may be notched (as in Figure 92g)

or smooth (as in Figure 92A). (One female had

both endopods notched; two females had only the

right endopod notched.) The endopod bears terminally a fringed spine, 44 n. The outer margins of

both segments of the endopod are haired.

Leg 5 (Figure 93a) has an unornamented free

segment 37 x 29 n in greatest dimensions. The two

terminal elements are very unequal, the long naked

seta is 96 n and the short spine is 17 n with a fringe

on its outer side. On the body near the insertion

of the dorsal seta there is a group of small spinules.

Leg 6 is probably represented by the two setae

near the attachment of each egg sac (Figure 91c).

The color in life in transmitted light is opaque,

the eye red.

MALE.—The body (Figure 936) is a little more

slender than in the female. The length (not including the ramal setae) is 0.66 mm and the greatest width 0.20 mm, based on the allotype in lactic
SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

FIGURE 92.—Debruma clavelinae, new species. Female: a, second maxilla, posterior (c); b,

maxilliped, antero-inner (c); c, area between maxillipeds and first pair of legs, ventral (E) ; d,

leg 1 and intercoxal plate, anterior (r) ; e, leg 2 and intercoxal plate, anterior (E) ; /, endopod

of leg 3, anterior (E) ; g, leg 4 and intercoxal plate, anterior (E) ; h, endopod of leg 4, anterior

(F) . Scale: c, 0.02 mm; E, 0.1 mm; F, 0.05 mm.
NUMBER 127 165

FIGURE 9$.—Debruma clavelinae, new species. Female: a, leg 5, dorsal (F) . Male: b, dorsal

(c); c. maxilliped, first segment incomplete, posterior (c); d, endopod of leg 1, anterior (F) .

Scale: c, 0.02 mm; F, 0.05 mm; c, 02 mm.

acid. The ratio of the length to the width of the

prosome is 2.0:1. The ratio of the length of the

prosome to that of the urosome is 1.58:1.

The allotype was not dissected and therefore not

all parts were studied in detail. The following

notes are based on examination of the entire animal

in lactic acid.

The rostrum is like that of the female. The first

antenna resembles that of the female but has three

additional aesthetes (Figure 93fr) so that the formula is 4, 13 -|- 2 aesthetes, 6, 3 + 1 aesthete,

4 -|- 1 aesthete, 2 -f 1 aesthete, and 7+ 1 aesthete.

The second antenna is like that of the female.

The labrum resembles that of the female.

The maxilliped (Figure 93c) has a long recurved

claw.

Legs 1-4 are like those in the female. There is

only slight sexual dimorphism in the endopod of

leg 1, where the formula for the armature is the

same as in the female, but the spine on the third

segment (Figure 93d) is a little longer in relation

to the length of the segment.

The caudal ramus resembles that of the female.

The color is similar to that of the female.

ETYMOLOGY.—The specific name is the genitive

form of the generic name of the host.

Genus Dor idle ola Ley dig, 1853

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded.
166 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

First antenna 7-segmented, in the female with the

armature 4, 13, 6, 3, 4 -f- 1 aesthete, 2 + 1 aesthete,

and 7 + 1 aesthete; in the male with 4, 13 -f- 2

aesthetes (1 aesthete in D. astrophyticus), 6, 3 -f- 1

aesthete, 4 -f 1 aesthete, 2 + 1 aesthete, and 7

-f- 1 aesthete (so far as known). Second antenna

4-segmented, the formula 1, 1, 3, and two terminal

claws -|- several small elements (in D. fishelsoni

with one claw and a long spine).

Labrum incised medially. Mandible with the

basal region beyond the indentation having on its

convex side a scalelike area with spinules often

followed by a serrated fringe and on its concave

side a row of spinules; lash long. Paragnath a small

hairy lobe. First maxilla with three or four elements. Second maxilla of the usual lichomolgid

type. Maxilliped in the female 3-segmented with

a pointed tip; in the male 4-segmented (assuming

that the proximal part of the claw represents a

fourth segment).

Legs 1-4 with 3-segmented rami except leg 4

endopod which is 2-segmented. Armature of the

typical lithomolgidiform type. Leg 4 exopod with

the third segment having 111,1,5. Leg 4 endopod

with the formula 0—I ;II, the seta being feathered

(except in D. astrophyticus, D. foxi, and perhaps

D. buddhensis). Leg 1 endopod of the male with

the third segment, as far as known, having the formula 1,1,4 instead of 1,5 as in the female, except

in D. magnificus where both sexes have 1,5. (In D.

gemmatus this segment is 1,11,3 in the male). Leg

5 with a free segment bearing two terminal eleKey to Species of the Genus Doridicola

FEMALES

1. Body length S mm; caudal ramus with ratio 7.8:1 D. mmgnificus

Body length 2 mm or less; caudal ramus with ratio less than 3.5:1 2

2. Second antenna terminally with a claw and a long spine D. fnheltoni

Second antenna with two claws 3

3. Second maxilla with second tooth on lash much larger than the others in the series

D. astrophyticus

Second maxilla with teeth on lash evenly graded 4

4. Two claws on second antenna setiform, long and slender D. brevipet

Two claws on second antenna stouter and clearly unguiform 5

5. Free segment of leg 5 with an inner proximal expansion sharply set off by a marginal

indentation 6

Free segment of leg 5 without an inner proximal expansion thus delimited 7

6. Two terminal elements on second segment of leg 4 endopod very unequal (2.7:1); inner

seta on first segment of this endopod naked D. foxi

Two terminal elements on second segment of leg 4 endopod more nearly equal (2.1:1);

inner seta on first segment of this endopod feathered D. bulbipts

7. Inner margin of free segment of leg 5 with a distinct notch. D. audens

Inner margin of free segment of leg 5 smooth 8

8. Two setae on second segment of maxilliped nearly equal in length D. buddhensu

Two setae on second segment of maxilliped very unequal in length 9

9. Genital segment in dorsal view with subparallel sides 10

Genital segment in dorsal view with rounded lateral expansions 12

10. Caudal ramus with ratio 1.8:1; free segment of leg 5 with many small spinules D. gemmatus

Caudal ramus only a little longer than wide, or about as long as wide, free segment of leg

5 apparently smooth 11

11. Caudal ramus a little longer than wide; leg 4 endopod with second segment having the

two elements very unequal in length (2:1) D. rigidus

Caudal ramus about as long as wide; leg 4 endopod with second segment having the two

elements more nearly equal in length (1.3:1) D. trispinosus

12. Caudal ramus about as long as wide; body length 12 mm D. agilis

Caudal ramus clearly longer than wide (13-2:1); body length 15-2.3 mm 13

13. Two claws on second antenna nearly equal; two setae on second segment of maxilliped

about 2:1; medial margin of caudal ramus straight D. actiniae

Two daws on second antenna distinctly unequal; two setae on second segment of maxilliped about 5.6:1; medial margin of caudal ramus slightly bowed D. ptilosarci
NUMBER 127 167

merits and usually ornamented with spinules,

spines, or setules, and often having an inner proximal expansion. Leg 6 represented in both sexes

by the two setae near the genital openings.

Other features as in the species below.

Associated with actiniarians, octocorals, opisthobranch mollusks, crinoids, ophiuroids, and polychaetes.

TYPE-SPECIES.—Doridicola agilis Leydig.

REMARKS.—The histories of the genera Doridicola and Lichomolgus have been rather confused.

Leydig (1853) established his Doridicola agilis for

a copepod species found on the nudibranch Doris

lugubris at Trieste. His description and figures

were very incomplete. Six years later Thorell

(1859) described his new genus Lichomolgus based

on several copepods found in ascidians. Later,

various authors transferred Leydig's agilis to Lichomolgus (for example, Aurivillius, 1882b, 1883,

Canu, 1892, etc.).

The genus Doridicola was used by relatively few

authors after Leydig's original description. Both

Claus (1875) and Aurivillius (1882a) employed

this generic name. The work of Claus (1889) is of

special interest, since it contains a description (under the name Lichomolgus doridicola) of D. agilis

from Doris at the type locality. In this description

Claus confirmed the nature of the mandible (with

a sea lei ike area on the convex side of the base) and

the presence of two claws on the second antenna.

Canu (1892) also described these features.

It has been apparent for several years, following

the studies by Stock on European D. agilis, that

this species belongs to Doridicola, and that this

generic name should be retained. Other species

such as Thorell's Lichomolgus albens, having a

slender mandible without a scalelike area and lacking sexual dimorphism in leg 1, remain in Lichomolgus.

A key to the males of Doridicola is not included

here because sufficient information is not available

in existing descriptions and figures. The male is

unknown in four species: D. brevipes, D. buddhensis, D. fishelsoni, and D. trispinosus.

Doridicola agilis Leydig, 1853

Doridicola agilis Leydig, 1853, pp. 377-382, pi. 14, figs. 1-8

[from the nudibranch Doris lugubris, Trieste, Adriatic

Sea] .-Aurivillius, 1882a, pp. 34-39, pi. 5, figs. 1-12 [from

the nudibranchs Doris tuberculata Cuvier and Doris verrucosa Penn., Gullmarn, Sweden]; 1883, pp. 4-9, pi. 4,

fig. 9 [from Doris tuberculata and Doris verrucosa, Gullmarn, Sweden].-Carus, 1885. p. 352.-Graeffe, 1900, p. 41

(^Lichomolgus doridicola) [from Doridopsis limbata and

Doris tuberculata, Trieste, Italy].—Stock and Rleeton,

1963b, p. 258.

Doricola agilis.—Odhner, 1922, p. 33 [from the nudibranch

Facelina drummondi Thompson, Farsund, Norway].

Ergasilus.—Alder and Hancock, 1855, p. 27 [on the nudibranchs Doris tuberculata and Antiopa cristata, England].

Eolidicola tenax M. Sars, 1862, p. 135.

Doridicola.—Hancock and Norman, 1863, p. 49.—Claus, 1875,

pp. 348, 349, pi. 24, figs. 29, 29* [from Doris lugubris,

Trieste, Italy].

Lichomolgus chromodoridis Delia Valle, 1880a, pp. 101,

102, pi. 6, figs. 43-48 [from the nudibranch Chromodoris

elegans, Naples, Italy]; 1880b, pp. 120, 121, pi. 2, figs.

43-48 [from Chromodoris elegans, Naples, Italy].—Carus,

1885, p. 351.

Lichomolgus agilis.—Aurivillius, 1882b, pp. 42, 43, pi. 13,

fig. 9 [from Doris tuberculata and Doris verrucosa, Gullmarsfjord, Sweden]; 1883, pp. 38, 39, pi. 1, figs. 1-12 [from

Doris tuberculata and Doris verrucosa, Gullmarsfjorden,

Sweden].-Canu, 1892, pp. 228-230, pi. 22. figs. 1-14 [from

the gills of the nudibranchs Doris tuberculata Cuvier,

Doris millegrana Alder and Hancock, and Doris johnstoni

Alder and Hancock, on the dorsal papillae of Eolis corountil Forbes, Antiopa cristata Delia Chiaje, and Antiopa

hyalina Alder and Hancock, Boulogne coast of France, and

from Doris tuberculata and Doris johnstoni, Concarneau,

France]; 1894a, p. 3; 1894b, p. 137 [among algae, Isles

Chausey, northern France]; 1898, p. 399 [on Eolis drummondi W. Thompson, Saint-Marcouf, France].—T. Scott,

1894b, pp. 233, 258 (=Lichomolgus concinnus T. Scott);

1905, p. 207 [from Doris tuberculatus, Firths of Forth and

Clyde, Scotland]; 1906, pp. 352, 353 [from St. Monans,

Scotland, and from the gills of Doris (?) tuberculatus,

Granton, Scotland].—Herdman, 1895, p. 45 [in tow-net,

Morecambe Bay, near Port Erin, England].—Thompson,

1895, p. 102 [Morecambe Bay, England].—Gadeau de Kerrville, 1898, pp. 342, 343 [on Aeolis drummondi].—Cu£not,

1903, p. 20.-Farran, 1913, p. 6 [on Eolis papillosa, Ballynakill Harbor, Clare Island, Ireland].—G. O. Sars, 1917a,

pp. 161, 162, pi. 90 [from the nudibranchs Doris, Eolis,

and Polycera, Norway].—Flattely and Walton, 1922, p.

146.-Pelseneer, 1928, p. 179.—Monod and Dollfus, 1932a,

pp. 137, 138, figs. 3B, 8 [from the nudibranch Armina

verrucosa (Cantraine), coast of Morocco, 3O°34'3O'' N,

9

o

49/30w W; hosts listed as Aeolidia papillosa (Linnaeus),

Facelina longicornis (Montagu) = Eolidia coronata Forbes

and Goodsir, Facelina curia (Alder and Hancock) = Eolis

drummondi Thompson, Janolus hyalinus (Alder and Hancock) = Antiopa hyalina Alder and Hancock, Janolus

cristatus (Delia Chiaje) = Antiopa cristata Alder and

Hancock, Idulia coronata (Gmelin) = Doto coronata

(Gmelin), Armina verrucosa (Cantraine) = Pleurophyllidia pustolosa Bergh, Sphaerostoma hombergi (Cuvier) =

Tritonia hombergi Cuvier, Staurodoris verrucosa (Cuvier)

Bergh = Dorodigitata derelicta (Paul Fischer), Proctonotus mucronifer Alder and Hancock, Eolis auctorum,
168 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

sensu lato, Archidoris tuberculata (Cuvier), Archidoris

britannica (Johnston) Bergh = "Doris tuberculata Cuvier" Alder and Hancock, non Cuvier, Thordisa millegrana (Alder and Hancock) Bergh, Jorunna tomentosa

(Cuvier) = Doris johnstoni Alder and Hancock, Glossodoris elegans Cantraine = Chromodoris elegans (Cantraine), Dendrodoris lugubris (I. L. C. Gravenhorst),

Euphurus claviger (O. F. Miiller) = Triopa clavigera

(O. F. M.) Alder and Hancock, Palio lessoni (d'Orbigny)

Bergh = Polycera ocellata Alder and Hancock, and Polycera sensu lato].—Leigh-Sharpe, 1933, p. 113 [from the

nudibranch Archidoris britannica (=tuberculata), Plymouth, England]; 1935, p. 48 [from the nudibranchs Aeolidia papillosa (Linnaeus), Archidoris britannica (Johnston) , Facelina longicornis (Montagu), Idulia (Doto)

coronata (Gmelin), Janolus cristatus (Delia Chiaje),

Janolus hyalinus (Alder and Hancock), Jorunna tomentosa (Cuvier), and Sphaerostoma (Tritonia) hombergi

Cuvier, England].-Wilson, 1936, p. 268 [free, 66° to 67°

off Cape Penryhn, Melville Peninsula, Canada].—Lang,

1949, p. 8 [from Archidoris tuberculata (Cuvier), Gullmar

Fjord, Bohus, Sweden].—Stock, 1952. p. 58 [from the nudibranch Aeolidia papillosa (Linnaeus), The Netherlands];

1959a, p. 45 [from the tectibranch Aplysia punctata Cuvier, Gulf of Naples, Italy]; 1960c, pp. 241, 242 [from the

gills of nudibranchs and sometimes tectibranchs, Archidoris tuberculata Cuvier, Doris ("StaurodorisT) verrueosa

(Linnaeus), Facelina drummondi (Thompson), and

Gasteropteron rubrum (Rafinesque) = G. meckeli (de

Blainville), Banyuls, Mediterranean coast of France];

1964a, p. 67.-Pruvot-Fol, 1954, pp. 25. 26 [cites ex litt.

the following hosts: Dendrodoris limbata, Limacia clavigera, Polycera tin bin, Polycera sp., Doris millegrana, Jorunna tomentosa, Doris verrueosa, Archidoris tuberculata,

Glossodoris valenciennesi, Tritonia hombergi, Janolus

hyalinus, Antiopella cristata, Facelina drummondi, Zephyrina pilosa, Armina tigrina, Armina verrueosa, Doto coronata, Facelina coronata].—Marine Biological Association,

1957, p. 176 [from Archidoris tuberculata (=Archidoris

pseudoargus), Archidoris britannica, and Jorunna tomentosa, Rum Bay, Plymouth, England].—Bresciani and

Liitzen, 1962, p. 377.—Bocquet and Stock, 1963. p. 295.-

Bocquet. Stock, and Kleeton, 1963, p. 38 [from the polychaete Polynoe sp., Rade de Brest, France].—Bruce, Colman, and Jones, 1963, p. 127 [on 11 species of nudibranchs.

Isle of Man, England].—Crothers, 1966, p. 52 [from Archidoris pseudoargus, Wales].—Swennen, 1967, p. 108 [on

Archidoris tuberculata, The Netherlands].-Glac,on, 1971,

p. 23 (unnumbered).—Shih, Figuiera, and Grainger. 1971,

p. 214.

Lichomolgus (Lichomolgus) agilis.—Capart, 1941, p. 172

[from the nudibranch Armina tigrina Rafinesque, Kasamanse, Senegal, 12°47' N, 17°7' W].

Lichomolgus doridicola.—Claus, 1889, pp. 2, 12, 13, pi. 2,

figs. 1-12.-Canu, 1891a, p. 478 [from the gills of the

nudibranchs Doris tuberculata, Doris johnstoni, and Doris

millegrana and from the dorsal papillae of the nudibranchs Antiopa hyalina and Eolis coronata, Boulogne

and Goncarneau, France]; 1899, p. 73.—Voigt, 1892, p.

35.—Hecht, 1895, pp. 624, 625 [from the nudibranchs

Doris tuberculata, Doris johnstoni, Triopa clavigera, Eolis

papillosa, Eolis coronata, Proctonotus mucroniferus, and

Antiopa cristata, Roscoff. France].—Eliot, 1910, p. 26.—

Hoffmann, 1926, p. 50.

non Lichomolgus agilis.—Herdman, 1893, p. 83.—T. Scott,

1892, pp. 266, 267. (=Herrmannella rostrata.)

Lichomolgus diordicola.—Cami, 1892, p. 338.

Lichomolgus concinnus (provisional name).—T. Scott, 1892,

p. 261 [off St. Monans, Firth of Forth, Scotland]; 1893b,

p. 212, pi. 7, figs. 12-15 [from Doris (T) tuberculata,

Granton. Scotland].

Doridicola sp. (cf. agilis Leydig, 1853)

Lichomolgus sp. (cf. agilis Leydig) .—Stock, 1960c, p. 242

[from the gills of cephalopod Ommastrephes sagittatus

(Lamarck), in the market at Rosas. Catalanian coast of

Spain].

REMARKS.—Stock (1960c) was unable to make a

definitive determination of the single female that

he found. Since I), agilis has thus far been found

only on opisthobranchs, its occurrence on a cephalopod would be very unusual. Because of the uncertain status of this form it has not been included

in the key to species of Doridicola.

Doridicola actiniae (Delia Valle, 1880)

non Doridicola antheae Ridley. 1879. p. 458 [from the ac

tiniarian Anthea cereus, near Ilfracombe, North Devon,

England (perhaps = Paranthessius anemoniae Claus, 1889) ].

Lichomolgus actiniae Delia Vallc. 1880a, pp. 97-99, pi. 6,

figs. 27-32 [from the actiniarian Actinia concentrica var.

viridis, Naples, Italy]; 1880b, pp. 118-119. pi. 1, figs.

27-30, pi. 2, figs. 31-32 [from Actinia concentrica var.

viridis, Naples, Italy].-Carus, 1885, p. 351.—Canu. 1898. pp.

399-401, pi. 6, figs. 1-8 [in plankton, Grandchamp-lesBains, France]; 1899, p. 76.—Gadeau de Kerville, 1898, p.

343.-De Zulueta, 1912, pp. 9, 10, 50-Stock, 1960c. pp.

234-238, figs. 9, 10 [from the actiniarians Actinia equina

Linnaeus and Anemonia sulcata (Penn.) , Banyuls, Mediterranean coast of France, and from Anemonia sulcata,

Split, Jugoslavia].—Carton, 1963, p. 1148 [from Anemonia

sulcata and Actinia equina, Banyuls, France, and from

Anemonia sulcata, Roscoff, France].—Bouligand, 1966, p.

269, pp. 301, 302 [from Actinia equina and Anemonia

sulcata, Banyuls, France].

Lichomolgus anemoniae Claus, 1889, pp. 13, 14, pi. 3, figs.

1-7 [from the actiniarian Anemonia sulcata, Trieste, Italy].

Lichomolgus fucicolus (non Brady).—Canu. 1899, p. 76.

Doridicola astrophyticus, new species

FIGURES 94-96

TYPE MATERIAL.—91 9 $, 98 cf cf from eight

basket stars, Astrophyton muricatum (Lamarck), in
NUMBER 127 169

7 m, Oistin's Bay, Christchurch, Barbados, 15 July

1959, collected by AGH and RUG. Holotype ? ,

allotype, and 183 paratypes (88 9 9, 95 cf cf) deposited in USNM; other paratypes (dissected) in

the collection of AGH.

OTHER MATERIAL EXAMINED (all from Astrophyton muricatum).—Barbados: 12 9 9> 9 copepodids

from single host, in 2 m, Six Men's Bay, St. Peter,

21 June 1959; and 8 $ $, 13 <$ <$ from single

host, in 5 m, Carlisle Bay, Bridgetown, 16 June

1959.

Puerto Rico: 7 9 9, 7 cf cf. and 16 copepodids

from single host, in 2 m, Corona del Diablo Reef,

south of La Parguera, southwestern Puerto Rico,

24 August 1959; 23 $ ?, 32 cf cf, and 5 copepodids

from 19 hosts, in 4 m, lee side of Terremoto Reef,

southeast of La Parguera, 6 August 1959; 2 cf cf

from two hosts, in 9 m, south side of Terremoto

Reef, 6 August 1959; 6 9 9, 17 cf cf from 20 hosts,

in 3 m, Terremoto Reef, 9 August 1959; 27 9 ?,

37 cf c? from 18 hosts, in 3 m, Laurel Reef, west

of La Parguera, 13 August 1959; 4 ? $, 11 cf cf.

and 2 copepodids from 19 hosts, in 7 m, north side

of Terremoto Reef, 15 August 1959; and 4 9 $,

13 cf cf from three hosts, in 4 m, Terremoto Reef,

6 August 1959.

Jamaica: 14 9 9» 41 cf cT from seven hosts, in

3 m, Lime Cay, near Kingston, 30 August 1959;

and 9 9 9» 14 cf cf from single host, in 1.5 m,

Maiden Cay, near Kingston, 4 September 1959. All

collected by AGH and RUG.

FEMALE.—The body (Figure 94a) has the prosome not unusually expanded. The length (not

including the setae on the caudal rami) is 1.30

mm (1.22-1.40 mm) and the greatest width 0.53

mm (0.48-0.56 mm), based on ten specimens in

lactic acid. The ratio of the length to the width

of the prosome is 1.44:1. The ratio of the length

of the prosome to that of the urosome is 1.43:1.

The segment of leg 1 is separated from the cephalosome by a transverse dorsal furrow. The epimeral

areas of the pedigerous segments are more or less

rounded.

The segment of leg 5 (Figure 94b) is 99 x 172 i*.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is elongated, 213 x 156 n. The areas of

attachment of the egg sacs are located dorsolaterally

just in front of the middle of the segment. Each

area (Figure 94c) bears two small naked setae, 7 n

and 14 n. The three postgenital segments are 78 x

86 n, 67 x 78 n, and 70 x 81 n from anterior to posterior. The anal segment has on its posteroventral

border on each side a row of minute spinules.

The caudal ramus (Figure 94d) is 66 x 31 n, or

2.13 times longer than wide. The outer lateral

seta is 165 n and naked, the dorsal seta 45 v and

lightly feathered, the outermost terminal seta 130

n with hairs along the inner side proximally, the

innermost terminal seta 208 n and plumose proximally. The two long median terminal setae are

440 n (outer) and 640 n (inner), both plumose and

inserted between dorsal (unornamented) and ventral (with a row of minute spinules) flaps.

The egg sac (Figure 94a) is globular, scarcely

reaching to the anal segment, in dorsal view 340 x

240 n in a specimen 1.28 mm long, and 260 x 187 n

in a specimen 1.20 mm long. There are numerous

eggs about 52 n in diameter.

The rostrum (Figure 94e) is broadly rounded

posteroventrally.

The first antenna (Figure 94/) is 516 n long.

The lengths of its seven segments (measured along

their posterior nonsetiferous margins) are 44 (83 n

along the anterior margin), 126, 31, 88, 78, 62, and

48 (i respectively. The formula for the armature

is 4, 13, 6, 3, 4 -j- 1 aesthete, 2 + 1 aesthete, and

7 —f— 1 aesthete. All the setae are naked.

The second antenna (Figure 94g) is 4—segmented.

The first two segments bear a small inner seta.

The third segment bears three setae, one of them

bent. The fourth segment, 110 n along its outer

edge, 77 n along its inner edge and 26 |i wide, has

two terminal claws, the more slender one 42 n

along its axis, the stouter one 43 n, and five slender

setae. All the setae are naked.

The labrum (Figure 94/i) has two linguifonn

posteroventral lobes.

The mandible (Figure 95a) has on the convex

edge of the base a scalelike area with a curved row

of spines followed by a serrated lamella; the

concave edge has a shallow indentation followed

by a transverse row of slender spinules which become coalesced. The lash is moderately long and

barbed. The paragnath (Figure 94fi) is a slender

hairy lobe. The first maxilla (Figure 956) has three

elements. The second maxilla (Figure 95c) has an

unornamented first segment. The second segment

has a proximal seta on the outer (ventral) surface,

a surficial posterior barbed seta, and a distal inner
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 94.—Doridicola astrophyticus, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ram us, dorsal (o) ; e, rostrum,

ventral (E) ; /, first antenna, with arrows indicating positions of additional aesthetes in male,

dorsal (E); g, second antenna, anterior (F) ; h, labrum, with paragnaths indicated by broken

lines, ventral (D) . Scale: A, OJ} mm; B, E, 0.2 mm; c, 0.05 mm; D, F, 0.1 mm.
NUMBER 127

FIGURE 95.—Doridicola astrophyticus, new species. Female: a, mandible, posterior (c); b, first

maxilla, posterior (c); c, second maxilla, posterior (c); d, maxilliped, posterior (c); e, area

between maxillipeds and first pair of legs, ventral (F) ; /, leg 1 and intercoxal plate, anterior

(

F

); & kg 2 and intercoxal plate, anterior (F) ; h, endopod of leg S, anterior (F) . Scale:

c, 0.05 mm; F, 0.1 mm.
172 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

(dorsal) seta with slender spinules along one edge,

and it terminates in a short lash whose second

tooth is much larger than the rest. The maxilliped

(Figure 95d) has an unornamented first segment.

The second segment bears two very unequal, finely

barbed setae and a group of posterior surficial

spinules. The third segment has a barbed spine and

a slender setule and terminates in a barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 95<?) is not protuberant.

A sclerotized line connects the bases of the maxillipeds.

Legs 1-4 (Figures 95f-h, 96a) have the usual

segmentation and armature of the genus. The inner

coxal seta is long and plumose in legs 1-3 but short

(17 n) and naked in leg 4. The inner margin of

the basis is haired in legs 1-3 but naked in leg 4.

The three spines on the last segment of the endopod

of leg 3 (Figure 95/J) are 46, 50, and 64 n from

outer to inner. The exopod of leg 4 is 177 M- long.

The endopod of leg 4 (Figure 96a) has the first

segment 51 x 37 n (without the spiniform processes), with its distal inner naked seta 29 n. The

second segment is 110 n long (including the

spinous processes) and 21 n wide at the middle, its

two terminal fringed spines 64 n (outer) and 83 n

(inner). Both segments bear outer hairs, and the

second segment has spinules near the insertions

of the spines.

Leg 5 (Figure 96b) has a recurved free segment,

156 x 47 n, one of its two terminal elements 55 u

and naked, and the other 96 n with a slight lamella.

The segment is ornamented outwardly with strong

spines. The adjacent seta on the body is relatively

short and finely haired, with a group of spinules

near its insertion.

Leg 6 is represented by the two setae near the

area of attachment of each egg sac (Figure 94c).

The color in life in transmitted light is very

light brown, the eye red, the egg sacs dark brown.

MALE.—The body (Figure 96c) resembles that

of the female in general form. The length (without

the ramal setae) is 1.04 (1.01-1.10 mm) and the

greatest width 0.36 mm (0.35-0.37 mm), based on

ten specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.52:1. The ratio of

the length of the prosome to that of the urosome

is 1.18:1.

The segment of leg 5 (Figure 96d) is 44 x 112 i*.

There is no ventral intersegmental stierite. The

genital segment is 195 x 188 n. The four postgenital

segments are 47 x 70 n, 47 x 68 n, 45 x 62 \i, and 57

x 65 n from anterior to posterior.

The rostral area is like that of the female.

The first antenna is like that of the female but

has two additional aesthetes (at points indicated

by arrows in Figure 94/) so that the formula is

4, 13 -4- 1 aesthete, 6, 3 -f- 1 aesthete, 4 -f- 1 aesthete, 2 -+- 1 aesthete, and 7 -+- 1 aesthete.

The second antenna resembles that of the female but the inner surface of the second segment

is squamous (Figure 96c).

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 96/) is slender. The first

segment has a small inner distal spiniform process.

The second segment is expanded inwardly, bears

two setae, one slender and naked and the other

stouter and finely barbed, and two rows of spines.

The third segment is unornamented. The claw is

200 (i along its axis, with a weak division about

midway, and bears two very unequal proximal

setae.

Leg 1 has an exopod like that in the female; the

endopod (Figure 96g) has the last segment with

the formula 1,1,4, the outer spine being stouter

than in the female and having more slender barbules. Leg 2 has an exopod like that in the female; the endopod (Figure 96/J) has the middle

spine on the last segment curiously bent. Legs 3

and 4 are like those of the female.

Leg 5 (Figure 96d) has a rectangular free segment, 43 x 12 M, ornamented with a few small

spines; its two terminal elements are 46 n and 35 I*.

Leg 6 (Figure 96d) consists of a posteroventral

flap on the genital segment, bearing two naked

setae 30 n and 44 n.

The spermatophore is unknown.

The color resembles that of the female.

ETYMOLOGY.—The specific name astrophyticus

is a modification of the generic name of the host

(adapted from cpuxixos, of plants).

REMARKS.—Doridicola astrophyticus differs from

all other members of the genus in the much enlarged tooth on the lash of the second maxilla in

both sexes. Another distinctive character is the

naked seta on the first endopod segment of leg 4.

In all species except D. foxi (Gurney, 1927) and
NUMBER 127 173

s

FIGURE 96.—Doridicola astrophyticus, new species. Female: a, leg 4 and intercoxal plate, anterior (F) ; b, free segment of leg 5, inner (F) . Male: c, dorsal (A) ; d, urosome, dorsal (E) ; e,

second segment of second antenna, posterior (c); /, maxilliped, inner (F) ; g, endopod of leg

1, anterior (D) ; h, endopod of leg 2, anterior (D). Scale: A, 0.5 mm; c, 0.05 mm; D, F, 0.1 mm;

E, 02 mm.
174

D. buddhensis (Thompson and A. Scott, 1903) this

seta is feathered. These two species may be separated from D. astrophyticus, however, by the ratio

of the length to the width of the caudal ramus

(quadrate in D. buddhensis, 1.16:1 in D. foxi).

Doridicola audens (Humes, 1959)

Lichomolgus audens Humes, 1959, pp. 305-309, figs. 97-124

[from the nudibranch Platydoris scabra (Cuvier), region

of Nosy Be. Madagascar].

Doridicola brevipes (Shen and Lee, 1966)

Macrochiron (Macrochirori) brevipes Shen and Lee, 1966,

pp. 218, 219, 223, figs. 18-26 [from plankton, Chaikiang

River, Kwantung Province, China].

REMARKS.—The male is unknown.

Doridicola buddhensis (Thompson and A. Scott,

1903)

Lichomolgus buddhensis Thompson and A. Scott, 1903, p.

279, pi. 15, figs. 18-24 [from washings of dredged invertebrates, Ceylon].-A. Scott, 1909, pp. 263, 264 [in washings

of dredged invertebrates, Siboga Stations 164 and 226,

midway between the Lucipara and Schildpad Islands].

Macrochiron buddhensis.—Stock, 1957, p. 381.

REMARKS.—The male is unknown.

Doridicola bulbipes (Stock and Kleeton, 1963)

Lichomolgus bulbipes Stock and Kleeton, 1963b, pp. 251-257,

figs. 4-6 [from the alcyonaceans Alcyonium acaule Marion

and Parerythropodium coralloides (Pall.), vicinity of Banyuls, France, and Cap Creus, Spain].

Doridicola fishelsoni (Stock, 1967)

Lichomolgus fishelsoni Stock, 1967b, pp. 574-578, figs. 4, 5

[from the crinoid Oligometra serripinna (P. H. Carpenter),

Gulf of Aqaba, Israel].

REMARKS.—The male is unknown.

Doridicola foxi (Gurney, 1927)

Lichomolgus foxi Gurney, 1927, pp. 468, 469, fig. 113 [from

Suez Canal].-Humes and Ho, 1968c, pp. 655-662, figs.

67-88 [from the alcyonaceans Cladiella krempfii Hickson,

Cladiella ladniosa (Tixier-Durivault), and Cladiella

pachyclados (Klunzinger), region of Nosy Be\ Madagascar].

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

NEW RECORD.—From Cladiella laciniosa: 25 9 9»

9 cT cT» and 4 copepodids, in 2 m, west of Pte. de

Tafondro, Nosy Be, Madagascar, 30 May 1967.

NEW HOSTS.—From Cladiella latissima (TixierDurivault): 55 9 $, 43 o* cf» and 45 copepodids, in

1 m, Pte. de Tafondro, Nosy Be, Madagascar, 12

July 1967; 2 9 9 , 2 cf d\ »" 18 m, Bane du Touareg, near Nosy Be, Madagascar, 11 July 1967.

From Cladiella sphaerophora (Hemprich and

Ehrenberg): 14 9 9 , 24 <$ cf, and 24 copepodids,

in 1 m, Nosy N'Tangam, near Nosy Be, Madagascar,

21 July 1967. All collected by AGH.

Doridicola gemmatus (Humes, 1964)

Lichomolgus gemmatus Humes, 1964, pp. 60-62, fig*. 1-42

[from the actiniarian Stoichactis giganteum (Forskal).

Nosy Bt\ Madagascar].-Bouligand, 1966, p. 269.

REMARKS.—This species is unusual in its formula

for the third segment of leg 1 endopod in the

male (1,11,3).

Doridicola magnificus (Humes, 1964)

Lichomolgus magnificus Humes, 1964, pp. 63-65, figs. 43-73

[from the actiniarian Stoichactis giganteum (Forskal).

Nosy Hr, Madagascar]—Bouligand, 1966, p. 269.

REMARKS.—This species is unique in Doridicola

(so far as known) in lacking sexual dimorphism in

the formula for the third segment of leg 1 endopod.

Doridicola ptUosarci, new species

FICURES 97-100

TYPE MATERIAL.—47 9 9 , 11 cf cf» and 2 copepodids between the flaps and from the surface of the

stem of the sea pen Ptilosarcus gurneyi Gray

[=LeioptiIum quadrangulare (Moroff)], Friday

Harbor, San Juan Island, Washington, 17 August

1949, collected by J. L. Mohr. USNM 184568.

FEMALE.—The body (Figure 97a) has a moderately broad prosome. The length (not including

the ramal setae) is 1.94 mm (1.82-2.08 mm) and

the greatest width 0.85 mm (0.83-0.90 mm), based

on ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.36:1. The

ratio of the length of the prosome to that of the

urosome is 1.74:1. The segment of leg 1 is separated

from the cephalosome by a transverse dorsal furrow. The epimera of legs 1-4 are rounded.
NUMBER 127

FIGURE 97.—Doridicola ptilosarci, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, egg sac, with

outline of adjacent urosome, dorsal (A) ; /, rostrum, ventral (E); g, first antenna, with arrows

indicating positions of additional aesthetes in male, dorsal (E) ; h, second antenna, posterior

(F) ; i, labrum, ventral (D) . Scale: A, B, 0.5 mm; c, D, 0.1 mm; E, F, 0.2 mm.
176 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The segment of leg 5 (Figure 976) is 109 x 224 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 292 x 260 n in greatest dimensions, a

little longer than wide, in dorsal view swollen

laterally in its anterior half. The areas of attachment of the egg sacs are located dorsolaterally near

the middle of the segment. Each area (Figure 97c)

bears two naked setae about 20 p. long. The three

postgenital segments are 127 x 174 n, 96 x 164 p,

and 130 x 182 n from anterior to posterior. The

posteroventral border of the anal segment bears on

both sides a row of small, slender spinules.

The caudal ramus (Figure 97d) is 107 x 70 n,

about 1.5 times longer than wide. The outer lateral

seta is 114 n, the dorsal seta 66 ji, the outermost

terminal seta 130 n, the innermost terminal seta

177 ji, and the two long median terminal setae

470 n (outer) and 640 n (inner), both inserted

between smooth dorsal and ventral flaps. All these

setae are naked except the innermost terminal one

which has an inner proximal row of slender spinules. The dorsal surface of the ramus bears a few

hairs.

The body surface has a few hairs (sensilla) as

indicated in Figure 97a,b.

The egg sac (Figure 97e) is ovoid, 640 x 450 >i,

reaches as far as the anal segment, and contains

many eggs about 57 n in diameter.

The rostrum (Figure 97/) is rounded, with its

posteroventral margin incompletely defined.

The first antenna (Figure 97g) is slender, 597 n

long. The lengths of the seven segments (measured

along their posterior nonsetiferous margins) are

47 (99 n along the anterior margin), 159, 44, 94,

86, 65, and 50 n respectively. The formula for the

armature is 4, 13, 6, 3, 4 -{- 1 aesthete, 2 -f 1

aesthete, and 7 -f 1 aesthete. All the setae are

naked except four on the terminal segment which

are lightly haired.

The second antenna (Figure 97h) bears a very

finely barbed inner seta on each of the first two

segments. The third segment bears two slender

setae and a spiniform seta, all naked. The fourth

segment, 140 n along its outer edge, 104 n along

its inner edge, and 39 n wide, bears terminally two

unequal claws 58 » and 36 n along their axes and

five small hyaline elements.

The labrum (Figure 97i) has two rounded posteroventral lobes.

The mandible (Figure 98a) resembles in general

features that of Paradoridicola squamiger (Humes

and Frost, 1964). The transverse row of spinules

beyond the indentation turns anteriorly at the base

of the lash. The paragnath (Figure 986) is a small

hairy lobe. The first maxilla (Figure 98c) bears

four elements. The second maxilla (Figure 98d)

has a row of a few minute spinules on the outer

(ventral) surface of the first segment. The second

segment bears proximally on its outer (ventral)

margin a short, naked setule, on its posterior surface a seta with a few hairs, and on its inner

(dorsal) margin a spinelike seta with long strong

spinules. The terminal lash has a row of graduated

teeth. The maxilliped (Figure 98^) has two very

unequal setae on the second segment, the longer

one finely barbed, the shorter one naked. The third

segment, bearing a slender setule and a barbed

spine, terminates in a barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 98/) is only slightly

protuberant, with a fine line connecting the bases

of the maxillipeds.

Legs 1-4 (Figures 98g-i, 99a) have the same

segmentation and armature as in P. squamiger. The

inner coxal seta of legs 1-3 is long and plumose,

but in leg 4 this seta is short (16 io and naked. The

inner margin of the basis is haired in legs 1-3 but

naked in leg 4. The exopod of leg 4 is 265 n long,

with the third segment having the formula 111,1,5.

The first segment of the endopod (Figure 99a) is

60 x 36 n (not including the spiniform processes)

and its inner distal plumose seta 51 u. The second

segment is 128 x 30 n, including the spiniform

processes. The two terminal barbed spines are 70 n

(outer) and 120 n (inner). Both segments have an

outer row of hairs.

Leg 5 (Figure 996) has a free segment 125 I*

long, 47 n in greatest width proximally, and 23 n

in least width distally. The two terminal naked

setae are 143 n and 88 n. The segment is ornamented with small spinules along its outer surface.

The adjacent seta on the body is 65 n and naked.

Leg 6 is represented by two setae near the attachment of each egg sac (Figure 97c).

The color in life is unknown.

MALE.—The body (Figure 99c) resembles in general shape that of the female. The length (without

the setae on the caudal rami) is 1.57 mm (1.44—

1.63 mm) and the greatest width 0.58 mm (0.55-
NUMBER 127

FIGURE 98.—Doridicola ptilosarci, new species. Female: a, mandible, posterior (c); b, paragnath,

posterior (c); c, first maxilla, posterior (c); d, second maxilla, posterior (D) ; e, maxilliped,

anterior (c); /. area between maxillipeds and leg 1, ventral (E) ; g, leg 1 and intercoxal plate,

anterior (E) ; h, leg 2, anterior (E) ; i, endopod of leg 3, anterior (E) . Scale: c, D, 0.1 mm;

E, 02 mm; c, 0.05 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 99.—Doridicola ptilosarci, new species. Female: a, leg 4 and intercoxal plate, anterior

(E) ; b, leg 5, dorsal (D) . Male: c, dorsal (A) ; d, urosome, dorsal (B) ; e, second segment of

second antenna, posterior (D) ; /, maxilliped, postero-inner (F) ; g, second segment of maxilliped,

inner (D) ; h, barbed seta on second segment of maxilliped, outer (H) ; i, third segment of

endopod of leg 1, anterior (D) . Scale: A, B, 05 mm; D, 0.1 mm; E. F, 02 mm; H, 0.02 mm.
NUMBER 127 179

0.63 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome

is 1.54:1. The ratio of the length of the prosome

to that of the urosome is 1.52:1. The form of the

epimera of the pedigerous segments is slightly

different from that in the female.

The segment of leg 5 (Figure 99d) is 55 x 159 n.

There is no ventral intersegmental sclerite. The

genital segment is 308 x 326 »A. The four postgenital

segments are 55 x 122 n, 57 x 117 », 42 x 109 n, and

78 x 112 M- from anterior to posterior.

The caudal ramus resembles that of the female,

but is smaller, 78 x 50 n.

The rostrum is like that of the female.

The first antenna is similar to that of the female, but there are two additional aesthetes on

the second segment and one additional aesthete on

the fourth segment (at locations indicated by

arrows in Figure 97g), so that the formula is 4,

IS -f- 2 aesthetes, 6, 3 -f 1 aesthete, 4 -f 1 aesthete, 2 + 1 aesthete, and 7 -f 1 aesthete.

The second antenna is similar to that of the

female, but the inner surface of the second segment bears small slender needlelike spinules (Figure 99<r).

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 99/) is long and slender.

The second segment is stouter in an inner view

(Figure 99g) than in a postero-inner view. It bears

two rows of spines and two setae, one of them

naked, the other barbed (Figure 99h). The claw,

280 n along its axis and weakly divided about

midway, is slightly swollen proximally and carries

the usual two unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented as in the female and have

the same spine and setal formula except for the

endopod of leg 1 where the last segment (Figure

99t) is 1,1,4. Slight sexual dimorphism occurs in

the endopod of leg 2 where the outer spine is

stouter than in the female (Figure 100a). Legs 3

and 4 are like those of the female.

Leg 4 (Figure 1006) has a free segment 54 x 20 n,

slenderer than in the female. The two terminal

setae are 72 n (naked) and 69 i* (with outer

barbules). The segment is outwardly spinulose as

in the female.

Leg 6 (Figure 100c) consists of a posteroventral

flap on the genital segment that bears two naked

setae, 75 n and 55 n, and shows an outer group

of undulating striations.

The spermatophore (Figure lOOd), attached to

the female in pairs, is 234 x 112 n, not including

the neck.

The color in life is unknown.

ETYMOLOGY.—The specific name, ptilosarci, is

based on the name of the pennatulacean genus

serving as host.

REMARKS.—Most species of Doridicola are distinctly smaller than D. ptilosarci, and only one, D.

magnificus (Humes, 1964), is larger. Two species,

D. fishelsoni (Stock, 1967) and D. actiniae (Delia

Valle, 1880), have a size similar to that in the new

species. Both of these species, however, may be

FIGURE 100.—Doridicola ptilosarci, new species. Male: a, third segment of endopod of leg 2,

anterior (D) ; b, leg 5, dorsal (c); c, leg 6, ventral (E) ; d, spermatophores, attached to female,

dorsal (E) . Scale: D, 0.1 mm; E, 02 mm; c, 0.05 mm.
180 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

readily distinguished from D. ptilosarci. In D. actiniae the length to width ratio of the female

caudal ramus is about 2.2:1 and the ramal setae

are haired. The relative length of the two setae

on the second segment of the female maxilliped

also serves as a distinguishing feature. In D. fishelsoni the last segment of the second antenna bears

a claw and a long spine, instead of two claws as

in the new species.

Doridicola rigidus (Ummcrkutty, 1962)

Macrochiron (Macrochiron) rigida Ummerkutty, 1962, pp.

47-52, pi. 9, figs. 6-9, pi. 10, figs. 1-6 [from washings of

weeds and invertebrates, mainly sponges. Gulf of Mannar,

southeastern India].

Doridicola trispinosus (Stock, 1959)

Lichomolgus trispinosus Stock, 1959b, pp. 64-66, figs. S, 4

[from the pennatulacean Pennatula rubra Ell., Gulf of

Naples, Italy].-Bouligand, 1966, p. 269.

REMARKS.—The male is unknown.

Genus Epimolgus Bocquet and Stock, 1956

DIAGNOSIS.—Body of the female somewhat transformed, with a moderately swollen prosome. Body

of the male cyclopiform. Urosome in the female

5-segmented, in the male 6-segmented. Caudal

ramus with six setae. Rostrum broadly rounded.

First antenna 7-segmented. Second antenna 4-segmented, robust, with terminally a large claw, a

smaller clawlike spine, and two setae.

Labrum indented medially. Mandible with a

slender base distally with a row of spinules on its

concave side and on its convex side; lash long and

pectinate. First maxilla with three elements. Second

maxilla of the usual lichomolgid type. Maxilliped

in the female 3-segmented, in the male 1-segmented

(assuming that the proximal part of the claw

represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod, which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1 ;II, the seta being feathered.

Leg 1 endopod of the male with an armature like

that in the female. Leg 5 with a free segment

bearing two terminal elements. Egg sacs longer

than the body.

Other features as in the species below.

Associated with gastropod mollusks.

TYPE-SPECIES.—Epimolgus trochi (Canu).

REMARKS.—The male of E. orientalis is unknown.

Key to Species of the Genus Epimolgus

FEMALES

Prosomal segments bulging laterally; egg sacs arcuate E. orientalis

Prosomal segments not bulging laterally; egg sacs almost straight B. trochi

Epimolgus trochi (Canu, 1899)

FIGURE 101

Lichomolgus trochi Canu, 1899, pp. 75-79. pi. 8, figs. 1-10

[from the gastropod Trochus umbilicatus Linnaeus =

Gibbula urrbtltcalis (da Costa), Wimereux, channel coast

of France].-Dollfus, 1927, pp. 120, 121 [from Gibbula

(Steromphalus) cineraria (Linnaeus), Roscoff, northern

France].—Pelseneer, 1929, p. 44.—Gotto, 1962, p. 103.—

Arvy, 1963, p. 146.

Lichomolgus (Macrochiron) trochi.—Monod and Dollfus,

1932a, p. 141, figs. IA-F, 24G [from the pallial cavity of

Gibbula cineraria Linnaeus, Roscoff, northern France].

non Lichomolgus troc/ii.-Gaillard, 1953, p. 16 (—Trochicola

entericus) .

Lichomolgus (Epimolgus) (roc/i/.-Bocquet and Stock. 1956,

pp. 10-16, figs. 1,2 [from the gastropods Gibbula cineraria

(Linnaeus), Gibbula umbilicalis (da Costa), and Monodonta lineata (da Costa), northern France]—Stock, 1960c,

pp. 246, 247, fig. 15 [from pallial cavity of Gibbula varia

(Linnaeus), Banyuls, Mediterranean coast of France].

Epimolgus trochi.—Stock, 1957, p. 381.

Macrochiron trochi.—Stock, 1957, p. 381.

Epimolgus orientalis (Heegaard, 1962)

Alimeda orientalis Heegaard, 1962, pp. 155, 156, figs. 27-36

[from the gill flaps of the opisthobranch Aplysia, Port

Jackson, New South Wales].-Ho, 1970, p. 113.

REMARKS.—Heegaard placed this species in the

family Chondracanthidae, but it is obviously a

lichomolgid and as far as can be learned from the

original description it belongs in the genus Epimolgus.
NUMBER 127 181

FICURE 101.—Epimolgus trochi (Canu). Female: a, dorsal; b, lateral; c, second antenna; d,

mandible; e, leg 4; /, leg 5. (a, b, d, f from Bocquet and Stock, 1956, figs. la,b,e, 2f; c, e from

Canu, 1899, pi. 8. figs. 5, 9.) Length of female 1.95 mm.

The two females given the provisional name

Strongylopleura (?) pruvoti by Monod and Dollfus

(1932a, p. 204; 1932b, p. 295; 1934, pp 315, 316)

probably belong to Epimolgus (Bocquet and Stock,

1958c) but the available information leaves some

doubt about their generic position. Monod and

Dollfus (1934) record two molluscan hosts, Aplysia

sp., Noumea, New Caledonia, and Acclesia striata

Quoy and Gaimard, He des Pins, New Caledonia.

The male is unknown.

Genus Gelastomolgus Humes, 1968

DIAGNOSIS.—Body modified, with elongated prosome which in the female is pointed anteriorly and

broadened posteriorly and in the male is rounded

anteriorly and narrowed posteriorly. Segment of

leg 4 with a pair of ventrolateral lobes. Urosome

5-segmented in the female, 6-segmented in the

male. Caudal ramus with six setae. Rostrum broadly

rounded. First antenna 7-segmented, the armature

in both sexes being 4, 12, 6, 3, 4 -f- 1 aesthete,

2 -j- 1 aesthete, and 7 —|- 1 aesthete. Second antenna

4-segmented, but the first and second segments not

distinctly separated. Formula: 1, 1, 4, III -|- one

spine + three setae, there being three terminal

claws.

Lab rum with two lobes separated by a median

indentation. Mandible a broad elongated bipectinate blade rather abruptly attenuated distally;

toward the distal end of the blade a well-sclerotized

marginal spinous process. Paragnath a minute spinulose lobe. First maxilla with three setae. Second

maxilla of the usual lichomolgid type. Maxilliped

of the female with very weak armature and apparently 2-segmented, though cuticular creases
182 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

suggest further segmentation; in the male maxilliped 4-segmented (the proximal part of the claw

probably representing the fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Third exopod

segments of legs 1-4 bearing three spines, one fewer

than usual in lichomolgids. Leg 4 endopod with

0-1; 1. Formula for legs 1-4 in the male as in the

female. Leg 5 in both sexes with a free segment

bearing two terminal setae. Leg 6 in the female

represented by a spine and a seta near the area of

attachment of each egg sac; in the male by a weak

ventrolateral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated with bivalve mollusks.

TYPE-SPECIES.—Gelastomolgus spondyli Humes.

Gelastomolgus spondyli Humes, 1968

FIGURE 102

Gelastomolgus spondyli Humes, 1968b, pp. 66-74, figs. 1-36

[in the mantle cavity of the bivalve Spondylus variant

Sowcrby, region of Nosy Be, northwestern Madagascar].

NEW RECORD.— 1 9 from Spondylus variant, in

18 m, in lagoon west of beach between Parry Island

and Eniwetok Island, Eniwetok Atoll, Marshall

Islands, 10 July 1969, collected by AGH.

NEW HOST.—10 9 9 , 12 <? cf from the madreporarian Plerogyra sp., in 22 m, Tany Kely, near

Nosy Be, Madagascar, 17 June 1967, collected by

AGH. This host may be accidental. The coral was

taken from the type-locality of G. spondyli in close

proximity to living Spondylus varians.

Genus Haplomolgus Humes and Ho, 1968

DIAGNOSIS.—Body cyclopiform. Urosome 5-*egmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum linguiform.

First antenna 7-segmented, the armature in the

female being 4, IS, 6, S, 4 + 1 aesthete, 2 -f 1

aesthete, and 7 + 1 aesthete; in the male 4, 13 -f 2

aesthetes, 6,3+ 1 aesthete, 4 -j- 1 aesthete, 2 -{• 1

aesthete, and 7 -f 1 aesthete. Second antenna 4-

segmented; the armature 1, 1, 3, and I, there being

a single terminal claw without accessory elements.

FIGURE 102.-Gelastomolgus spondyli Humes. Female: a, dorsal; b, lateral; c, mandible; d,

maxilliped; e, leg 4 and intercoxal plate with adjacent process; /, endopod of leg 4. Male:

g, dorsal; h, lateral. (From Humes, 1968b, figs. 1, 2, 13. 17, 22, 23, 25, 26.) Length of female

1.16 mm, of male 0.59 mm.
NUMBER 127 183

Labrum with two widely divergent lobes separated by a median notch. Mandible having the

base divided by a constriction into a slender unornamented proximal part and a broadened distal

part, the latter bearing on its concave edge two

lobes, each with a row of spinules, and on its convex edge a large, pointed, posteriorly directed process followed by a row of small serrations; lash

moderately long. Paragnath a small hairy lobe.

First maxilla with four setae. Second maxilla of

the usual lichomolgid type. Maxilliped of the female 3-segmented, but in the male 4-segmented

(assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with S-segmented rami except for leg

4 endopod, which is a single unarmed segment that

often bears a small distal lobe. Leg 3 endopod

0-l;0-2;II,2. Formula of legs 1-4 of the male like

that of the female. Leg 5 in both sexes with a free

segment bearing two setae. Leg 6 in the female

represented by two setae near the area of attachment of each egg sac, in the male by a posteroventral flap on the genital segment with two setae.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Haplomolgus montiporae Humes

and Ho.

Haplomolgus montiporae Humes and Ho, 1968

FlCURE 103

Haplomolgus montiporae Humes and Ho, 1968a, pp. 375-

378, figs. 211-240 [associated with the hard corals Montipora sinensis Bernard, Montipora sp., and Montipora sp.

cf. stcllata Bernard, region of Nosy B£, northwestern

Madagascar].

NEW RECORD.—32 $ $, 12 cf cf from Montipora

sp., in 3 m, Ambatoloaka, Nosy B£, northwestern

Madagascar, 23 June 1967, collected by AGH.

Genus Indomolgus Humes and Ho, 1966

DIAGNOSIS.—Body modified, elongated. Urosome

5-segmented in the female, 6-segmented in the

male. Caudal ramus with six short setae. Rostrum

FIGURE 103.—Haplomolgus montiporae Humes and Ho. Female: a, dorsal; b, mandible; c, leg

4. Male: d, dorsal. (From Humes and Ho, 1968a. figs. 211, 220, 228, 233.) Length of female

0.86 mm, of male 0.87 mm.
184

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

linguiform or not well defined. First antenna 7-

segmented with short setae, the formula not the

same in all species; in the female 4, 11, 6, 3, 4 -j- 1

aesthete, 2 -+- 1 aesthete, and 7 + 1 aesthete, or,

3, 12, 6, 3, 4 -j- 1 aesthete, 2 -f aesthete, and

7 + 1 aesthete, or 4, 13, 6, 3, 4 -f 1 aesthete,

2 -j- 1 aesthete, and 7 + 1 aesthete; and in the

male 4, 11 -f 2 aesthetes, 6, 3 -f 1 aesthete, 4

+ 1 aesthete, 2 + 1 aesthete, and 7 -f- 1 aesthete, or 3, 12 -f 2 aesthetes, 6, 3 + 1 aesthete,

4 -{- 1 aesthete, 2+ 1 aesthete, and 7 -j- 1 aesthete, or 4, 13, 6, 3, 4 + 1 aesthete, 2 -f 1 aesthete,

and 7 -j- 1 aesthete, with no aesthetes added. Second antenna 4-segmented, the formula 1, 1, 3, and

II -|- several small elements, there being two terminal claws.

Labrum with the two lobes separated by an

indentation. Mandible with the basal part distal

to the indentation having on its convex side a

scalelike area either smooth or with spinules and

a fringelike process followed by a serrated fringe

and on its concave side a row of spinules; lash

moderately short or long. Paragnath a small hairy

lobe. First maxilla with three or four elements.

Second maxilla of the usual lichomolgid type.

Maxilliped in the female 3-segmented with pointed

tip, in the male 4-segmented (assuming that the

proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Leg 4 exopod

with the third segment 111,1,4, or 111,1,5, or 1,1,1,5.

Leg 4 endopod with the first segment having the

formula 0-1, but the second segment variable with

11,2, or 1,3, or 11,3. Armature of the legs in the

male like that in the female. Leg 5 with a free

segment bearing two setae. Leg 6 represented in

the female by the two setae and the process near

the area of attachment of each egg sac, in the male

by the ventral posterolateral flap on the genital

segment bearing two setae.

Other features as in the species below.

Associated with artiniarian and zoanthidean coelenterates.

TYPE-SPECIES.—Indomolgus brevisetosus Humes

and Ho.

REMARKS.—The four species assigned to the genus Indomolgus show considerable interspecific

variation in the armature of the first antenna and

leg 4, and on this basis a possible division of the

genus might be suggested. However, since the armature of both these appendages varies at times

intraspecificially, it would be unwise, as Humes and

Ho (1966a) have mentioned, to make generic distinctions based on the armature.

The following key to the females applies also

to the males of this genus, but information on the

male of /. panikkari is scanty.

Key to Species of the Genus Indomolgus

FEMALES

1. Caudal ram us longer than last two postgenital segment together /. panikkari

Caudal ramus shorter than last two postgenital segments together 2

2. Last segment of exopod of leg 4 with three outer spines; last segment of endopod of leg 4

with either 11,3 or 11,2 3

Last segment of exopod of leg 4 with 1,1,13; last segment of endopod of leg 4 with 13

I. mutatus

3. Last segment of exopod of leg 4 with 111,13; last segment of endopod of leg 4 with 113

I. brevisetosus

Last segment of exopod of leg 4 with 111,1,4; last segment of endopod of leg 4 with 11,2

l.dtoersus

Indomolgus brevisetosus Humes and Ho, 1966

FIGURE 104

Indomolgus brevisetosus Humes and Ho, 1966a, pp. 11-14,

figs. 56-85 [from the zoanthideans Patythoa tuberculosa

(Esper) and Palythoa liscia Haddon and Duerdon, region

of Nosy B£, Madagascar].

Indomolgus divers us Humes and Ho, 1966

Indomolgus diversus Humes and Ho, 1966a, pp. 14-17, figs.

86-114 [from Palythoa tuberculosa and Palythoa liscia,

region of Nosy Ik'-, Madagascar].
NUMBER 127 185

FICURE 104.—Indomolgus brexnsetosus Humes and Ho. Female: a, dorsal; b, lateral; c, mandible;

d, leg 4 and intercoxal plate. Male: e, dorsal. (From Humes and Ho, 1966a, figs. 56, 57, 67,

78, 80.) Length of female 2.02 mm, of male 2.20 mm.

Indomolgus mutatus Humes and Ho, 1966

Indomolgus mutatus Humes and Ho, 1966a, pp. 17-20, figs.

115-143 [from Palylhoa tuberculosa, Nosy Be, Madagascar].

Indomolgus panikkari (Gnanamuthu, 1955)

Lichomolgus panikkari Cnanamuthu, 1955, pp. 151-156, figs.

1-3 [from the actiniarian Phytocoeteopsis ramunni Panikkar, Madras, India].-Bouligand, 1966, p. 269.

Indomolgus panikkari.—Humes and Ho, 1966a, p. 11.

Genus Kelleria Gurney, 1927

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5—segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, in the

female with the armature 4, 13, 6, 3, 4 -f- 1 aesthete, 2+ 1 aesthete, and 7+ 1 aesthete; in the

male with 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete,

4 + 1 aesthete, 2+ 1 aesthete, and 7+ 1 aesthete. Second antenna 4-segmented, with the formula 1, 1, I + 2, and II + 5, the claws being

slender and setiform.

Labrum with a deep indentation separating the

two lobes. Mandible with the basal area distal to

the weak indentation, bearing on its convex side

a patch of spinules followed by a row of conspicuous spiniform teeth which grade into small spinules

on the lash and on its concave side a row of stout

spines; lash moderately long. Paragnath a small

lobe with hairlike spinules. First maxilla with four

elements. Second maxilla 2-segmented, with the

usual lichomolgid components but the lash short

and stout, forming an angle with the segment and

bearing irregularly sized spines; the element next

to the lash a strongly barbed spine. Maxilliped in

the female 3—segmented, with the proximal seta on

the second segment bearing one or two spinules

on its proximal margin, and with the small third

segment having four elements; in the male 4-seg-
186

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

merited (assuming that the proximal part of the

claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 1-segmented. Third segment

of leg 4 exopod 11,1,5. Leg 4 endopod with the

formula 11,1. Armature similar in both sexes except for leg 1 endopod in the male where the third

segment is 1,1,4 instead of 1,5 as in the female.

Leg 5 with a free segment bearing two terminal

elements. Leg 6 in the female represented by the

two setae and the spiniform process near the area

of attachment of each egg sac; in the male by a

posteroventral flap on the genital segment bearing

two setae.

Other features as in the species below.

Free-living, in intertidal burrows, or associated

with crinoids.

TYPE-SPECIKS.—Kelleria regalis Gurney.

REMARKS.—Lack of information on the males of

this genus prevents the construction of a key for

that sex.

Key to Species of the Genus Kelleria

FEMALES

1. Proximal spine on second segment of maxillipcd with a large lateral tpinule producing

a bifid appearance .... 2

This spine regularly pectinate or plumose, without such a ipinulc S

2. Body length not less than 1.25 mm, average 1.3 mm K. rcfallt

Body length 0.7 mm Jt. rubimmculmU

3. Medial margin of leg 5 with one or two pointed processes (

Medial margin of leg 5 with an obtuse swelling or straight 6

4. Medial margin of leg 5 with one pointed process 5

Medial margin of leg 5 with two pointed processes K. muttrmlUntU

5. Mediobasal tooth of mandibular blade much larger than the others K. propinqua

This tooth not oversized K. purpurodncta

6. Medial teeth of lash of second maxilla graded in size, with 5 or 6 teeth in all K. gradata

Medial teeth of lash of second maxilla irregular in size (long and short teeth mixed), with

7-10 teeth in all 7

7. Basal spine of second segment of maxilliped unilaterally pectinate 8

This spine bilaterally with barbules K. camortentU

8. Free segment of leg 5 about five times as long as wide K. andamanensis

Free segment of leg 5 about three times as long as wide K. pectinate

Kelleria regalis Gurney, 1927

FIGURE 105a-/

Kelleria regalis Gurney, 1927, pp. 471-474, figs. 116-117

[from Suez Canal].—Marques, 1961, p. 47, pis. lc-f, 2a

[from Portuguese Guinea].—Humes and Ho, 1969a, pp.

221-225, figs. 1-30 [from intertidal burrows, region of

Nosy Be1, Madagascar].—Anonymous, 1969, p. 414.

Kelleria andamanensis Sewell, 1949

Kelleria andamanensis Sewell, 1949, pp. 112-114, fig. 29a-i

[in surface tow-net, Andaman Islands].

REMARKS.—The male is unknown.

Kelleria australiensis Bayly, 1971

Kelleria australiensis Bayly, 1971, pp. 111-116, 2 figs [from

brackish water, Victoria, Australia].

Kelleria camortensis Sewell, 1949

Kelleria camortensis Sewell, 1949, pp. 114-117, fig. 30a-m

[in weed-washings, Nicobar Islands and Maldive Archipelago].

Kelleria gradata Stock, 1967

Kelleria gradata Stock, 1967b, pp. 569-573, figs. 1-3 [from

the crinoid Heterometra savignyi (J. Muller), Gulf of

Aqaba, Israel].

REMARKS.—The male is unknown.

Kelleria pectinate, (A. Scott, 1909)

FIGURE 105g

Pseudanthessius pectinatus A. Scott, 1909, pp. 268-269, pi.

68, figs. 21-27 [in plankton, 8°0.3' S, 116°59' E. Bali Sea].
NUMBER 127 187

Fir.i RF 105.—a-f, Kellcria regain Gurney. Female: a, dorsal; b, second antenna; c, mandible;

d, second maxilla; e, maxilliped. Male: /, dorsal. Length of female 1.32 mm, of male 1.05 mm.

g, Kelleria pectinata (A. Scott). Female: leg 4. (From Humes and Ho, 1969a, figs. 1, 8, 10,

IS. 15, 22. 43.)

Kelleria pectinata.—Humes and Ho, 1969a, pp. 225-228, figs.

31-53 [from intertidal burrows, including those of the

shrimp Axius (Neaxius) acanthus A. Milne Edwards,

region of Nosy Be, Madagascar].—Anonymous, 1969, p.

414. figs, a, b.

Kelleria rubimaculata Krishnaswamy, 1952

Kelleria rubimaculata Krishnaswamy, 1952, pp. 326-330, fig.

3 [in plankton, Madras, India]; 1953, pp. 66, 67 [in plankton, Madras, India].

Kelleria propmqua (T. Scott, 1894)

Pseudanthessius propinquus T. Scott, 1894a, p. 121, pi. 13,

figs. 49-56, pi. 14, figs. 1, 4 [in surface tow-net, Loanda,

Gulf of Guinea].

Kelleria purpurocincta Gurney, 1927

Kelleria purpurocincta Gurney, 1927, pp. 474-476, fig. 118

[from Suez Canal].

Genus Lichomolgella G. O. Sars, 1918

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. First antenna 7-segmented. Second antenna 4-segmented, with the

formula 1, 1, 3 -\- a slender claw, and 5 -j- a slender claw. Mandible and first maxilla unknown.

Second maxilla of the usual lichomolgid type. Maxilliped in the female 3-segmented with a pointed
188 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tip, in the male 4-segmented (the proximal part

of the claw representing the fourth segment).

Legs 1^1 in both sexes with 3-segmented rami

except leg 4 endopod which is a single segment

armed with two terminal spines. Leg 4 exopod

with the third segment having 11,1,5. Apparently

no sexual dimorphism in the formula of leg 1 endopod. Leg 5 with a free segment bearing two

terminal elements.

Other features as in the species below.

Free-living or associated with algae.

TYPE-SPECIES.—Lichomolgella pusilla G. O. Sars.

REMARKS.—The two species comprising this genus should be carefully restudied as existing descriptions are inadequate. The male of L. pusilla

is unknown.

Key to Species of the Genus Lichomolgella

FEMALES

Second segment of second antenna relatively short and stout, ratio about 2:1 L. fnuilla

Second segment of second antenna long and slender, ratio about 3.5:1 ... L. iueli

Lichomolgella pusilla G. O. Sars, 1918

FIGURE 106

Lichomolgella pusilla G. O. Sars, 1918, pp. 216, 217, pi. 118,

fig. 1 [in 20 fathoms, Skutesnaes, southwestern coast of

Norway]; 1921, pp. 111. 112. pi. 73. fig. 2 [from Skutesnaes.

Norway].-Gallingani, 1952, p. 72, fig. A.

REMARKS.—The male is unknown.

Lichomolgella isseli Gallingani, 1952

Lichomolgella isseli Gallingani, 1952, pp. 71-75, figs. 1-7

(p. 74) [from the alga Posidonia, Paraggi, Italy],

Genus Lichomolgides Gotto, 1954

DIAGNOSIS.—Body of the female modified, with

the prosome strongly inflated dorsally and with

FIGURE \06.-Lichomolgella pusilla G. O. Sars. Female: a, dorsal; b, second antenna; c, maxilliped; d, leg 4. (Redrawn from G. O. Sars. 1918, pi. 118, fig. 1.) Length of female 0.45 mm.
NUMBER 127 189

the cephalic area sharply delimited. Body of the

male cyclopiform. Urosome in the female 5-segmented, in the male probably 6-segmented.4

 Caudal ramus with six setae. Rostrum present. First

antenna 7-segmented. Second antenna 4-segmented, with the formula 1, 1, 2 —|— one claw, and

3 -f- four terminal claws.

Labrum with a shallow median cleft. Mandible

with a long lash. First maxilla with two elements.

Second maxilla of the usual lichomolgid form.

Maxilliped of the female 3-segmented with a

pointed tip; in the male 4-segmented (assuming

that the proximal part of the claw represents a

fourth segment). Postorally with a median, ventrally produced structure terminating in two small

hooks (this structure absent in the male).

Legs 1-4 with 3-segmented rami except leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

4

 Although in the text of the original description of the

male the number of postgcnital segments is given as three,

Gotto's (1954b) furore* 8 and 9 suggest the presence of four.

with the formula 0-1 ;II, the seta being feathered.

Leg 1 endopod with the same formula as in the

female. Leg 5 with a minute free segment bearing

a spine and a seta.

Other features as in the species below.

Lives in ascidians.

TYPE-SPECIES.—Lichomolgides cuanensis Gotto.

Lichomolgides cuanensis Gotto, 1954

FlCURF. 107

Lichomolgides cuanensis Gotto, 1954b, pp. 379-385, figs. 1-20

[in the cloacal cavities of the ascidian Trididemnum tenerum (Verrill), Strangford Lough, Northern Ireland];

1960b, p. 223, fig. 28; 1966, p. 194.

Genus Lichomolgus Thorell, 1859

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. Rostrum rounded though

sometimes elongated. First antenna 7-segmented,

in the female with the armature 4, 13, 6, 3, 4 -f- 1

FIGURE 107.—Lichomolgides cuanensis Gotto. Female: a, dorsal; b, lateral; c, second antenna;

d, mandible and first maxilla; e, distal portion of postoral hooked organ; /, leg 4. Male: g,

dorsal. (From Gotto, 1954b, figs. 1, 2, 4, 5, 8, 12, 16.). Length of female 1.5-2.1 mm, of

male 0.74 mm.
2QQ SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

aesthete, 2 + 1 aesthete, and 7 + 1 aesthete; in the third and fourth segments is obscure); forthe male with 4, 13 + 2 aesthetes, 6, 3 + 1 mula 1, 1, 3, and 1-4 terminal claws + several

aesthete, 4 + 1 aesthete, 2 -f 1 aesthete, and 7 setae (last segment in L. minor with five slender

-f- 1 aesthete. Second antenna 4-segmented setae, no claws),

(though in L. chamarum the articulation between Labrum deeply incised medially. Mandible with

Key to Species of the Genus Lichomolgus

FEMALES

1. Second antenna terminally with five slender setae I~ minor

Second antenna terminally with 1-4 claws 2

2. Second antenna terminally with one claw 3

Second antenna terminally with more than one claw 8

3. Rostrum elongated, moderately slender, and beaklike, extending between the bases of the

second antenna L- chamarum

Rostrum broadly rounded, not extending between the bases of the second antennae ..... 4

4. Caudal ramus with length to width ratio 4:1 or less 5

Caudal ramus with length to width ratio 5:1 or greater 7

5. Genital segment in dorsal view widest posterior to middle; free segment of leg 5 elongated

with ratio 2.73:1 L. tridocno*

Genital segment in dorsal view widest anterior to middle; free segment of leg 5 short,

ratio less than 2:1 6

6. Leg 5 minute, without an adjacent dorsal lobe; segment bearing leg 5 with a pair of

small (13 n long) posterodorsal processes L. mrtmmu

Leg 5 minute, with an adjacent dorsal lobe; segment bearing leg 5 without posterodorsal

processes L. tpondyli

7. First antenna with smooth setae; segment bearing leg 5 with a pair of small posterodorsal

spinous processes (13 fi long); caudal ramus with ratio 5.7:1 L. otophidit

First antenna with many setae haired; segment bearing leg 5 apparently without posterodorsal processes; caudal ramus with ratio a little more than 5:1 L. inflatus

8. Second antenna terminally with two claws 9

Second antenna terminally with more than two claws 17

9. Anal segment elongated, as long as or longer than the first two postgenital segments

combined 10

Anal segment short, much shorter than the first two postgenital segments combined II

10. Caudal ramus very long, tapered, ratio about 12:1, somewhat bent in the middle where it

carries two setae L. forficukt

Caudal ramus very long, less tapered, about 10:1, straight, carrying one seta near the

middle L. mmginatm

11. Caudal ramus wider than long L. imitcut

Caudal ramus longer than wide 12

12. Caudal ramus long, ratio about 7:1 L. ieversi

Caudal ramus shorter, ratio 6:1 or less IS

13. Longest seta on caudal ramus much longer than the ramus L. longieauda

Longest seta on caudal ramus shorter or only as long as the ramus 14

14. Caudal ramus with ratio 5:1-6:1 15

Caudal ramus with ratio less than 45:1 16

15. Longest seta on caudal ramus less than one-half as long as the ramus; ramus about 6:1

L. atbens

Longest seta on caudal ramus about as long as the caudal ramus; ramus about 5:1

L. diaumat

16. Free segment of leg 5 about 2.4:1, with a notch in the middle of the outer edge

L. furdUatus

Free segment of leg 5 about 1.8:1, with the inner edge forming a conspicuous bulge

L. canui

17. Second antenna terminally with three claws L. leptodermatus

Second antenna terminally with four claws L. elegantuhu
NUMBER 187 191

a usually slender base, merging gradually into a

long, slender, attenuated pectinate lash. Paragnath a small lobe with hairs or spinules. First

maxilla with two or three elements. Second maxilla of the usual lichomolgid form. Maxilliped in

the female 3-segmented; in the male 4-segmented

(assuming that the proximal part of the claw rep

resents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Armature of the

typical lichomolgidiform type. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1 ;II, the seta being feathered.

Leg 1 endopod of the male with the third segment

having the same formula as in the female (as far

as known). Leg 5 with a free segment bearing two

terminal elements and usually unornamented. Leg

6 represented in both sexes by the two setae near

the genital openings.

Other features as in the species below.

Associated mainly with ascidians and mollusks.

TYPE-SPECIES.—Lichomolgus albens Thorell.

REMARKS.—The seventeen species of Lichomolgus show considerable variation in the number of

claws (1-4) on the second antenna, yet otherwise

they form a cohesive group, especially in the form

of the mandible. One might suppose that generic

subdivisions could be made on the number of claws

combined with host preferences. Most of the species associated with bivalve mollusks, for example,

have only one claw (L. arcanus, L. asaphidis, L.

chamarum, L. inflatus, and L. spondyli). Two

other species from bivalves, however, have more

than one claw (L. leptodermatus with three claws

and L. elegantulus with four). Thus, a division on

this basis does not seem natural.

All the species associated with ascidians have two

claws on the second antenna (L. albens, L. canui,

L. diazonae, L. forficula, L. furcillatus, and L. marginatus). The single remaining species (L. longicauda) with two claws is associated with cephalopod mollusks.

Since subdivisions of the genus on the number

of second antennal claws and host preference appear to be artificial at present, we have broadly

interpreted the genus to embrace all the species

listed below.

Mariscal (1967) found several undetermined species of copepods of the genus Lichomolgus associated with sea anemones in the Indo-Pacific. Since

he used Lichomolgus in its older, broad sense, however, these copepods may in actuality belong to

other lichomolgid genera. This would seem probable in view of the species of Lichomolgus in its

restricted sense being found associated with ascidians or mollusks, rather than with actiniarians.

A key for the males of Lichomolgus has not been

prepared because sufficient information is not available from the existing descriptions and figures.

Lichomolgus albens Thorell, 1859

Lichomolgus albens Thorell, 1859, pp. 340, 341; 1860, pp..

69-71, pis. 10, 11, 13 [from the ascidians Ascidia parallelogramma, Ascidia mentula, and Ascidia canina, BohuslSn,

Sweden].—Mobius, 1874, p. 275 [from the ascidian Molgula,

Zuiderzee, The Netherlands; this may represent L. canui,

below].-Aurivillius, 1882b, pp. 98-100, pi. 16, figs. 9, 10

[from the ascidians Phallusia patula O. F. Muller, Phallusia xrirginea O. F. Muller, and Corella parallelogramtna

Alder. Gullmarsfjord, Sweden]; 1883, pp. 94-%, pi. 7, figs. 9,

10 [from Phallusia patula, Phallusia virginea, and Corella

parallelogramtna, BohuslMn, Sweden].—Carus, 1885, pp. 350,

351.-Canu, 1891a, p. 478 [from Ciona intestinalis O. F. Muller, Molgula socialis Alder, and Cynthia lurida Thorell,

northern France]; 1892. pp. 230, 231, pi. 22, figs. 15, 23, pi.

23, figs. 1-4 [from Ciona intestinalis, Molgula socialis, Cynthia lurida, and Perophora listeri M. Don., Boulogne coast

of FranceJ.-Thompson, 1893. p. 207, pi. 15, fig. 3c [from

Puffin Island, Liverpool Bay, England]; 1889, pp. 182, 189

[in tow-net. Puffin Island, Liverpool Bay, England].—T.

Jfcott, 1894b, p. 258; 1898, p. 269 [from Otter Spit, Clyde,

Scotland]; 1907, p. 371.—Farran, 1913, p. 6 [in Ascidiella

aspersa].-G. O. Sars, 1917a, pp. 151-153, pi. 83 [from the

branchial cavity of various ascidians, most frequently

Corella parallelogramma, Norway].—Lang, 1949, p. 8 [from

the ascidian Corella parallelogramma, Sorgrundsberget,

Sweden].-Gotto, 1960b, pp. 213, 222.-Bresciani and Lutzen, 1962, p. 377.

REMARKS.—This species has been designated the

type-species following G. O. Sars (1917, p. 153).

Lichomolgus arcanus Humes and Cressey, 1958

Lichomolgus arcanus Humes and Cressey, 1958a, pp. 938-

941, figs. 127-156 [from the mantle cavity of the bivalves

Area senilis Linnaeus, Ostrea tulipa Lamarck, and Pitar

tumens Gmelin, Senegal].-Cheng, 1967, pp. 315, 343.

Lichomolgus asaphidis Humes, 1959

Lichomolgus asaphidis Humes, 1959, pp. 309-315, figs. 125-

159 [from the bivalves Asaphis rugosa Lamarck and Standelta solanderi Gray, Nosy B£, Madagascar].
192

NEW HOST.—3 9 9,1c ? from 30 bivalves, Solen

conneus Lamarck, intertidal, west of Pte. Mahatsinjo, Nosy B6, Madagascar, 4 October 1960, collected by AGH.

SMITHSONIAN CONTRIBUTIONS TO /OOLOGY

Lichomolgus diazonae Gotto, 1961

Lichomolgus diazonae Gotto, 1961b, pp. 85-88, fig. la-r [from

the ascidian Diazona violacea Savigny. Plymouth, England];

1962, p. 104.

Lichomolgus canui G. O. Sars, 1917

Lichomolgus canui G. O. Sars, 1917a, pp. 157-158, pi. 87

[from ascidians, Norway].—Klie, 1933, p. 15 [in the ascidian

Styelopsis (Dendrodoa) grossularia, Kiel, Germany].—

Stock, 1960b, pp. 72, 73 [from the ascidians Molgula manhattensis (de Kay), Molgula sp., Ascidiella aspersa (Miiller), and Ciona intestinalis (Linnaeus), The Netherlands].-Gotto, 1960b, p. 227; 1961b, p. 88 [from the

ascidian Clavelina lepadiformis (O. F. Miiller), northern

Ireland]; 1966, p. 194.-Leloup et al., 1963, p. 29; 1964, p.

30.-Leloup and Polk, 1967, pp. 17, 86.-Costanzo, 1968, pp.

27-37, figs. 1-6 [from the ascidians Clavelina lepadiformis

and Styela plicata, Lago di Faro, Messina, Sicily]; 1969,

pp. 143-158.

Lichomolgus albens.—Mobius, non Thorell, 1873, p. 275

[from Zuiderzee, The Netherlands; probably = L. canui,

cf. Stock, 1960b].

Lichomolgus chamarum

FIGURE 108

Lichomolgus chamarum Humes, 1968b, pp. 74-81, figs. 37-64

[from the bivalve Chama iostoma Conrad, region of Nosy

Be, Madagascar].

Lichomolgus eUgantulus Stock, 1960

Lichomolgus elegantulus Stock, 1960c, pp. 238-241, figs. 11,

12 [from the bivalve Pteria hirundo (Linnaeus), Banyuls,

Mediterranean coast of France].

Lichomolgus forficula Thorell, 1859

Lichomolgus forficula Thorell, 1859, p. 340; 1860, pp. 73. 74,

pis. 12, 13, fig. 19 [from the ascidian Ascidia canina,

Bohuslan. Sweden].-Claus, 1875, p. 347.-Kossmann. 1877.

pp. 18, 19, pi. 4, fig. 2 [from various ascidians, Naples,

Italy].-Brady, 1880. pp. 50-52. pi. 85. figs. 12-16, pi. M.

figs. 14-18, Scotland and Irdand].-Aurivillius. 1882b, pp.

100. 102. pi. 16. figs. II, 12 [from Phallusia mentula, Bohuslan, Sweden]; 1883. pp. 96-98, pi. 7. figs. II, 12 [from

Phallusia mentula, Hohiislan. Swcdcn].-Gicbrccht, 1882,

pp. 295. 296 [from the ascidians Phallusia mentula O. F.

Miiller and Phallusia mamillata Cuvier, Naples, Italy].—

Carus. 1885. p. S50.-Gourret, 1887. p. 187; 1889, p. 474

(/.. elongatus) .-T. Scott, 1894b, p. 258; 1897a. p. 153 [from

the branchial chambers of large ascidians, Loch Tyne and

East Loch, Tarbert. Scotland]; 1907, p. 370 [from large

ascidians, Scotland].-Herdman. 1896. p. 48.-Graffe, 1900,

p. 41 [from ascidians, Trieste, Italy].—Pearson, 1905, p. 163

[from simple ascidians, Ascidia sp., Ireland].—Norman and

f

FIGURE 108.—Lichomolgus chamarum Humes. Female: a, dorsal; b, second antenna; c, mandible;

d, leg 4; e, leg 5. Male: /, dorsal. (From Humes, 1968b, figs. 37, 43. 46, 55, 57, 58.) Length of

female 1.66 mm, of male 1.34 mm.
NUMBER 127 193

T. Scott. 1906, p. 198 [from the branchial chambers of

Phallusia mamillata, Plymouth, England].—Farran, 1913,

p. 6 [in Ascidia men tula, Ireland].—G. O. Sars, 1917a, pp.

153, 154, pi. 84 [from the ascidians Ascidia mentula and

Ascidia canina, Norway].—Bocquct, 1952, p. 497 [from the

echinoid Psammechinus miliaris, Beclem, Roscoff, northern

France].—Fouque and Franc, 1953, p. 24 [from Ascidia

aspersa Miiller and Ascidia mentula Miiller, northern

France].—Marine Biological Association, 1957, p. 175.—

Gotto. 1960b, pp. 213, 222, fig. 29; 1966, p. 193.-Bresciani

and Liitzen, 1962, p. 3"7.-Bruce, Colman, and Jones, 1963,

p. 127.

Lichomolgus elongatus Buchholz, 1869, pp. 151-153, pi. 11,

fig. 9 [from the ascidian Phallusia mamillata and several

others, Naples, Italy; probably=/.. forficula, cf. Kossmann,

1877, p. 18, Delia Valle, 1880a, p. 84, and Giesbrecht, 1882.

p. 295].

Lichomolgus forficulus.—D'Arcy Thompson, 1901, p. 47.

Lichomolgus furcillatus Thorell, 1859

Lichomolgus furcillatus Thorell, 1859, p. 341; 1860, pp. 74,

75, pi. 13, fig. 20 [from the ascidian Ascidia intestinali (sic),

BohusUn, Sweden].—Brady and Robertson, 1873, p. 140

[from Roundstone Bay, western Ireland].—var. mediterranea

Kossmann, 1877, p. 19, pi. 4, fig. 1 [from Mediterranean

Sea].-Brady, 1880. pp. 49-50, pi. 88. figs. 10-14 [from the

branchial sac of the ascidian Corella paratlelogramma,

England).-Aurivillius, 1882b, pp. 102-105 [from Ciona intestinalis, Gullmarafjord, BohuslSn, Sweden]; 1883, pp.

98-101 [from Ciona intestinalis, Gullmarsfjorden, Bohuslan,

Sweden].—T. Scott, 1891, p. 304 [from the branchial cavity

of an ascidian, west of May Island, Firth of Forth, Scotland]; 1894b, pp. 233. 258; 1897a, p. 153 [off Inveraray,

Upper Loch Fyne, Scotland]; 1902, p. 470 [from Inchkeith,

Scotland]; 1906, p. 352 [from the branchial cavity of large

ascidians, May Island and Inchkeith, Firth of Forth, Scotland]; 1907, p. 371.-Thompson, 1893, p. 207, pi. 25, fig. 3

Liverpool Bay, England].—Herdman, 1896, p. 66.—Canu,

1898, p. 415.—Pearson, 1905, p. 162 [in tow-net, Ireland].—

G. O. Sars, 1917a, pp. 156, 157, pi. 86 [from the branchial

cavity of the ascidian Stye la intestinalis, Norway].—Marine

Biological Association, 1957, p. 175.—Gotto, 1960b, pp.

213, 223; 1966. p. 194.-Bresciani and Liitzen, 1962. p. 377.

Pseudanthessius furcillatus.-Gorman and T. Scott, 1906, p.

198 [from England].-Farran, 1913, p. 6 [from Ireland].

Lichomolgus ieversi Thompson and A. Scott, 1903

Lichomolgus ieversi Thompson and A. Scott, 1903, p. 278,

pi. 15, figs. 10-17 [from washings of invertebrates including

pearl oysters, Ceylon].

Lichomolgus indicus Ummerkutty, 1962

Lichomolgus indicus Ummerkutty, 1962, pp. 62-66, pi. 12,

figs 4-10, pi. 13, figs. 1-4 [from starfish and sponge washings. Gulf of Mannar, southeastern India].

REMARKS.—According to its describer, this species has the very unusual formula of 11,1,4 on the

third segment of leg 4 exopod, with only four

setae instead of five as in other lichomolgids.

Lichomolgus inflatus Tanaka, 1961

Lichomolgus inflatus Tanaka, 1961, pp. 263-266, pi. 30, figs.

1-8, pi. 31, figs. 1-10, pi. 32, figs. 1-5 [from the bivalve

Paphia, Sasebo Bay, Japan].—K6, Murakami, and Daiku,

1962, p. 114 [from the bivalves Tapes japonica, Saxidomus

purpuratus (Sowerby) and Pinctada martensii (Dunker),

Japan.-Kd, 1969b, pp. 83-86, 90, 91 [from Tapes japonica,

Sasebo Bay, Japan].

Lichomolgus leptodermatus Gooding, 1957

Lichomolgus leptodermatus Gooding, 1957, pp. 207-212, figs.

4, 5a-m, [from the gills of the bivalve Laevicardium erassum (Gmelin), Plymouth. England].—Marine Biological

Association, 1957, p. 176.—Bocquet and Stock, 1959c, pp.

119, 120 [from Cardium crassum Gmelin (=Laevicardium

crassum (Gmelin)=Cardium norvegicum Spengler), Channel

coast of France, and Plymouth, England].

Lichomolgus longicauda (Claus, 1860)

Sepicola longicauda Claus, 1860, pp. 4, 5, pi. 1, figs. 7-9

[from the gills of the cephalopod Sepia officinalis, Nice

(Nizza), France].

Lichomolgus sepicola.-Claus, 1875, p. 348.-Wiezejski, 1877,

pp. 574-580, pi. 34, figs. 18-32 [from Sepia officinalis,

Trieste, Italy].—Stossich, 1880, p. 250 [from the gills of

Sepia officinalis, Trieste, Italy].—Pesta, 1909, pp. 262, 263,

pi. 2, fig. 7 [from the gills of Sepia officinalis, Trieste, Italy].

Lichomolgus sepiicola.—Czrm, 1885, p. 351.—Graeffe, 1900,

p. 41 [from the gills of Sepia officinalis var. filliouxi Lafont,

Trieste, Italy].

Lichomolgus longicauda.-Cuenot, 1927, p. 287 [from Sepia

officinalis and Sepia filliouxi, Arcachon, France].—Stock,

1956, pp. 117-120, figs. 1-12 [from the gills and mantle

cavity of Sepia officinalis Linnaeus, The Netherlands];

1960c, p. 242 [from the gills of Sepia officinalis Linnaeus,

Banyuls, Mediterranean coast of France]; 1964a, p. 67.—

Gotto, 1962, p. 102.—Glacpn, 1971, p. 23 (unnumbered).

Lichomolgus (Lichomolgus) longicauda.—Monod and Dollfus,

1932a, p. 138 [on Sepia filliouxi Lafont].

Lichomolgus (Sepicola) longicauda.—Bresciani, 1970, p. 11.

Lichomolgus marginatus Thorell, 1859

Lichomolgus marginatus Thorell, 1859, p. 340; 1860, pp.

71-73, pi. 12, fig. 18 [from the ascidians Ascidia venosa

and Ascidia canina, Bohusla"n, Sweden].—G. O. Sars, 1917a,

pp. 155, 156, pi. 85 [from Norway].—Fouque and Franc,

1953, pp. 24, 25 [from Ascidia mentula Miiller, northern

France]-Boschma, 1956, p. 8.-Mulder, 1956, p. 11 [in
194 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

tunicate, The Netherlands].-Gotto, 1960b, p. 213, p. 223;

1966, p. 194.—Bresciani and Liitzen, 1962, p. 377.

Lichomolgus minor A. Scott, 1902

Lichomolgus minor A. Scott, 1902, pp. 417, 418, pi. 1, fig. 17,

pi. 2, figs. 15-24 [in plankton from bathroom tap on ship,

Red Sea and Indian Ocean].—Thompson and A. Scott,

1903, p. 277 [between Port Said and Suez, and in washings

of a young pearl oyster, Ceylon].—Gurney, 1927, p. 465

[from Suez Canal].

Lichomolgus spondyli Yamaguti, 1936

Lichomolgus spondyli Yamaguti, 1936, pp. 124, 125, pi. 13,

figs. 76-84 [from the mantle cavity of the bivalve Spondylus japonicus Kuroda, Pacific coast of Wakayama Prefecture, Japan].-Tanaka, 1961, p. 249.

Lichomolgus tridacnae Humes, 1971

Lichomolgus tridacnae Humes, 1971, pp. 348-357, figs. 1-S2

[from the mantle cavity of the bivalves Tridacna gigas

(Linnaeus) and Tridacna squamosa Lamarck, Eniwetok

Atoll, Marshall Islands].

Genus Macrochiron Brady, 1872

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. Rostrum with a slender,

needle-like process. First antenna 7-segmented, in

the female with the armature 4, 13, 6, 3, 4 + 1

aesthete, 2+ 1 aesthete, and 7 —f- 1 aesthete; in

the male 4, 13 + 2 aesthetes, 6, 3 + 1 aesthete,

4 + 1 aesthete, 2+ 1 aesthete, and 7 + 1 aesthete. Second antenna 3-segmented, the third segment resulting from the fusion of the original third

and fourth segments; formula 1,1, and 3 + terminally one pectinate claw and one clawlike seta, or

two pectinate claws and five setae.

Labrum incised medially. Mandible with the

basal region beyond the indentation having on its

convex side a scalelike area with spinules followed by a serrated fringe and on its concave side

a row of spinules; lash long. Paragnath a small

hairy lobe. First maxilla with four elements. Second maxilla of the usual lichomolgid type, bearing

near the base of the apical lash an ornamented

seta and a smooth, hyaline, often blunt seta. Maxilliped in the female 3-segmented, slender, the

first two segments elongated and held at a sharp

angle to each other, the last segment small and

indistinctly separated from the second segment; in

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 1- or 2-segmented. Leg 4 exopod with the third segment having 11,1,5 or 111,1,5.

Leg 4 endopod with the formula 0-0;2 or, if 1-segmented, simply 2. Leg 1 endopod in the male

1,1,4, so far as known, instead of 1,5 as in the female. Leg 5 with a free segment having two terminal setae; leg 6 represented by two setae near the

genital openings.

Other features as in the species below.

Associated mainly with algae or hydroids, in certain cases with ascidians or echinoids.

TYPE-SPECIES.—Macrochiron fucicolum Brady.

Key to Species of the Genus Macrochiron

FEMALES

1. Last segment of exopod of leg 5 with armature 11,13 2

Last segment of exopod of leg 5 with armature 111,1,5 6

2. Endopod of leg 4 a single segment (or at most with only a weak and incomplete indication

of division) 3

Endopod of leg 4 composed of two distinct segments M. fucicolum

3. Genital segment abruptly constricted in its posterior third; rostrum with a subconical beak

M. rostratum

Genital segment broadest anteriorly and tapering posteriorly; rostrum with a needle-like

process 4

4. With a pair of posterior protuberances on segment of leg 5 medial to fifth leg . M. vervoorti

Without such protuberances 5

5. Free segment of leg 5 tapered distally; endopod of leg 4 without an outer marginal notch

M. chetiferum

Free segment of leg 5 not tapered; endopod of leg 4 with an outer marginal notch

M. valgum
NUMBER 127 195

6. Endopod of leg 4 composed of two distinct segments M. mutatum

Endopod of leg 4 a single segment (although in Af. echinicolum and M. lytocarpi there

may be a very indistinct line of division) 7

7. Second antenna with one claw (though a jointed seta may be present also) 8

Second antenna with two claws 9

8. With a pair of conspicuous, well-sclerotized posterior lobes on segment of leg 5 medial to

fifth legs M. lobatum

Without such lobes M. lytocarpi

9. Claws on last segment of second antenna nearly equal to length of segment; maxilliped with

armature dentiform in appearance M. sargasfi

Claws on last segment of second antenna much shorter than segment; maxilliped with usual

setiform or spiniform armature M. echinicolum

MALES

1. Last segment of exopod of leg 4 with armature 11,13 2

Last segment of exopod of leg 4 with armature 111,13 6

2. Endopod of leg 4 a single segment (or at most with only a weak and incomplete indication

of division) . 3

Endopod of leg 4 composed of two distinct segments M. fucicolum

3. Rostrum with a subconical beak M. rottratum

Rostrum with a needle-like process 4

4. Without sexual dimorphism in formula of endopod of leg 1 (Vervoort, 1964) . M. cheliferum

With sexual dimorphism in this ramus (last segment, 1,1,4) 5

5. Caudal ramus 54 x 23 ft (2.35:1); a single claw on second antenna M. valgum

Caudal ramus 36 x 20 p (1.8:1); two claws on second antenna M. verooortx

6. Endopod of leg 4 composed of two distinct segments M. mutatum

Endopod of leg 4 a single segment 7

7. Second antenna with one claw (though a jointed seta may be present also) 8

Second antenna with two claws 9

8. Outermost spine on last segment of endopod of leg 1 with proximal half swollen medially

and abruptly delimited from distal portion M. lobatum

Outermost spine on last segment of endopod of leg 1 swollen medially along most of its

length M. lytocarpi

9. Claws on second antenna nearly equal to length of last segment M. sargassi

Claws on second antenna distinctly shorter than last segment M. echinicolum

Macrochiron fucicolum Brady, 1872

Macrochiron fucicolum Brady, 1872a, pp. 9, 10, pi. S, figs.

9-18 [among Fucus, between Ryhope and Sunderland,

England; spelled "Macrocheiron" on plate].—Sewell, 1949,

pp. 99, 152, 156, 185-187.-Stock. 1957. pp. 380, 381, figs.

8-10 [from The Netherlands]-Stock and de Vos, 1960,

p. 206 [among Fucus and edible mussels. The Netherlands].

Macrochiron fucicolus.—Sewell, 1949, pp. 94, 95.

Macrochiron (Macrochiron) fucicolum.—Shen and Lee, 1966,

p. 223.

Macrocheiron fucicolum.-Brady, 1872a, pi. 3, figs. 9-18;

1872b, p. 434, pi. 18, figs. 9-18 [among Fucus, between

Ryhope and Sunderland, England].—G. O. Sars, 1917a, pp.

163-165, pi. 91 [from Norway].-C. B. Wilson, 1932, pp. 349,

350, fig. 209 [from Katama Bay, Martha's Vineyard,

Massachusetts].—Gallingani, 1952, p. 75.—Marine Biological

Association, 1957, p. 176.—Bruce, Colman, and Jones, 1963,

p. 128.-Vervoort, 1964. p. 47.

Macrocheiron fuciculum.—Brian, 1923, p. 131 [from Rovigno,

Italy].—Vatova, 1928, p. 179 [on Sargassum, Bagnole. near

Rovigno, Italy].

Lichomolgus fucicolus.-Brady and Robertson, 1873, p. 140

[among weeds, western Ireland; England]; 1876, p. 197

[from England] .-Brady, 1880, pp. 41-i4, pi. 85, figs. 1-11

[from England]; 1902, p. 55 [off Newcastle, County Down;

Roundstone Bay; Portpatrick Harbour, Wigtownshire].—

Thompson, 1889, p. 189 [among algae, Puffin Island, Liverpool Bay, England]; 1893, p. 207, pi. 25, fig. 1 [from

Liverpool Bay, England]; 1897b, p. 98 [from St. Aubin's

Bay, English Channel].-T. Scott, 1894b, pp. 233, 258;

1897a, p. 153 [from Upper Loch Fyne, Scotland]; 1901,

p. 252 [from Sound of Mull. Scotland]; 1906, p. 352 [from

Firth of Forth, Scotland].—Herdman, 1896, p. 66.—Timm,

1896, p. 384 [from Cuxhaven, Germany].-Pearson, 1905,

pp. 162, 163 [among weeds and in tow-net, Ireland].—

Conseil Permanent International pour l'Exploration de

la Mer, 1906, p. 95 [from English Channel].-L. W. Williams, 1907, p. 75 [from Wickford and Charlestown Pond,

Rhode Island].-Sharpe, 1910. pp. 408, 421, 422, fig. 16 [in

surface net, Buzzards Bay, Woods Hole, Massachusetts].-

Farran, 1913, p. 6 [from Ireland].—Sumner, Osburn, and

Cole, 1913, p. 639 [from Buzzards Bay, Massachusetts].
1% SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lichomolgus fucicola.—Norman and T. Scott, 1906, p. 198

[from England].-T. Scott, 1912, p. 571 [from floating

seaweed, 33°53' N, 32°27' W].

Mcurochiron cheliferum (Thompson and A. Scott,

1903)

Pseudanthessius chelifer Thompson and A. Scott, 1903, p.

277. pi. 14, figs. 12-18 [from Ceylon].

Macrocheiron chelifer.—Gurney, 1927, p. 469 [from Suez

Canal].

Macrocheiron cheliferum.—Vervoort, 1964, pp. 47-54, figs.

lla, 12-14 [from Ifaluk Atoll, Caroline Islands].

Macrochiron cheliferum.—Stock, 1957, p. 381.

Macrochiron (Paramacrochiron) cheliferum.—-Sewell, 1949, p.

108.

Lichomolgus (Macrochiron) chelifer.—Monod and Dollfus,

1932a, p. 139.

Macrochiron echinicolum, new species

FIGURES 109, 110

MATERIAL EXAMINED.—From Lytechinus variegatus (Leske), the green-white sea urchin (determined by Dr. W. Schroevers, Amsterdam): Bonaire

(collected by JHS): many specimens (holotype, allotype, and paratypes), sea-grass field, depth 1-2

m, Lac, near Cai, 23 December 1958, ZMA Co.

100.647a-c. Curasao (collected by JHS): 16 specimens, partly ovigerous, dredged in about 3 m in

Piscadera Innerbay, near the former turtle hatchery, 17 December 1958, ZMA Co.100.648; many

specimens, on sandy bottom, depth about 3 m,

Fuikbay, 10 December 1958, ZMA Co. 100.649.

Puerto Rico (collected by AGH and RUG): 7 ? ? ,

4 cT cT from 63 hosts, Cayo Corral, south of La

Parguera, 30 July 1959; 107 $ $, 150 cf tf, and

43 copepodids from 150 hosts, depth 1 m, Cayo

Majimo, near La Parguera, 2 August 1959. Jamaica

(collected by AGH and RUG): many specimens,

depth 1.5 m, Rackham's Cay, near Kingston, 31

August 1959.

From Echinometra viridis A. Agassiz, the dark

sea urchin (determined by Prof. H. Engel, Amsterdam): Curagao (collected by JHS): 1 cf, depth

about 1 m, Fuikbay, 10 December 1958, ZMA Co.

100.650a-b.

From Tripneustes ven.tricosus (Lamarck), the

white sea urchin (determined by Dr. W. Schroevers,

Amsterdam): Curacao (collected by JHS): 1 cf

from 20 hosts, depth about 1 m, Piscadera Bay, 27

September 1958, ZMA Co. 100.651. Bahamas (this

and the following eight collections made by AGH

and RUG): 1 $ from single host, off the Lerner

Marine Laboratory, North Bimini, 1 June 1959.

Barbados: 1 o*» 3 copepodids from 9 hosts, Six

Men's Bay, 18 June 1959; 24 ? 9, 13 cf cf from 3

hosts, depth 8 m, near wreck in Carlisle Bay, 6

July 1959; 116 $ $, 46 cf <f from 11 hosts, depth

5 m, Carlisle Bay, 16 July 1959; 5 9 9, 1 cf from

a single host, depth 6 m, Oistin's Bay, 15 July

1959; 20 $ ?, 20 cf cf. and 1 copepodid from 6

hosts (white variety), depth 1.5 m, Carlisle Bay,

20 July 1959; and 133 9 9 , 91 cf cf. and 21 copepodids from 5 hosts (black and white variety), depth

1.5 m, in front of Bellairs Research Institute, St.

James, 17 July 1959. Puerto Rico: 255 99 , 618

cf cf, and 20 copepodids from 25 hosts (plus many

copepods unsorted from debris), Cayo Corral, south

of La Parguera, 30 July 1959. Jamaica: 274 99 ,

420 cfcf. and 331 copepodids (1,025 in all) from

7 hosts, depth about 1.5 m, Rackham's Cay, near

Kingston, 31 August 1959.

The collections were made by AGH and RUG

during field work supported by the National Science Foundation (G-8628), and by JHS during a

stay at the Caribbean Marine Biological Institute,

Piscadera Bay, Curasao, subventioned by the former WOSUNA (now Netherlands Organization

for the Advancement of Scientific Research in the

Tropics, The Hague).

FEMALE.—The total length, without the furcal

setae, is 1.49-1.41 mm (mean, 1.32 mm), based on

six specimens. The metasome (Figure 109a) is

elongately oval in outline, not very enlarged. The

first pedigerous segment is completely fused with

the cephalic segment. The ratio of the length to

the width of the prosome is 1.57:1. The ratio of

the length of the prosome to that of the urosome

is 1.31:1.

The urosome consists of the usual five segments.

The genital segment is 230 x 187 n. The caudal

ramus (Figure 1096) is 113 x 35 n, about three

times as long as wide, and more than 1.5 times as

long as the anal segment. Each ramus bears five

long, feathered setae—one at about % of the outer

margin, one lateral subterminal, two terminal, and

one internal subterminal. A short, smooth dorsal

seta is borne near the base of the latter. A row of

minute spinules occurs on the posterior border of

the anal segment on both sides.
NUMBER 127 197

The egg sac (Figure 109fl) is short, not reaching

the extremity of the caudal rami and in some

specimens hardly reaching the second postgenital

segment, is oval in shape, and contains numerous

small eggs about 46 n in diameter.

The first antenna (Figure 109rf) is 343 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 22 (64 n

along the anterior margin), 88, 24, 61, 46, 32, and

28 ji respectively. The formula for the armature

is the same as in Macrochiron lytocarpi Humes,

with each of the three terminal segments bearing

an aesthete. The rostrum (Figure 109c) is a

pointed, tongue-shaped lobe. The second antenna

(Figure 109<*) is 3-segmented. The stout basal

segment bears a spiniform seta which is finely pectinate on the inner margin. A similar seta occurs

at three-f.)urths of the inner margin of the second

segment. The third segment is slender, being nearly

three times as long as wide, and bears (1) on the

proximal inner margin two setae (one smooth and

short, the other finely plumose and long) and (2)

before the middle a pectinate seta. There is a complex terminal armature consisting of three smooth

setae, two setae whose margins are provided with

widely spaced, short ciliations, and two claws with

finely and uniformly denticulated inner margins.

The longer of these claws (58 \*. along its axis) is

strongly curved, the shorter (47 v) is indistinctly

2-segmented.

The labrum is similar to that in M. lytocarpi.

The mandible (Figure 109/) is of the rather complex structure typical for Macrochiron. The paragnath is a small hairy lobe as in other species of

the genus. The first maxilla (Figure 109/) consists

of a simple lobe armed with one lateral smooth

seta and three terminal compound setae. The second maxilla (Figure 109g) has a large basal segment; the terminal main lash bears more than 20

denticulations on its medial margin; the auxiliary

lash also bears nearly 20 slender denticulations; a

crenulated spinelike element arises near the implantation of the auxiliary lash.

The maxilliped (Figure \09h) is 3—segmented,

the first segment being rather short and obscure.

The second segment is smooth and rectangular.

The third segment has an irregular shape and bears

(1) a strong denticulated spine on its medial margin and (2) a terminal projection armed with two

rows of denticulations and with two spinules at

its base.

The ventral area between the maxillipeds and

the first pair of legs resembles that of M. lytocarpi.

Legs 1-4 are biramous, each ramus being trimerous except the endopod of leg 4 which is a

single segment. Legs 1 and 2 are as figured (Figures 109», 110a). Leg 3 resembles leg 2 but has

the armature of the third endopod segment slightly

different (Figure 1106). Leg 4 (Figure 110c) has

a 3-segmented exopod 213 n in length. The endopod, 98 x 31 n in greatest dimensions including

the spinous processes, is functionally 1-segmented

(Figure 110c), although an indication of a segmentation line may sometimes exist (Figure HOd).

The place of the former articulation is indicated

by a distinct indentation of the outer margin of

the segment. The only armature of the fourth

endopod consists of two spines, the outer 52 n,

the inner 75 n. The inner side of the basipods of

legs 2-4 bears a strong bicuspidate process. On

leg 1 a similar process is usually lacking, but in

one specimen it was observed, though only on one

side and thus probably representing an anomaly.

The chaetotaxis of the rami of the legs is shown

by the following formula:

P, exp I-O; 1-1; 111,1,4

enp 0-1; 0-1; 1,5

Ps exp 1-0; 1-1; III.U

enp 0-1; 02; 1,113

P, exp 1-0; 1-1; 111,13

enp 0-1; 0-2; 1,11.2

P4 exp 1-0; 1-1; 111,1,5

enp II

Legs 1-4 have well-developed intercoxal plates,

each with a medial coxopod seta (plumose, reduced

in leg 4 to only 18 n in length) and a lateral

basipod seta (plumose).

The 1-segmented fifth leg, 114 x 26 n in greatest

dimensions, is clavate, long, slender, and curved

(Figure HOe). The distal part of its lateral margin

bears short spinules; there are two distal spines of

very unequal length (39 n and 83 n). The adjacent

seta on the body is 40 n and lightly feathered.

Leg 6 is represented by two small, naked setae

10 n and 8.5 n and a small spinous process near

the area of attachment of each egg sac.

MALE.—The total length, without the furcal

setae, is 1.00-1.18 mm (mean, 1.09 mm), based on

six specimens. In general aspect (Figure 110/) the
FIGURE lQ&.—Macrochiron echinicolum, new species. Female: a, dorsal (A) ; b, anal segment

and caudal rami (B) ; c, rostrum (c); d, first antenna, with asterisks indicating positions of

additional aesthetes in male, (D); e, second antenna (F.) ; /, mandible and first maxilla (F) ; g,

second maxilla (c); h, maxilliped (E) ; i, leg 1 (H) ; /, intercoxal plate of leg 1 (i). Scale:

A, 0.25 mm; B, C, 0.1 mm; D-F, U, I, 0.05 mm; c, 0.025 mm. (Drawn from sample ZMA

Co.100.647.)
NUMBER 127 199

FIGURE 110.—Macroch iron echinicolum, new species. Female: a, leg 2 (j); b, third segment

of endopod of leg 3 (E) ; c, leg 4 and intercoxal plate (H) ; d, endopod of leg 4 of another

female (H) ; f, kg 5 (E) . Male: /, dorsal (A) ; g, distal segments of second antenna (K) ; h,

maxilliped (E) ; i, third segment of endopod of leg 1 (K) ; /, leg 5 (F) ; *, leg 6 (H) . Scale:

A, 035 mm; E, F, H, K, 0.05 mm; j , 0.1 mm. (Drawn from sample ZMA Co.100.647.)
200 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

body is more slender than in the female. The

ratio of the length to the width of the prosome is

1.57:1. The ratio of the length of the prosome to

that of the urosome is 1.11:1. The urosome is 6-

segmented. The genital segment, 253 x 215 n, is

striking. Left and right on the posterior margin

of this segment we find a pointed prominence (Figure 110*) armed with two setae, 29 n and 31 n,

presumably representing the rudimentary sixth legs.

The caudal ramus is 89 x 30 u.

There are several secondary sexual differences in

the structure of the appendages. The first antenna

bears, in addition to the three aesthetes present in

the female, two long aesthetes on the second segment and one such aesthete on the fourth segment

(the position of these aesthetes being indicated by

an asterisk in Figure \09d). The second antenna

has a more slender terminal main claw (78 \i along

its axis), which is moreover less curved than the

corresponding claw in the female (compare Figures

109e and HOg). The male maxilliped (Figure

110//) is strongly prehensile; it consists of a smooth

basal segment, a large second segment, a very small

third segment, and a long, curved claw 198 n along

its axis. The palmar edge of the second segment

is armed with a row of sharp needlelike spines,

with a strong compound spine, and with a slender

smooth spine. Two setae arise near the base of

the claw. The cutting edge of the claw is smooth,

not crenulated.

The first leg of the male is similar to that of

the female except that the outer distal seta of the

third endopod segment (Figure HOi) has been replaced by a spine in the male. Consequently, the

chaetotaxis formula of the endopod of leg 1 of the

male is 0—1;0—1;II,4. The second and third legs

are similar to those of the other sex. The fourth

leg differs only slightly from that of the female—

in having the third exopod segment a trifle more

elongate.

The fifth leg (Figure 110;), 64 x 13 n, is less

slender and straighter than that of the female, with

its two terminal elements 30 n and 57 n; its outer

margin is smooth.

Leg 6 is shown in Figure 110A.

The spermatophore is unknown.

COLORATION.—Specimens associated with Lytechinus and Echinometra have an opaque, grayish

or ivory prosome; the egg sacs have the same color;

the eye is pale red; the urosome is colorless. The

specimens found on Tripneustes have a dark red

eye, an opaque, colorless body, and a somewhat

yellowish intestine.

ETYMOLOGY.—The specific name echinicolum

(Latin echinus, a sea urchin, and colo, to inhabit)

refers to the association of this species with echinoids.

REMARKS.—This is a common, sometimes abundant, associate of two edible sea urchins in the

West Indian region. The number of copepods associated with a host may be considerable, as in the

case of the seven Tripneustes ventricosus from

Kingston, Jamaica, which had an average number

of 146 copepods, including adults and copepodids,

per urchin.

From the other West Indian species, Macrochiron

sargassi, this species differs in the following details:

leg 5 of the female is more elongate and less clavate

in M. sargassi; the second antenna of the female

of M. echinicolum has a hooked and unsegmented

main claw; the second maxilla shows a slightly different apical armature in the two species; the

female maxilliped differs markedly in the ornamentation of the terminal segment; the caudal

rami of Af. echinicolum are more elongate.

Macrochiron lobatum Humes and De Maria,

1969

Macrochiron lobatum Humes and De Maria, 1969, pp. 139-

144, figs. 1-21 [from the hydroid Lytocarpus phoeniceus

Busk (=Lytocarpus spectabilis Allman), region of Nosy Be,

Madagascar].

Macrochiron lytocarpi Humes, 1966

Macrochiron lytocarpi Humes, 1966a, pp. 6-14, figs. 1-32

[from the hydroid Lytocarpus philippinus (Kirchenpauer),

region of Nosy Be, Madagascar]; 1969, p. 152 [from Lytocarpus philippinus, region of Nosy B6, Madagascar].

Macrochiron mutatum Stock, 1957

Macrochiron mutatum Stock, 1957, p. 381.

Macrocheiron fucicolum.—G. O. Sars, non Brady, 1917, pp.

163-165, pi. 91 [from Norway].

Macrocheiron mutatum.—Vervoort, 1964, p. 47.

Macrochiron rostratum Humes, 1966

Macrochiron rostratum Humes, 1966a, pp. 18-24, figs. 61-88

[from the hydroids Lytocarpus philippinus and Lytocarpus

spectabilis Allman (=Lytocarpus phoeniceus Busk), region
NUMBER 127 201

of Nosy Bo, Madagascar].—Humes and De Maria, 1969, pp.

1">2, 153 [from the liydroids l.ytocarpus philippinus, Lytoenrpus phoeniceus, Gymnangium hians (Busk), Gymnangium gracilicaulis (Jaderholm], and Aglaophenia delicatula

(Busk), region of Nosy Be, Madagascar].

Macrochiron sargassi G. O. Sars, 1916

KlC l RE S 111 , 11 2

Macrocliriron sargnssi G. O. .Sars, 1916, pp. 3, 10, 11, pi. 2,

[from Sargasso Sea].

Macrochiron iargassi.—Campbell, 1930, p. 181, pi. 1, figs.

13-16 [from Vancouver Island, Canada].—Clemens, 1933, p.

44.—Yeatman, 1962, pp. 2.r>6-2.r)9, figs. 1-17 [from Sargassum

washings. 27*49' N, 48° 29' W, Atlantic Ocean].-FuUon,

1968, p. 46-Shih. Figuicra, and Grainger, 1971, p. 155.

MATERIAL EXAMINED.—1 9» 2 cf cf» and 3 copepodids from washings of blackish brown compound

ascidians, Didemnum candidum Sav. (determined

by Dr. R. H. Millar, Millport), on mangrove roots,

Piscadera Inncrbay, Curasao, 6 October 1958 (ZMA

Co. 100.644); 3 9 9 from washings of orange synascidians Tridemnum ? orbiculare (Van Name) (determined by Dr. R. H. Millar), same locality and

date (ZMA Co.100.645); 1 copepodid, probably this

species, from a coelenterate, Renilla reniformis

(Pall.) (determined by F. M. Bayer, Washington),

in sandy bottom, depth about 1 m, Little Bay, St.

Martin, 2 February 1959 (ZMA Co. 100.646).

REMARKS.—There can be little doubt that the

species described by Sars from material collected

by Prince Albert of Monaco in the Sargasso Sea

is the same as the present form, found in washings

of compound ascidians. The very characteristic

female maxilliped and, to a lesser degree, the second antenna and the second maxilla agree well

with Sars' illustrations. On the other hand, there

are a few slight deviations from the original description. For example, Sars figures on the inner

side of the basipod of leg 1 a slender seta which

arises near the base of the endopod. Such a seta

never occurs, as far as we know, in the family

Lichomolgidae to which Macrochiron belongs, but

it is characteristic in certain siphonostome families. At any rate, neither our specimens nor those

described by Yeatman (1962) possess this seta, and

we can but suppose that the seta figured by Sars

is an error. The original illustrations do not show

aesthetes on the first antenna, but we must point

out that often they are discernible only with a

high-magnification or phase-contrast microscope

(phase-contrast being unknown in Sars' time).

All appendages of the female reillustrated here

(Figures 111, 112a-c) are on a somewhat larger

scale than in Sars' and Yeatman's papers. The

male (Figure \\2d-f) differs in the usual characters

of external anatomy (the shape of the genital segment, the number of urosomal segments); in the

presence of long, perhaps aesthete-like setae (two

on the second segment and one on the fourth segment); in the maxilliped (Figure 112d); in the less

elongate leg 5 (Figure 112/); and in the different

chaetotaxis of the third endopod segment of leg 1

(Figure \\2e).

The four females in our collection measure 0.84,

0.85, 0.85, and 1.06 mm in length, the two males

0.61 and 0.64 mm, and are consequently slightly

smaller than the specimens recorded by Sars (female 1.03 mm) and Yeatman (female 1.05-1.12

mm, male 0.82-0.92 mm).

The fifth leg in the female copepodid V is very

short (Figure Hid).

The association of M. sargassi with ascidians re*

ported here is unique so far as known in the genus.

The various species of Macrochiron hitherto have

been known only as free-living animals or as associates of hydroids (except for M. echinicolum

described above from echinoids).

It is a matter of considerable doubt whether

Campbell's record from the Canadian Pacific coast

actually belongs to this species. The descriptions,

illustrations, and locality do not provide sufficient

basis for a positive recognition of this record.

Macrochiron valgum Humes, 1966

Macrochiron valgum Humes, 1966a, pp. 14-18, figs. 33-60

[from the hydroid Lytocarpus philippinus, Nosy Be, Madagascar].-Humes and De Maria, 1969, p. 152 [from the

hydroids Gymnangium hians and Gymnangium gracilicaulis, region of Nosy Be, Madagascar],

Macrochiron vervoorti Humes and De Maria,

1966

Macrochiron vervoorti Humes and De Maria, 1966, pp. 144-

152, figs. 22-52 [from the hydroid Aglaophenia cupressina

Lamouroux, region of Nosy Be, Madagascar].
FIGURE 111.—Macrochiron sargassi G. O. San. Female: a, first antenna (A); b, rostrum (A); C,

mandible (B) ; d, leg 5 of copepodid V (c); e, leg 5, adult (D) ; /, first maxilla (B) ; g, second

antenna (A) ; h, labrum (D) ; i, maxilliped (D) ; ;', anal segment and caudal rami (D) ; k,

second maxilla (B) ; /, leg 1 (E) . Scale: A, 0.07 mm; B, 0.03 mm; c, 0.025 mm; D, 0.05 mm;

E, 0.1 mm. (All drawn from sample ZMA Co.100.645, except leg 5 of copepodid V, which is

drawn from sample ZMA Co.100.644.)
NUMBER 127 20S

FIGURE 112.—Macrochiron sargassi G. O. Sars. Female: a, third segment of endopod of leg 2

(E); b, leg S (E) ; c, leg 4 (E) . Male: d, maxilliped (F) ; e, leg 1 (c); /, leg 5 (F) . Scale: E,

0.1 mm; F, 0.025 mm; c, 0.05 mm.
204

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Meringomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum broadly

rounded posteroventrally. First antenna 7-segmented, with the formula in the female 4, 13, 6,

3, 4 + 1 aesthete, 2 -f- 1 aesthete, and 7 + 1

aesthete; in the male 4, 13 + 2 aesthetes, 6, 3 -j- 1

aesthete, 4 + 1 aesthete, 2 + 1 aesthete, and

7 -f 1 aesthete. Second antenna 4-segmented,

with the armature being 1, 1, 3, and I -|- 6 small

elements, there being a single terminal claw.

Labrum with a deep cleft between the two lobes.

Mandible with the base divided by an indentation

into an unornamented proximal part and a distal

area having on its convex side a scalelike region

with a row of spinules; lash long. Paragnath a

small hairy lobe. First maxilla with four elements.

Second maxilla of the usual lichomolgid type, with

the terminal lash and the adjacent seta of about

equal length, but sexually dimorphic, the outer

(ventral) proximal element on the second segment

in the female being minute, but in the male an

unusually large, proximally directed seta. Maxilliped in the female 3-segmented; in the male slender and 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Coxa of leg 1

with a posterior outer lobe. Armature of the usual

lichomolgidiform type, with leg 4 endopod being

0-I;II. Third segment of leg 4 exopod 111,1,5. Inner element on the coxa of leg 4 spiniform. Sexual

dimorphism in leg 1 endopod where the third segment is 1,1,4 instead of 1,5 as in the female (in Af.

devotus the inner spine is, however, only slightly

spiniform). Leg 5 with a free segment bearing two

terminal setae. Leg 6 in the female with two small

spines and a spiniform process; in the male a

posteroventral flap on the genital segment bearing

two setae.

Other features as in the three species described

below.

Associated with alcyonaceans.

TYPE-SPECIES.—Meringomolgus facetus, new species.

ETYMOLOGY.—The name is a combination of the

Greek words unoivfc (bristle), alluding to the long

proximal anteroventral element on the second segment of the second maxilla in the male, and

JIOXYO;. Gender masculine.

REMARKS.—Meringomolgus resembles Acanthomolgus in having a spine instead of a seta on the

first segment of the endopod of leg 4. It differs

prominently from that genus, however, in having

only one claw on the second antenna, the lash and

the adjacent seta on the second segment of the second maxilla being about equal in length (and in

the male this segment bearing an unusually large

outer proximal seta), the armature of the last segment of the exopod of leg 4 being 111,1,5, and the

inner element on the basis of leg 4 being a barbed

spine.

Key to Species of the Genus Meringomolgus

FEMALES

1. Genital segment not sharply incised laterally M. facetus

Genital segment sharply incised laterally 2

2. Second antenna with outer side of second segment smooth, first segment of leg 4 endopod

with inner element plumose proximally but barbed distally M. hamatus

Second antenna with outer side of second segment with spinules; first segment of leg 4

endopod with inner element barbed M. devotus

MALES

1. Third segment of leg 1 endopod with inner spine setiform and plumose M. devotus

Third segment of leg 1 endopod with inner spine barbed 2

2. Second antenna with both outer and inner sides of second segment bearing spinules

M. facetus

Second antenna with only inner side of second segment bearing spinules ilf. hamatus
NUMBER 127 205

Meringomolgus facetus, new species

FICURES 113-117

TYPE MATERIAL.—7 9 9,42 <$ cf > and 51 copepodids washed from a single colony of an alcyonacean, Sinularia polydactyla (Hemprich and Ehrenberg), in 2 m, at Ambariobe, a small island almost

between Nosy Komba and Nosy B£, northwestern

Madagascar, 22 August 1967, collected by AGH.

Holotype 9» allotype, and 43 paratypes (4

9 9, 39 <S d) deposited in USNM, the remaining

paratypes (dissected) and the copepodids in the

collection of AGH.

OTHER MATERIAL EXAMINED.—From Sinularia

polydactyla: 6 9 9» 5 cfcf. and 49 copepodids

from single colony, in 12 m, west of the harbor at

Hellville, Nosy B6, 4 August 1967. From Sinularia

minima Verseveldt: 6 9 9» 3 cf cf» and 6 copepodids from seven colonies, in 15 m, Bane du Touareg, south of Nosy Be, 13°32'05//

 S, 48°15'20" E,

1 September 1967.

FEMALE.—The body (Figure 113a) is cyclopiform. The length (not including the ramal setae)

is 1.36 mm (1.28-1.44 mm) and the greatest width

0.57 mm (0.59-0.68 mm), based on ten specimens

in lactic acid. The ratio of the length to the

width of the prosome is 1.43:1. The ratio of the

length of the prosome to that of the urosome is

2:1.

The segment of leg 5 (Figure 113&) is 99 x

200 n. There is no ventral intersegmental sclerite

between that segment and the genital segment.

The genital segment is 200 x 172 n in greatest dimensions, in dorsal view constricted in its posterior

half where the width is 104 n. The areas of attachment of the egg sacs are situated dorsolaterally

in the anterior half. Each area (Figure 113c) bears

two small, spiniform, finely barbed elements, 6.5 n

and 10 ^ in length, with a pointed process between them. The three postgenital segments are

73 x 94 fi, 44 x 100 n, and 76 x 88 n from anterior to posterior.

The caudal ramus (Figure 113d) is somewhat

wider than long, 32 x 41 n. The six setae are

naked except for the dorsal seta which is 72 n

long and delicately feathered. The outer lateral

seta and the outermost terminal seta are distally

weak and hyaline, and they were broken in all

specimens examined. The innermost terminal seta

is 320 n. The two long median terminal setae are

490 n (outer) and 680 n (inner), and both are

inserted between dorsal (unornamented) and ventral (with a few minute crenations but no spinules) flaps. The ramus bears an inner dorsal

setule and its outer margin has a very minute hair.

The body surface bears hairs (sensilla) as indicated in Figure ll$o,b. The posteroventral margin

of the anal segment is smooth.

The egg sac (Figure 113e) is oval, 495 x 242 n,

reaches slightly beyond the caudal rami, and contains many eggs about 45 n in diameter.

The rostrum (Figure 113/) is broadly rounded

posteroventrally.

The first antenna (Figure 113g) is 536 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 41 (88 n

along the anterior margin), 164, 39, 101, 57, 44,

and 43 n respectively. The formula for the armature is 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete,

and 7 -}- 1 aesthete. All the setae are naked.

The second antenna (Figure 113/J) is 4—segmented. Each of the first two segments bears an

inner seta. The third segment bears three setae.

The last segment, 91 n along its outer edge, 66 n

along its inner edge, and 26 n wide, bears a single

terminal claw 53 n along its axis and six small hyaline elements. All the setae are naked. The outer

surface of the second segment is ornamented with

a few small spines.

The labrum (Figure 114a) has two broad posteroventral lobes.

The mandible (Figure 1146) has on the concave

side of the base a deep indentation, distal to which

there is a row of long spinules; the convex margin

of the base bears a scalelike area with a row of

spinules followed by a serrated hyaline fringe. The

lash is moderately long and barbed on both sides.

The paragnath (Figures 114a,c) is a small hairy

lobe with a somewhat attenuated tip. The first

maxilla (Figure 114d) bears four naked setae. The

second maxilla (Figure 114e) is 2-segmented. The

first segment is large and unornamented. The second segment has only a vestige of an outer (ventral) proximal element; more distally it bears a

posterior surficial seta finely barbed along one edge

and a prominent, unilaterally spinose seta which

is about as long as the terminal lash (also spinose

along one edge and with a few slender spinules

distally along the opposite edge). The maxilliped

(Figure 114/) is 3-segmented. The first segment
206

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE MS.—Meringomolgus jacetus, new species. Female: a, dorsal (A); b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (c); e, egg sac,

dorsal (D) ; /, rostrum, ventral (£); g, first antenna, dorsal (B) ; h, second antenna, anterodorsal

(F) . Scale: A, D, 0.5 mm; B, E, 0.2 mm; c, 0.05 mm; F, 0.1 mm.
NUMBER 127

FIGURE 114.—Menngomolgus facetus, new species. Female: a, labrum, with paragnaths indicated

by broken lines, ventral (c); b, mandible, posterior (c); c, paragnath, ventral (H) ;

d, first maxilla, anterior (c); e, second maxilla, posterior (c); /, maxilliped, postero-inner (c);

g, area between maxillipeds and first pair of legs, ventral (B) ; h, leg 1 and intercoxal plate,

anterior (F) ; i, outer edge of coxa of leg 1, posterior (c). Scale: B, 0.2 mm; c, u, 0.05 mm,

F, c, 0.1 mm.
208

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1,5

111,1,5

1,11,3

111,1,5

1,11,2

111,13

is unornamented; the second segment bears two

unequal setae, the shorter one naked, the longer

one bilaterally spinose; and the third segment bears

a stout, naked spine, a small, naked spinule, and

a terminal spiniform barbed process lacking a distinct articulation.

The ventral area between the maxillipeds and

the first pair of legs (Figure 114g) is only slightly

protuberant, just behind the level of the maxilliped. A sclerotized line connects the bases of the

maxillipeds.

Legs 1-4 (Figures l\4h, 115fl-c) have trimerous

rami, except for the endopod of leg 4 which is

2-segmented. The armature is as follows:

Pt coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; 0-1

P, coxa 0-1 basis 1-0 exp 1-0; 1-1:

enp 0-1; 0-2

P, coxa 0-1 basis 1-0 exp 1-0; 1-1;

enp 0-1; 0-2

P4 coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; II

The inner element on the coxa of legs 1-3 is

large, setiform, and feathered, but in leg 4 this

element is shorter (31 ]>•), spiniform, and barbed.

The coxa of leg 1 bears an outer posterior lobe

(Figures H4h,i). The inner margin of the basis of

all four legs bears a row of hairs. The endopod

of leg 4 has hairs along the outer margins of both

segments. The first segment is 55 x 34 n (the

length not including the spinous processes) and

bears an inner distal barbed spine 52 n (Figure

l\5e). The second segment is 110 n long (including the terminal spiniform processes) and 28 n in

greatest width; the outer terminal spine is 31 n,

the inner spine 77 IA, both barbed. One female

showed slightly different proportions in this endopod (Figure 115d): the first segment 66 x 36 n

with its spine 70 n, and the second segment 143 x

25 fi (19 n in least width) with its terminal spines

44 n and 94 n. Although the lobe of the basis

overlying anteriorly the first segment of the exopod is usually smooth, in one female this lobe

bore a row of spinules (Figure 115/).

Leg 5 (Figure 115g) has a moderately elongated

free segment 104 x 22 n (29 n wide at the slight

proximal expansion and 16 \i wide at the tip),

slightly recurved, and not reaching the level of

the middle of the genital segment (Figure 1136).

The dorsal outer surface of the segment has large,

broad, scalelike spines, in flat view (Figure l\5h)

each scale being about 11 x 9 i». One terminal

seta is weak and hyaline, and broken in all specimens seen; the other seta is 83 n long and very

finely barbed along the middle of one edge. The

seta on the body near the insertion of the free segment is 30 n and feathered.

Leg 6 is probably represented by the two elements near the attachment of each egg sac (Figure

113c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.

MALE.—The body (Figure 116a) is more slender

than in the female. The length is 1.15 mm (1.09-

1.20 mm) and the greatest width 0.36 mm (0.35-

0.39 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome

is 2.08:1. The ratio of the length of the prosome

to that of the urosome is 1.71:1.

The segment of leg 5 (Figure 116b) is 52 x 112 ^.

There is no ventral intersegmental sclerite. The

genital segment is 185 x 169 n, slightly longer than

wide. The four postgenital segments are 49 x 81 n,

47 x 74 n, 34 x 70 n, and 50 x 73 n from anterior

to posterior.

The caudal ramus is similar to that in the female, but smaller, 28 x 34 n.

The surface of the body is ornamented with hairs

as in the female.

The rostrum is like that of the female.

The first antenna (Figure 116c) is segmented

and armed as in the female, but there are two

additional aesthetes on segment 2 and another on

segment 4, so that the formula is 4, 13 -\- 2 aesthetes, 6, 3 -L- 1 aesthete, 4 -\- 1 aesthete, 2

-f- 1 aesthete, and 7+ 1 aesthete. All the setae

are naked.

The second antenna (Figure 116d) is a little

more slender than in the female, with the fourth

segment markedly so. The dimensions of the fourth

segment are 105 n along the outer edge, 77 n along

the inner edge, and 17 n wide. In addition to the

armature and ornamentation already noted in the

female, the first segment bears a row of spines near

the insertion of the seta, the second segment bears

inwardly a row of slender spinules and three rows

of broad spines, and the fourth segment bears an

inner marginal row of small spinules and scattered

minute spines on its antero-inner surface. The

terminal claw is 57 n along its axis.
NUMBER 127

FIGURE 115.—Meringomolgus jacetus, new species. Female: a, leg 2, anterior (F) ; b, third

segment of endopod of leg 3, anterior (F) ; c, leg 4 and intercoxal plate, anterior (F) ; d,

endopod of leg 4, anterior (F) ; e, inner spine on first segment of endopod of leg 4, anterior

(H) ; /, lobe of basis overlying first segment of exopod of leg 4, anterior (c); g, leg 5, dorsal

(G) ; h, three scales on leg 5, antero-inner (i) . Scale: c, H, 0.05 mm; F, G, 0.1 mm; i, 0.02 mm.
210

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 116.—Meringomolgus facet us, new species. Male: a, dorsal (D) ; b, urosome, dorsal (B) ;

c, first antenna, dorsal (B) ; d, second antenna, antero-inner (F) ; e, second maxilla, posterior

(c); /, maxilliped, inner (B) . Scale: B, 02 mm; D, 0.5 mm; F, c, 0.1 mm.
NUMBER 127 211

a

FICUKE in.—Meringomolgus facetus, new species. Male: a, endopod of leg 1, anterior (c); b,

leg 5, dorsal (c); c, leg 6, ventral (c); d, suctorian attached to second segment of exopod of

leg 4, posterior (E) . Scale: c, 0.05 mm; E, 0.2 mm; c, 0.1 mm.

The labrum, mandible, paragnath, and first maxilla resemble those of the female. The second

maxilla (Figure 116<?) has an unusually large outer

(ventral) proximal seta on the second segment.

The maxilliped (Figure 116/) is slender and 4-segmented (assuming that the proximal part of the

claw represents a fourth segment). The first segment is unornamented. The second segment bears

two setae (one with a row of slender barbules

along one edge, the other smooth) and two rows

of spinules. The small third segment is unarmed.

The recurved claw is slender and elongated, 335 n

along its axis, with a small terminal lamella, and

with weak indication of division about midway.

Proximally the claw bears a long seta with unilateral subterminal barbules and only a trace of the

second seta usually seen in lichomolgids.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the female, except for the last segment of the endopod

of leg 1 (Figure 117a) where the formula is 1,1,4

instead of 1,5 as in the female. The outer spine,

recurved distally, is 56 fi and the inner straight

spine 23 \t. Legs 2-4 resemble those of the female.

Leg 5 (Figure H7b) has a slender, subrectangular, unornamented free segment 34 x 9 n. The two

naked terminal setae are 20 n and 65 n and the

slightly feathered seta on the body near the segment is 28 ix.

Leg 6 (Figure 117c) consists of a posteroventral

flap on the genital segment, bearing two naked

setae 26 n and 34 ji.

The spermatophore was not observed.

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—The specific name facetus (Latin,

elegant) alludes to the general form and ornamentation of this species.

REMARKS.—One female had a large suctorian

(Figure 117d) attached to the posterior surface of

the second segment of the exopod of the right leg 4.

The body of the suctorian is 244 x 138 n; the

tentacles and five buds are about 106 n long, making the overall length 350 n. It seems probable

that this suctorian is similar to the Ophryodendron

sp. reported from Lichomolgus singularipes by

Humes and Ho (1968c).

Meringomolgus devotus, new species

FICURES 118-120

TYPE MATERIAL.—49 $ ? , 41 o*c?» and 35 copepodids from three colonies of the alcyonacean
212 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Sinularia leptoclados (Hemprich and Ehrenberg),

in 1 m, Ankify, on the mainland of Madagascar

opposite Nosy Komba, 23 August 1967, collected

by AGH. Holotype $, allotype, and 84 paratypes

(46 $ $, 38 tfd1) deposited in USNM, the remaining paratypes (dissected) and the copepodids

in the collection of AGH.

OTHER MATERIAL EXAMINED (all from Sinularia

leptoclados).—15 ? $, 15 cf cf>

 anu<

 21 copepodids

from single colony, in 1 m, Ankify, 22 July 1967;

and 5 9 ? > 2 copepodids from single colony, in

1 m, Ankify, 11 August 1967.

FEMALE.—The body (Figure 118a) is cyclopiform. The length is 1.37 mm (1.18-1.46 mm) and

the greatest width 0.58 mm (0.53-0.64 mm), based

on seven specimens in lactic acid. The ratio of

the length to the width of the prosome is 1.51:1.

The ratio of the length of the prosome to that of

the urosome is 1.93:1.

The segment of leg 5 (Figure \\%b) is 94 x

180 n. Between this segment and the genital segment there is no ventral intersegmental sclerite.

The genital segment in dorsal view is broadened

in its anterior three-fourths (146 x 185 v) and

abruptly narrowed in its posterior fourth (50 x

114 \i). The overall length of the segment is 196 v-.

The margins of the expanded portion are slightly

lobulate as indicated in the figure. The areas of

attachment of the egg sacs are located dorsolaterally

at the middle of the anterior portion. Each area

(Figure 118c) bears two naked elements, 8 n and

17 n, with a spiniform process between them. The

three postgenital segments are 62 x 101 n, 39 x 88 n,

and 70 x 91 n from anterior to posterior.

The caudal ramus, 34 x 35 v, resembles that of

Af. facetus. The dorsal seta is 78 n and feathered.

The outer lateral seta and the outermost terminal

seta are broken. The innermost terminal seta is

210 n, and the two long median terminal setae 375 n

(outer) and 550 H (inner), both inserted between

unornamented flaps. All three of these setae are

naked.

The body surface is ornamented with hairs (sensilla) as in Figure 118aA The posteroventral margin of the anal segment is smooth.

The egg sac was not observed.

The rostrum is like that in M. facetus.

The first antenna, 477 n in length, is segmented

and armed as in Af. facetus. The lengths of the

seven segments (measured along their posterior

nonsetiferous margins) are 61 (89 n along the anterior margin), 140, 44, 75, 53, 42, and 34 n respectively. All the setae are naked.

The second antenna (Figure 118d) resembles that

of Af. facetus, but the fourth segment is more slender, 115 M along its outer edge, 88 n along its inner

edge, and 21 n wide. The claw is 62 fi along its

axis.

The labrum and paragnath are like those of Af.

facetus. The mandible (Figure 118e) resembles

closely that of Af. facetus. The first maxilla (Figure 118/) also resembles that of Af. facetus except

that the longest element has a few minute unilateral barbules. The second maxilla (Figure 118g)

is similar to that in Af. facetus, but the terminal

lash is armed with a dentiform spine followed by

a row of long, slender spines with slightly recurved

tips. The maxilliped (Figure lI8/») resembles that

of M. facetus, but the first and second segments are

ornamented with groups of spinules as shown in

the figure.

The ventral area between the maxillipeds and

the first pair of legs is like that in M. facetus.

Legs 1-4 are segmented and armed as in Af.

facetus. As in that species, the coxa of leg 1 has

an outer posterior lobe, the inner margin of the

basis in all four legs bears a row of hairs, and the

inner element on the coxa of leg 4 is short (32 n),

spiniform, and barbed, rather than long, setiform,

and feathered as in legs 1-3. The spine on the

last segment of the endopod of leg 1 (Figure 119a)

is straight. The endopod of leg 4 (Figure 1196)

closely resembles that of Af. facetus. The first segment is 51 x 37 n (not including the spinous processes) and its inner distal barbed spine is 66 ji. The

second segment is 113 n long (including the terminal spiniform processes) and 27 n in greatest width;

the outer terminal spine is 42 n, the inner spine

84 n, both barbed.

Leg 5 (Figure 119c) has a free segment somewhat

like that of Af. facetus, but the shape is slightly

different and the scalelike spines are fewer in number. The dimensions are 108 x 23 n (the width

taken at the middle). The three setae (two terminal and one on the body near the free segment) are

similar to those in Af. facetus.

Leg 6 is probably represented by the two elements near the attachment of each egg sac (Figure

118c).
NUMBER 127 21S

FIGURE lli.—Meringomolgus devotus, new species. Female: a, dorsal (A); b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, second antenna, antero-inner (D) ; e,

mandible, posterior (c); /, first maxilla, posterior (c); g, second maxilla, posterior (E) ; h,

maxilliped, postero inner (c). Scale: A, 1.0 mm; B, 02 mm; c, 0.05 mm; D, E, 0.1 mm.
214

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 119.—Meringomolgus devotus, new species. Female: a, endopod of leg 1, anterior (E) ; b,

leg 4, anterior (D) ; c, leg 5, dorsal (E) . Male: d, dorsal (F) ; e, urosome, dorsal (D) ; /, second

antenna, antero-inner (E) . Scale: D, K, 0.1 mm; r, 0.5 mm.
NUMBER 127 215

The color in life in transmitted light resembles

that of M. facetus.

MALE.—The body (Figure 119d) resembles that

of the female. The length is 0.88 mm (0.85-0.92

mm) and the greatest width 0.31 mm (0.30-0.33

mm), based on six specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.83:1. The ratio of the length of the prosome to

that of the urosome is 1.86:1.

The segment of leg 5 (Figure \\9e) is 34 x 84 n.

There is no ventral intersegmental sclerite. The

genital segment is 127 x 124 »i. The four postgenital segments are 33 x 63 n, 31 x 57 n, 22 x

52 »i, and 35 x 54 n from anterior to posterior.

The caudal ramus is like that of the female, but

smaller, 28 x 28 n.

The surface of the body is ornamented with hairs

as in the female.

The rostrum resembles that of the female.

The first antenna is similar to that of the female, but there are three additional aesthetes as

in the male of M. facetus. The second antenna

(Figure 119/) resembles that of Af. facetus. The

last segment is 83 n along its outer edge, 62 n along

its inner edge, and 16 |i wide; there are fewer

minute spines on the antero-inner surface than in

M. facetus. The terminal claw is 55 n along its

axis.

The labrumk mandible, paragnath, and first maxilla are like those in the female. The second maxilla (Figure 120a) has an unusually long outer

(ventral) proximal seta on the second segment.

The maxilliped (Figure 1206) resembles that of

M. facetus, but the claw is shorter (205 n along

its axis) and its proximal seta is relatively longer

than in that species.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female, except for the last segment of the endopod

of leg 1 (Figure 120c), where the spine (35 n long)

is slightly recurved rather than straight as in the

female and the adjacent element, though not distinctly spiniform as in M. facetus, is shorter than

in the female, straight, and feathered. Legs 2-4

are like those of the female.

Leg 5 (Figure 119e) resembles that of M. facetus, with the free segment 32 x 10 n.

FIGURE 120.—Meringomolgus devotus, new species. Male: a, second maxilla, posterior (E) ; b,

maxilliped, inner (D) ; c, endopod of leg 1, anterior (c) ; d, leg 6, ventral (E) ; e, spermatophores, attached to female, dorsal (D) . Scale: c, 0.05 mm; D, E, 0.1 mm.
216 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Leg 6 (Figure 120d) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 30 n and 16 n.

The spermatophore (Figure 120e), attached to

the female in pairs, is 150 x 44 n (not including

the neck).

ETYMOLOGY.—The specific name devotus (Latin,

attached or faithful) alludes to the association with

an octocoral.

REMARKS.—The female of this species may be

distinguished from M. facetus by the shape of the

genital segment; the male by the nature of the

sexual dimorphism in the armature of the last

segment of the endopod of leg 1. Af. devotus has

in both sexes the terminal lash on the second

maxilla ornamented differently than in Af. facetus.

Meringomolgus hamatus, new species

FIGURES 121-123

TYPE MATERIAL.—11 99 , 14 o* g from three

colonies of the alcyonacean Sinularia leptoclados

(Hemprich and Ehrenberg), in 1 m, Ankify, on the

mainland of Madagascar, opposite Nosy Komba,

23 August 1967, collected by AGH. Holotype $ ,

allotype, and 17 paratypes (7 $ $, 10 tfd1) deposited in USNM, the remaining paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED.—From Sinularia

leptoclados: 1 <? in 1 m, Ankify, 22 July 1967; 2

? ?. 5 cT cf, and 10 copepodids from single colony,

in 15 m, Nosy Iranja, southwest of Nosy Be",

13°35'1O" S, 48O49W' E, 9 August 1967; 3 $ ? ,

2 cT <^ from single colony, in 1 m, Ankify, 11

August 1967; and 3 $ ?, 4 <$ <$ and 3 copepodids

from single colony, in 20 m, west of Andilana,

Nosy Be, 13°18'S, 48°07' E, 24 August 1967. From

Sinularia maxima Verseveldt: 3 $ $, 13 cf tf, and

150 copepodids from single colony, in 1 m, Pte.

Lokobe, Nosy Be", 12 June 1967. From Sinularia

humesi Verseveldt: 3 ? ? , 6 o" o" from single

colony, in 18 m, Bane du Touareg, south of Nosy

Be\ 13°32'05" S, 48°15'2O" E, 11 July 1967; and

8 $ $, 12 cTcf, and 16 copepodids from single

colony, in 13 m, opposite Antsiabe, southern shore

of Nosy Komba, 2 September 1967.

FEMALE.—The body (Figure 121a) is more

slender than in either of the two preceding species.

The length is 1.32 mm (1.12-1.57 mm) and the

greatest width 0.57 mm (0.56-0.66 mm), based on

ten specimens in lactic acid. The ratio of the

length to width of the prosome is 1.74:1. The ratio

of the length of the prosome to that of the urosome is 1.92:1..

The segment of leg 5 (Figure 1216) is 99 x 224 n.

Between this segment and the genital segment there

is no ventral intersegmental sclerite. The genital

segment in dorsal view is 234 x 237 n, broadened

in its anterior three-fourths with regularly curved

margins but abruptly constricted in its posterior

fourth where the width is 135 n. The areas of

attachment of the egg sacs are situated dorsolaterally on the expanded anterior portion of the segment. Each area (Figure 121c) bears two unequal

elements, one 31 n and naked, the other 8.5 n with

a minute distal spur, and a spiniform process between them. The three postgenital segments are

73 x 117 n, 52 x 109 t», and 69 x 101 n from anterior

to posterior.

The caudal ramus, 42 x 45 n, resembles that of

Af. facetus and Af. devotus. The dorsal seta is 90 n

and feathered. The outer lateral seta and the outermost terminal seta are broken. The innermost

terminal seta is 295 n and the two long median

terminal setae 400 n (outer) and 510 n (inner),

both inserted between unornamented flaps. All

three of these setae are naked.

The body surface is ornamented with hairs (sensilla) as in Figure 12\a,b. The posteroventral margin of the anal segment is smooth.

The egg sac was not observed.

The rostrum (Figure 121d) has a broadly

rounded posteroventral margin in ventral view; in

lateral view (Figure 121^) it projects somewhat

ventrally.

The first antenna, 475 n long, is segmented and

armed as in the two preceding species. The lengths

of the seven segments (measured along their posterior nonsetiferous margins) are 44 (81»i along

the anterior margin), 135, 34, 72, 65, 52, and 36 n

respectively. All the setae are naked.

The second antenna (Figure 121/) is segmented

and armed as in the two previous species. The

second segment lacks ornamentation on its outer

margin. The longest of the three setae on the

third segment has extremely short barbules. The

fourth segment is 83 11 along its outer edge, 58 j*

along its inner edge, and 18 n wide. The claw is

61 ix along its axis.
NUMBER 127

FIGURE 121.—Meringomolgus hamatus, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, rostrum, ventral (D) ; e, rostrum, lateral

(E) ; /, second antenna, antero-inner (F) ; g, mandible, posterior (c); h, maxilliped, posteroinner (c). Scale: A, E, 03 mm; B, D, 0.2 mm; c, 0.05 mm; F, 0.1 mm.
218

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The labrum, mandible (Figure 121g), and paragnath resemble those in M. facetus and M. devotus.

The first maxilla is like that of M. devotus, but all

the elements are naked. The second maxilla is

similar to that of M. devotus. The maxilliped

(Figure 121 h) resembles in major respects that of

M. devotus. In one female the two setae on the

second segment were more unequal in length and

the unarticulated spiniform process on the third

segment had unusually long lateral spinules (Figure 122a).

The ventral area between the maxillipeds and

the first pair of legs is like that in the two preceding

species.

Legs 1-4 are segmented and armed as in M.

facetus and M. devotus. The detailed ornamentation closely resembles that in M. facetus. As in both

preceding species, the coxa of leg 1 has an outer

posterior lobe, and the inner element on the coxa

of leg 4 is short (39 »), spiniform, and barbed

rather than long, setiform, and feathered as in

legs 1-3. The inner margin of the basis in leg 4 is

naked, but in legs 1-3 it bears a row of hairs. The

endopod of leg 4 (Figure 1226) is more slender

than in either of the two preceding species. The

first segment is 12 x 29 n (not including the spinous

processes) and its inner distal spine is 61 n. This

spine (Figure 122c) is feathered with very slender

spinules in its proximal half but barbed in its

distal half. The second segment is 113 n long (including the terminal spiniform processes) and 17 (*

wide at the middle; the outer terminal spine is 25

\i and the inner spine 72 fi, both barbed.

Leg 5 (Figure 122d) has a slender, elongated free

segment reaching to about the posterior limit of

the expanded part of the genital segment. (In

Figure 1216 leg 5 appears to be shorter because

the legs were held at somewhat dorsal angle in the

specimen drawn.) The dimensions are 195 x 23 n

(the width taken at the middle), with the width

at the slight proximal expansion 31 n and the

terminal width 30 n. The three setae (two terminal

and one on the body near the free segment) resemble those in the two previous species, but the barbs

along one side of the intact terminal seta are more

prominent than in either of those two species.

Leg 6 is probably represented by the two elements

near the attachment of each egg sac (Figure 121c).

The color in life in transmitted light is similar

to that in the two previous species.

MALE.—The body (Figure 122?) is slender. The

length is 1.04 mm (0.90-1.16 mm) and the greatest

width 0.36 mm (0.31-0.43 mm), based on ten

specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.91:1. The ratio of

the length of the prosome to that of the urosome

is 1.69:1.

The segment of leg 5 (Figure 122/) is 47 x 117 ti.

There is no ventral intersegmental sclerite. The

genital segment is 205 x 179 n, a little longer than

wide, with its lateral margins in dorsal view

slightly flattened. The four postgenital segments are

47 x 83 \i, 47 x 78 n, 34 x 70 u, and 49 x 75 n from

anterior to posterior.

The caudal ramus is similar to that of the female,

but smaller, 33 x 34 n.

The surface of the body is ornamented with

hairs as in the female.

The rostrum is like that of the female.

The first antenna resembles that of the female,

but there are three additional aesthetes as in the

males of M. facetus and Af. devotus. The second

antenna (Figure 122g) has the same segmentation,

armature, and ornamentation as in those species.

The claw is 68 u along its axis.

The labrum, mandible, paragnath, and first

maxilla resemble those of the female. The second

maxilla (Figure 123a) has an unusually long outer

(ventral) proximal seta on the second segment. The

maxilliped (Figure 1236) resembles that in the two

previous species. The claw is 360 ti along its axis.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female, except for the last segment of the endopod

of leg 1 (Figure 123c), where the formula is 1,1,4

instead of 1,5 as in the female. The outer spine

on this segment is 65 n long, distinctly recurved

distally, and ornamented with prominent bilateral

stout spinules; the inner spine is 34 n, straight, and

barbed. Legs 2-4 are like those of the female. There

appears to be some variation in the dimensions of

the endopod of leg 4. The average dimensions for

three males are 37 x 26 >* for the first segment, with

its spine 62 n, and 95 x 14 n for the second segment.

The measurements for each of the three males are

as follows: (1) 34 x 21 »* with the spine 51 i*,

83 x 12 n, (2) 43 x 30 n with the spine 67 n, 98 x

13 n, and (3) 36 x 28 \i with the spine 67 n, 104

x 16.5 n.
NUMBER 127 219

FIGURE 122.—Meringomolgus hamatus, new species. Female: a, maxilliped, as seen in one individual, postero-inner (c); 6, leg 4, anterior (F) ; c, inner spine on first segment of endopod of

leg 4, anterior (c); d, leg 5, dorsal (F) . Male: e, dorsal (A) ; /, urosome, dorsal (B); g, second

antenna, antero inner (F) . Scale: A, 0.5 mm; B, D, 02 mm; c, c, 0.05 mm; F, 0.1 mm.
220

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 123.—Meringomolgus hamatus, new species.. Male: a, second maxilla (H) ; b, nuxilliped,

inner (B) ; c, endopod of leg 1, anterior (m; d, leg 6, ventral (c). Scale: B, 0.2 mm; c, 0.05

mm; H, 0.1 mm.

Leg 5 (Figure 122/) resembles that in the two

previous species, with the free segment 44 x 11 n.

Leg 6 (Figure 123d) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 44 n and 22 n.

The spermatophore was not observed.

ETYMOLOGY.—The specific name hamatus (Latin,

curved like a hook) refers to the recurved spine on

the last segment of the endopod of leg 1 in the

male.

REMARKS.—The female of M. hamatus may be

distinguished from both M. facetus and M. devotus

by the shape of the genital segment and the longer

leg 5; the male by the relatively long, hooked spine

on the last segment of the endopod of leg 1. In

both sexes of M. hamatus the spine on the first

segment of the endopod of leg 4 is half feathered

and half barbed rather than completely barbed as

in the two previous species. The second segment

of this endopod is more slender than in either

M. facetus or M. devotus.

Genus Metaxymolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded

posteroventrally or not well developed. First antenna 7-segmented, in the female with the armature

4, 13, 6, 3, 4 -f 1 aesthete, 2 + 1 aesthete, and

7 -f- 1 aesthete; in the male with 4, 13 -+- 2

aesthetes, 6, 3 -j- 1 aesthete, 4 -f- 1 aesthete,

2 + 1 aesthete, and 7 + 1 aesthete, so far as known.

Second antenna 4-segmented, with the formula

1, 1, 3, and II -j- several small elements, there

being two terminal claws.

Labrum incised medially. Mandible with the

basal region beyond the indentation having on its

convex side a scalelike area with spinules followed

by a serrated fringe and on its concave side a row

of spinules; lash long. Paragnath a small hairy

lobe. First maxilla with four elements. Second

maxilla of the usual lichomolgid type. Maxilliped

in the female 3-segmented, with a pointed tip; in
NUMBER 127 221

Partial Key to Species of the Genus Metaxymolgus

FEMALES

1. Genital segment in dorsal view constricted at junction of anterior two-thirds, forming a

"waist" Af. cinctus

Genital segment not constricted 2

2. Genital segment distinctly longer than wide; sides not expanded and nearly parallel 3

Genital segment not elongated; sides expanded to some degree 6

3. Leg 4 endopod functionally 1-segmented, with an incomplete line of articulation between

the two segments; caudal ramus nearly 4:1 M. claudus

Leg 4 endopod functionally 2-segmented, with a distinct line of articulation between the

two segments; caudal ramus at most 2:1 4

4. Longer of two claws in second antenna clearly longer than fourth segment Af. aculeatus

Longer of two claws on second antenna not longer than fourth segment, usually much

shorter 5

5. Spines on lash of second maxilla obtuse; caudal ramus subquadrate, 28 x 25 /i; body

length 1.10 mm Af. anthcliae

Spines on lash of second maxilla spiniform; caudal ramus about 1.8:1; body length 0.81 mm

Af. inaequalis

6. Free segment of leg 5 broad, 81 x 34 fi, its inner margin abruptly expanded a short

distance from distal end of segment, contour of margin from this point to base of

segment being irregular Af. singularipes

Free segment of leg 5 not so shaped 7

7. Genital segment clearly broader than long (ratio at least 1:156) 8

Genital segment not distinctly broader than long (ratio at most 1:1.1) 10

8. Free segment of leg 5 not ornamented with spinules; body length 130 mm Af. myorae

Free segment of leg 5 ornamented with spinules; body length less than 1.30 mm 9

9. Prosome broad and suborbicular; free segment of leg 5 76 u. long; setae on caudal ramus

barbed Af. patulus

Prosome not unusually broadened and not suborbicular; free segment of leg 5 49 p, long;

setae on caudal ramus naked. M. venustus

10. Caudal ramus 1.7:1 or longer 11

Caudal ramus less than 15:1 12

11. Caudal ramus 1.7:1; free segment of leg 5 not ornamented with spinules and without a

proximal inner expansion Af. cuspis

Caudal ramus 3.3:1; free segment of leg 5 ornamented with spinules and with a proximal

inner expansion Af. spinulifer

12. Caudal ramus quadrate or wider than long IB

Caudal ramus a little longer than wide 17

13. Free segment of leg 5 with a large proximal inner expansion Af. securiger

Free segment of leg 5 with only a slight expansion or none 14

14. Free segment of leg 5 not ornamented with spinules 15

Free segment of leg 5 ornamented with spinules 16

15. Body length 133 mm; lateral margins of genital segment smoothly rounded Af. gracilipes

Body length 0.84 mm; lateral margins of genital segment slightly irregular . Af. hetaericus

16. Free segment of leg 5 108 p. long; lateral margins of genital segment slightly irregular

Af • con it#lorfii s

Free segment of leg 5 77 /j. long; lateral margins of genital segment smoothly rounded

Af. inflatiseta

17. Body length 132 mm Af. sensilis

Body length less than 1 mm 18

18. Spines on lash of second maxilla very long, forming a crest; second antenna slender, with

two subequal claws nearly as long as fourth segment Af. bohdosus

Spines on lash of second maxilla short; second antenna robust, with two very unequal

claws distinctly shorter than fourth segment Af. chalmydis
222

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform pattern. Leg 4 exopod

with the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1 ;II, the seta feathered. Leg 1

endopod with the third segment having 1,1,4, so

far as known, instead of 1,5 as in the female. Leg

5 with a free segment having two terminal elements. Leg 6 represented by the two setae near

the genital openings.

Other features as in the species below.

Associated with opisthobranchs, bivalves, alcyonaceans, gorgonaceans, actiniarians, zoanthids, and

ophiuroids.

TYPE-SPECIES.—Metaxymolgus securiger (Humes).

ETYMOLOGY.—The name is a combination of the

Greek words Hexa^u (between, alluding to the rather

intermediate position of this genus within the

family Lichomolgidae) and H»/.Y<>;. Gender masculine.

REMARKS.—In the following key to the females,

four species (Af. brevifurcatus, Af. congoensis, Af.

hirsutipes, and Af. simplex) have been omitted

because of lack of information. A key to the males

has not been prepared. In many cases, the males of

this genus can be distinguished only by careful

study, and largely on the basis of qualitative

characters.

Metaxymolgus securiger (Humes, 1964)

Lichomolgus securiger Humes, 1964, pp. 68-71, figs. 109-138

[from the nudibranch Doris mabilla Abraham, Madagascar].

NEW HOST.—From the nudibranch Asteronotus

cespitosus (van Hasselt): 8 9 <j>, 48 Q" cf. and 49

copepodids from two hosts, intertidal, Ambatozavavy, Nosy Be\ Madagascar, 29 January 1964; IS

? ?, 23 o" cf, and 12 copepodids from single host,

in 15 cm, on sandy bottom with Cymodocea, Antafianambitry, Nosy Be\ 8 September 1964. Both

collections by AGH.

Metaxymolgus actdeatus (Humes and Ho, 1968)

Lichomolgus aculeatus Humes and Ho, 1968b, pp. 20-25,

figs. 97-113 [from the alcyonaceans Nephthea aberrans

Vcneveldt, Nephthea sphaerophora Rukenthal, Nephthea

crassa Kiikenthal. Nephthea tixierae Verseveldt, and Litth

phyton arboreum Furskal. region of Nosy Be, Madagascar].

NEW HOSTS (all from Madagascar, collected by

AH).—From Nephthea amentacea Studer: 33 9 9#

14 cf cf, and 6 copepodids, from single colony, in

2 m, west of Pte. de Tafondro, Nosy Be, 30 May

1967; 19 9 9 , 47 cf cf» and 23 copepodids, from

single colony, in 13 m, off Antsiabe, Nosy Komba,

near Nosy Be\ 2 September 1967.

From Nephthea bumasta Verseveldt: 64 9 $#

115 cf cT>

 an d 87 copepodids, from single colony,

in 8 m, Pte. Lokobe, Nosy Be, 25 July 1967.

From Nephthea filamentosa Verseveldt: 56 9 9»

45 cT cT> and 57 copepodids, from single colony, in

23 m, Tany Kely, near Nosy Be\ 30 July 1967.

From Nephthea galbuloides Verseveldt: 78 9 9 ,

72 0* cf. from single colony, in 1 m, Pte. Ambarionaomby, Nosy Komba, near Nosy Be\ 8 June 1967;

92 9 9 . 81 cf cf. and 21 copepodids, from single

colony, in 22 m, Tany Kely, near Nosy H<\ 17

June 1967; 17 9 9 , 22 cf cf. and 12 copepodids,

from single colony, in 25 m, Tany Kely, near Nosy

Be\ 14 August 1967.

From Nephthea lanternaria Verseveldt: 122 9 9>

17 cf cf • and 19 copepodids, from single colony, in

15 m, Bane de la Lanterne, Bay of Tsimipaika,

near Nosy Be\ 26 July 1967.

From Stereonephthya nosybearia Verseveldt: 1

9 . 5 cf cf > and 16 copepodids, from single colony,

in 10 m, Rocher du N.E., Nosy Be, 19 August 1967.

From Stereonephthya scaphis Verseveldt: 2 9 9»

11 cf cf > and 13 copepodids, from single colony,

in 24 rn, north of Ankazoberavina, near Nosy Be1,

13°27.6' S, 47°58.2' E, 25 August 1967.

Metaxymolgus antheliae, new species

FIGURES 124-126

TYPE MATERIAL.—24 9 9 , 67 cf cf» and 17 copepodids from single colony of the alcyonacean Anthelia glauca Lamarck, in 12 m, west of the harbor

at Hellville, Nosy Be, Madagascar, 4 August 1967,

collected by AGH. Holotype 9 , allotype, and 67

paratypes (17 9 9 , 50 cf cf) deposited in USNM,

the remaining paratypes and the copepodids in the

collection of AGH.

OTHER MATERIAL EXAMINED.—3 9 9, 5 cf cf» and

9 copepodids from single colony of Anthelia ternatana (Schenk), in 18 m, Bane du Touareg, south
NUMBER 127 223

of Nosy Be\ 13°32'05" S, 48°15'2O" E, 11 July 1967,

collected by AGH.

FEMALE.—The body (Figure 124a) has a moderately broad prosome. The length is 1.10 mm (1.02-

1.14 mm) and the greatest width is 0.49 mm (0.46-

0.52 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome

is 1.55:1. The ratio of the length of the prosome

to that of the urosome is 1.68:1.

The segment of leg 5 (Figure 1246) is 49 x 138 \i.

Between this segment and the genital segment there

is a short weak ventral intersegmental sclerite. The

genital segment is 172 x 130 n in greatest dimensions

in dorsal view, its sides not much expanded and

nearly parallel. The areas of attachment of the egg

sacs are located laterally at the middle of the segment; posterior to this level there is a transverse

dorsal sclerotized fold which does not continue ventrally. Each area of attachment (Figure 124c) bears

two slender naked setae, 7 n and 12 n, with an obscure spiniform process between them. The three

postgenital segments are 42 x 81 n, 29 x 73 n, and

36 x 68 n from anterior to posterior.

The caudal ramus (Figure 124d) is 28 x 25 n,

only a little longer than wide. The dorsal seta is

31 n and the outer lateral seta 68 n, both naked. The

outermost terminal seta is 109 » with inner proximal hairs. The innermost terminal seta is 151 n

with bilateral spinules. The two long median

terminal setae are 262 n (outer) and 400 n (inner),

both with spinules and both inserted between dorsal

(unornamented) and ventral (with a marginal row

of small spinules) flaps.

The body surface has a few hairs (sensilla) as

shown in Figure \24a,b. The posteroventral

margin of the anal segment bears a row of spinules

on both sides.

The egg sac (Figure 124a) is oval, 440 x 185 n,

reaches beyond the caudal rami, and contains 16-20

large eggs, each about 88 n in diameter.

The rostrum (Figure 124c) is broadly rounded

posteroventrally.

The first antenna (Figure 124/) is 338 n long. The

lengths of the seven segments (measured along their

posterior nonsetiferous margins) are 25 (55 n along

the anterior margin), 94, 24, 59, 49, 34, and 23 n

respectively. The formula for the armature is 4,

13, 6, 3, 4 -f- 1 aesthete, 2 + 1 aesthete, and

7 + 1 aesthete. All the setae are naked.

The second antenna (Figure 124g) is 4-segmented. Each of the first two segments bears a

seta. The third segment has three setae. The

fourth segment, 65 n along its outer edge, 35 n along

its inner edge, and 23 n wide, bears two short

unequal terminal claws. The stouter claw is 28 it

along its axis, the more slender claw 21 n. Near

the insertions of the claws there are five hyaline

elements, two of them large and three small. All

the setae are naked. The second segment has a few

small spines along its inner surface.

The labrum (Figure 124/i) has two rather elongated and rounded posteroventral lobes.

The mandible (Figure 124i) is deeply indented

on the concave margin of the base, beyond which

there is a row of long spinules; the convex margin

of the base is also indented, and distal to it is a

protruded scalelike area bearing a row of small

spinules and a hyaline, serrated fringe. The lash

is moderately long and barbed. On the posterior

surface of the mandible near the base of the lash

there is usually a group of long striations. (On

one mandible these striations were absent.) The

paragnath (Figure \24h) is a small hairy lobe. The

first maxilla (Figure 124;) has four elements, one

of the three terminal ones being unusually stout.

The second maxilla (Figure 125a) has an unornamented first segment. The second segment is armed

with a small setule on its proximal outer (ventral)

surface, a surficial posterior seta barbed along one

edge, an inner (dorsal) distal spine bilaterally

with large spinules, and is produced to form a lash

which is ornamented along one edge with a series

of large obtuse teeth grading distally into smaller

spines, and along the opposite edge with small

barbs. The maxilliped (Figure 1256) has two

groups of spinules on the first segment. The second

segment has two unequal setae, both with very

short barbules, and a group of long spinules. The

third segment has a slender, barbed spine and a

naked seta, and terminates in an unarticulated

spiniform barbed process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 125c) is moderately

protuberant; a sclerotized line connects the bases

of the maxillipeds.

Legs 1-4 (Figure 125d-g) have the same segmentation and spine and setal formula as in other

species in the genus—for example, M. cuspis

(Humes). The inner coxal element of leg 4 is a
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 124.—Metaxymolgus antheliae, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B); c, area of attachment of egg sac, dorsal (c); d, caudal ram us, dorsal (D) ; e, rostrum,

ventral (E) ; /, first antenna, dorsal (E) ; g, second antenna, anterior (F) ; h, labrum and paragnaths, ventral (E) ; i, mandible, posterior (F) ; ;, first maxilla, anterior (F) . Scale: A, 0.5 mm;

B, E, 0.1 mm; c, 0.02 mm; D, F, 0.05 mm.
NUMBER 127

FIGURE 125.—Atetaxymolgus antheliae, new species. Female: a, second maxilla, posterior (F) ; b,

maxilliped, posterior (F) ; c, area between maxillipeds and first pair of legs, ventral (B) ; d,

leg 1 and intercoxal plate, anterior (E) ; e, leg 2, anterior (E) ; /, third segment of endopod of

leg 3, anterior (E) ; g, leg 4, anterior (E) ; h, leg 5, dorsal (D) . Scale: B, E, 0.1 mm; D, F,

0.05 mm.
226 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

small, naked seta of 19 n instead of a long, feathered

seta as in legs 1-3. The inner margin of the basis

of leg 4 is smooth instead of having a row of hairs

as in the other legs. The exopod of leg 4 is 174 n

long. The endopod of leg 4 has hairs along the

outer margins of both segments. The first segment

is 36 x 29 n (without the spiniform processes) and

its inner distal seta is 33 n and lightly feathered.

(The feathering is visible only with high magnification, as with an oil immersion objective.) The

second segment is 73 n long (including the terminal

spiniform processes), 26 n in greatest width proximally, and 20 n in least width distally; the two

terminal spines are fringed, the outer S3 n, the

inner 55 \i.

Leg 5 (Figure 125/i) has an elongated free segment 75 n long. The greatest width at the proximal

inner expansion is 25 n. The only ornamentation

consists of a very few minute spines on the outer

surface of the segment. The two terminal naked

setae are 39 n and 60 n. The naked seta on the

body near the insertion of the segment is 32 n.

Leg 6 is probably represented by the two setae

near the areas of attachment of the egg sacs (Figure

124c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs grayish.

MALE.—The body (Figure 126a) is only slightly

more slender than in the female. The length is

0.81 mm (0.77-0.85 mm) and the greatest width

0.27 mm (0.25-0.29 mm), based on ten specimens

in lactic acid. The ratio of the length to the width

of the prosome is 1.78:1. The ratio of the length

of the prosome to that of the urosome is 1.49:1.

The segment of leg 5 (Figure 1266) is 36 x 86 |i.

There is no ventral intersegmental sclerite. The

genital segment is 190 x 156 n, longer than wide.

The four postgenital segments are 28 x 48 n, 23 x 47

n, 18 x 46 n, and 23 x 46 n from anterior to posterior.

The caudal ramus resembles that of the female

but is smaller, 23 x 19 n.

The surface of the body is ornamented with

hairs as in the female.

The rostrum is like that of the female.

The first antenna is segmented and armed as

in the female but there are two additional aesthetes

on segment 2 and one additional aesthete on segment 4. The second antenna (Figure 126c) shows

sexual dimorphism in its ornamentation. The first

segment has a group of small spinules on its inner

surface. The second segment has on its inner surface small spines proximal to the seta and minute

spinules distal to it. The fourth segment is 54 n

along its outer edge, 31 n along its inner edge, and

17.5 n wide; the two claws are 24 n and 20 n.

The labrum is like that of the female. The

mandible (Figure 126d) resembles that of the female, but the scalelike area on the convex margin

of the basal area is not as protruded. The paragnath

and first maxilla are similar to those in the female.

The second maxilla (Figure 126c) is much like

that of the female, but the large teeth on the lash

are spiniform rather than obtuse. The maxilliped

(Figure 126/) has an unornamented first segment,

the second segment has two slender naked setae

and two rows of long spinules, and the small third

segment is unornamented. The claw is 162 n along

its axis, shows a weak division about midway, and

proximally has two very unequal setae and a few

inner surficial striations.

The ventral area between the maxillipeds and

the first pair of legs resembles that of the female.

Legs 1-4 are segmented and armed as in the

female except for the last segment of the endopod

of leg 1 (Figure 126g) where the formula is 1,1,4

instead of 1,5 as in the female.

Leg 5 (Figure 1266) has a rectangular, unornamented free segment 31 x 7 n, its two terminal

naked setae being 19 n and 30 n, and the naked

seta on the body 22 n.

Leg 6 (Figure 1266) consists of a posteroventral

flap on the genital segment, bearing two naked

setae 20 n and 25 n.

The spermatophore (Figure \26h), attached to

the female in pairs, is oval, 159 x 79 n (not including the neck).

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—The specific name, antheliae, is

the genitive form of the generic name of the host.

REMARKS.—In nearly all species of Metaxymolgus

the lateral margins of the genital segment in the

female are expanded. The only species in which

this segment has almost parallel lateral margins,

as in the new species, is Af. aculeatus (Humes and

Ho). Af. antheliae may be easily separated from it,

however, by the proportions of the caudal ramus

(2.2:1 in Af. aculeatus). The mandible of the female

of Af. antheliae is unlike that of nearly all other
NUMBER 127 227

FIGURE 126.—Metaxymolgus antheliae, new species. Male: a, dorsal (c) ; b, urosome, dorsal (B) ;

c, second antenna, posterior (F) ; d, mandible, posterior (D) ; e, second maxilla, posterior (D) ; /,

maxilliped, inner (F) ; g, endopod of leg 1, anterior (>) ; h, spermatophores, attached to female,

dorsolateral (B) . Scale: B, 0.1 mm; D, F, 0.05 mm; c, 0.2 mm.

species. Only in M. singularipes (Humes and Ho)

does the scalelike area protrude in a somewhat

similar manner. The fifth leg of the female of Af.

singularipes, however, is wholly unlike that of the

new species. The obtuse teeth on the lash of the

second maxilla in the female of M. antheliae are

unique, all other species having pointed spiniform

teeth.

Metaxymolgus botulosus (Stock and Kleeton, 1963)

Lichomolgus botulosus Stock and Kleeton, 1963b, pp. 247-

251, figs. 1-3 [from the gorgonaceans Extnicella stricta

(Bert.), Banyuls, Mediterranean coast of France; and

Paramuricea chameleon (von Koch), Cap Creus, Spain].

Metaxymolgus brevifurcatus (Ummerkutty, 1962)

Lichomolgus brexnfurcatus Ummerkutty, 1962, pp. 59-62,

pi. 11, figs. 1-7, pi. 12, figs. 1-3 [from starfish washings, Gulf

of Mannar, southeastern India].

Metaxymolgus chlamydis, new species

FIGURES 127-129

TYPE MATERIAL.—6 9 $, 2 c? o"» and 2 copepodids from the bivalve Chlamys varia (Linnaeus),

Bane de St. Marc, Rade de Brest, France, 26 March

1956, collected by JHS. Holotype $, allotype, 6

paratypes (3 entire $ $, 2 dissected $ ?, and 1
228 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

dissected cf) and the 2 copepodids are deposited in

ZMA.

FEMALE.—The body (Figure 127a) has a moderately broadened prosome. The length and the

greatest width of two females in lactic acid (the

length not including the setae on the caudal rami)

are 0.98 x 0.44 mm and 0.97 x 0.51 mm. The ratio

of the length to the width of the prosome is 1.38:1.

The ratio of the length of the prosome to that of

the urosome is 2:1. The segment of leg 1 is distinctly set off from the head dorsally by a transverse furrow.

The segment of leg 5 (Figure 1276) is 78 x 160 n.

Between this segment and the genital segment there

is a short ventral intersegmental sclerite. The

genital segment is 156 x 172 n, a little wider than

long, in dorsal view only moderately expanded. The

areas of attachment of the egg sacs are situated

almost laterally but slightly dorsally just behind

the middle of the segment. Each area (Figure 127c)

bears two small, naked setae, 9 v and 7 n, with a

small spiniform process between them. The three

postgenital segments are 57 x 96 n, 42 x 91 n, and

48 x 94 n from anterior to posterior. The posteroventral border of the anal segment is smooth.

The caudal ramus (Figure 127d) is 43 x 35 n in

greatest dimensions. The outer lateral seta is 60 n,

the dorsal seta 33 n, and the outermost terminal

seta 68 n, all three of these setae being naked.

The innermost terminal seta is 140 n with inner

proximal hairs. The two long median terminal

setae are 300 n (outer) and 560 n (inner), both

with lateral spinules along their midregions and

both inserted between dorsal (unornamented) and

ventral (with a marginal row of minute spinules)

flaps.

The surface of the body bears a few small hairs

(sensilla) as in Figure 127a,6.

The egg sac (Figure 127a) is elongated oval,

450 x 230 n, reaches well beyond the caudal rami,

and contains many eggs about 52 n in diameter

but somewhat variable in shape.

The rostrum (Figure 127e) is broadly rounded

posteroventrally.

The first antenna (Figure 127/) is 392 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 22 (61 M.

along the anterior margin), 114, 24, 61, 57, 45, and

30 n respectively. The formula for the armature is

4, 13, 6, 3, 4 -f 1 aesthete, 2 -f 1 aesthete, and

7 _|_ l aesthete. All the setae are naked.

The second antenna (Figure 127g) is 4-segmented, with a small seta on the first and second

segments and three setae on the third segment, one

of them bent. The fourth segment, 87 >* along its

outer edge, 55 n along its inner edge, and 34 n

wide, bears two very unequal claws, 64 n and 23 n,

along their axes and five setae (two long and three

short). All the setae are naked.

The labrum (Figure 128a) has two divergent,

narrow lobes.

The mandible (Figure 1286) has on its concave

side a deep indentation followed by a row of long

slender spinules; on its convex side there is a

prominent scalelike area with spinules. The lash

is long and bilaterally spinulose. The paragnath is

a small I<>U- with a few hairs. The first maxilla

(Figure 128c) bears four elements. The second

maxilla (Figure 12H<1) has an unornamented first

segment. The second segment bears a hyaline outer

(ventral) setule, a surficial posterior seta barbed

along one edge, and an inner (dorsal) spiniform

element with prominent lateral spinules. The segment terminates in a long lash ornamented along

one edge with a row of large spines distally

diminishing in size. The maxilliped (Figure \2Se)

has an unornamented first segment, two finely

barbed setae on the second segment, and a naked

seta, a barbed spine, and a barbed, spiniform

terminal process on the third segment.

The ventral area between the maxillipeds and

the first pair of legs (Figure 128/) is not protuberant. A fine line connects the bases of the maxillipeds.

Legs 1-4 (Figure 128g-;) have trimerous rami

except for the endopod of leg 4 which is 2-segmented. The armature is like that in other species

of Metaxymolgus—for example, Af. cuspis (Humes).

The inner seta on the coxa of legs 1-3 is long and

plumose, but in leg 4 this seta is shorter (28 v)

and naked. The inner margin of the basis in leg 4

is smooth instead of haired as in the other legs.

The exopod of leg 4 is 125 n long. The endopod

varies slightly in form. In Figure 128; the first

segment is 28 x 23 n (not including the spiniform

processes) with its inner distal feathered seta 48 n.

The second segment is 47 x 22 n (greatest dimensions, including the terminal spiniform processes),

its two terminal fringed spines 27 n (outer) and 36
NUMBER 127 229

FIGURE 127.—Metaxymolgus chlamydis, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, areas of attachment of egg sac, dorsal and slightly lateral (c); d, caudal ramus, dorsal

(c); e, rostrum, ventral (B) ; /, first antenna, ventral (D); g, second antenna, posterior (D) .

Scale: A, 0.5 mm; B, 02 mm; c, 0.05 mm; D, 0.1 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 128.—Metaxymolgus chlamydis, new species. Female: a, labrum, ventral (E); b, mandible, posterior (c); c, first maxilla, ventral (c); d, second maxilla, posterior (c); e, maxilliped,

posterior (c); /, area between maxillipeds and first pair of legs, ventral (B) ; g, leg 1 and inter

coxal plate, anterior (D) ; h, leg 2, anterior (D) ; i, endopod of leg S, anterior (D) ; /, leg 4

and intercoxal plate, anterior (D) . Scale: B, 0.2 mm; c, 0.05 mm; D, r, 0.1 mm.
NUMBER 127

FIGURE 129.—Metaxymolgus chlamydis, new species. Female: a, endopod of leg 4, anterior (D) ;

b, leg 5, dorsal (E) . Male: c, dorsal (A) ; d, urosome, dorsal (B) ; e, maxilliped, inner (c); /,

endopod of leg 1, anterior (E) ; g, third segment of endopod of leg 2, anterior (E) ; h, third

segment of endopod of leg 3, anterior (E) ; i, leg 5, dorsal (c); /', leg 6, ventral (E) . Scale:

A, 0.5 mm; B, 0.2 mm; c, 0.05 mm; D, E, 0.1 mm.
232 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

n (inner). In Figure 129a the first segment is 33 x

32 n, the seta 58 n; the second segment is 65 x

26.5 n, its two spines 32 n and 52 n. Both segments

are haired along their outer margins; the second

segment bears minute terminal spinules.

Leg 5 (Figure 1296) has an unornamented free

segment, 58 x 24 n (greatest dimensions), with its

proximal inner margin slightly expanded. The two

terminal naked setae are 52 n and 68 n. The

adjacent seta on the body is 61 n.

Leg 6 is probably represented by the two setae

near the attachment of each egg sac (Figure 127c).

The color in life is unknown.

MALE.—The body (Figure 129c) resembles that

of the female in general form. The length and

the greatest width of two males in lactic acid (the

length excluding the ramal setae) are 0.88 x 0.39

mm and 0.91 x 0.40 mm. The ratio of the length

to the width of the prosome is 1.45:1. The ratio

of the length of the prosome to that of the urosome

is 1.5:1.

The segment of leg 5 (Figure 129d) is 39 x 105 n.

There is only an indistinct trace of the ventral

intersegmental sclerite. The genital segment is 166

x 172 n. The four postgenital segments are 29 x 68 n,

29 x 65 n, 26 x 60 n, and 28 x 65 ft from anterior

to posterior.

The caudal ramus is similar to that of the female

but smaller, 33 x 25 f*.

The body surface has a few hairs (sensilla) as in

the female.

The rostrum resembles that of the female.

The first antenna is like that of the female, but

there are three additional aesthetes (Figure 129c)

so that the formula is 4, 13 -f- 2 aesthetes, 6,

3 + 1 aesthetes, 4 -f 1 aesthetes, 2 -f- 1 aesthetes,

and 7+ 1 aesthete.

The second antenna resembles that of the female

but shows sexual dimorphism in having very small

spinules along the inner surfaces of the second and

fourth segments.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are similar to those in the

female. The maxilliped (Figure \29e) has an unornamented first segment. The second segment

bears two naked setae, a row of prominent spinules,

and a group of small spinules. The small third

segment is unornamented. The claw is 127 y. along

its axis, including the small terminal lamella, and

is weakly divided about midway. It bears proximally two very unequal setae, the longer one with

a few very small distal barbules.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented as in the female, with

the spine and setal formula as in that sex except

for the last segment of the endopod of leg 1 (Figure

129/) which has the formula 1,1,4 (the outer spine

being 25 n, the inner spine 34 n) instead of 1,5.

Slight sexual dimorphism is seen also in the last

segment of the endopod of leg 2 (Figure 129g)

where the three spines from outer to inner are 29 (i,

26.5 n, and 30 n (in the female these being 16.5

n. 16.5 n, and 21 HI and the spinifoim processes

are longer; and in leg 3 (Figure \29h) where the

spines are 22 ji, 31 n, and 34 i» (instead of 16.5 n,

19 n, and 24 u in the female) and the processes are

also longer than in the female. Leg 4 is like that of

the female.

Leg 5 (Figure 129*) has an unornamented, unexpanded, slender free segment, 33 x 10 n, bearing

terminally an inner fringed spiniform element 22 n

and a naked seta 44 n.

Leg 6 (Figure 129;) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 44 n and 45 n, near a small spiniform process.

The spermatophore was not observed.

The color in life is unknown.

ETYMOLOGY.—The specific name, chlamydis, is

the Latin genitive form of the generic name of the

host.

REMARKS.—Of all the species in the genus

Metaxymolgus, this species resembles most closely

M. commodus (Humes). However, that species from

nudibranchs in Madagascar differs in the caudal

ramus being wider than long and in the ornamented fifth leg in the female.

Metaxymolgus cinctus, new species

FIGURES 130, 131

TYPE MATERIAI 32 9 ? , 19 <? o", and 48 copepodids from single colony of the gorgonian Psammogorgia ramosa Kiikenthal, in 12 m, Rocher du

N.E., Nosy Be, Madagascar, 13°ir52" S, 48°19/24"

E, 19 August 1967, collected by AGH. Holotype $ .

allotype, and 39 para types (25 $ ? , 14 <$ tf) deposited in USNM, the remaining paratypes and

the copepodids in the collection of AGH.
NUMBER 127 233

FEMALE.—The body (Figure 130a) is moderately

slender. The length is 1.12 mm (1.06-1.19 mm) and

the greatest width 0.49 mm (0.48-051 mm), based

on ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.56:1. The

ratio of the length of the prosome to that of the

urosome is 2.15:1.

The segment of leg 5 (Figure 130b) is 60 x 132 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment.

The genital segment is 143 n long, in dorsal

view constricted sharply at the level of the junction

of the anterior two-thirds. The width anterior to

the constriction is 101 n, at the constriction 89 n,

and posterior to the constriction 106 n. The areas of

attachment of the egg sacs are situated laterally

near the constriction. Each area (Figure 130c) bears

two naked setae, 7 n and 10 n, with a spiniform

process between them. The three postgenital segments are 57 x 75 n, 42 x 66 n, and 37 x 71 n from

anterior to posterior.

The caudal ramus (Figure 130d) is 39 x 31 n,

only a little longer than wide. The dorsal seta is

47 n and the outer lateral seta 68 n, both naked. The

outermost terminal seta is 124 n, the innermost

terminal seta 177 n, both with bilateral spinules.

The two long median terminal setae are 319 n

(outer) and 374 n (inner), both with spinules and

both inserted between dorsal (unornamented) and

ventral (with a marginal row of minute spinules)

flaps.

The body surface has a few hairs (sensilla) as

shown in Figure 130a,6. The posteroventral margin

of the anal segment bears a row of minute spinules

on both sides.

The egg sac (Figure 130a) was incomplete in all

ovigerous specimens seen. Each egg is about 53 n

in diameter.

The rostrum is like that of M. antheliae.

The first antenna is 389 n long and resembles

that of M. antheliae. The lengths of the seven segments (measured along their posterior nonsetiferous

margins) are 30 (77 n along the anterior margin),

109, 31, 55, 50, 34, and 33 n respectively. The

formula for the armature is the same as in M.

antheliae. All the setae are naked.

The second antenna (Figure 130e) is segmented

and armed as in M. antheliae, with the formula 1,

1, 3, and II -(- 5 hyaline elements. The fourth segment is 72 n along its outer edge, 44 n along its

inner edge, and 23 n wide. Of the two unequal

terminal claws the more slender one is 32 n along

its axis, the stouter one 36 \i. All the setae are naked

and there is no fine ornamentation on the segments.

The labrum (Figure 130/) has two rather truncated and angular posteroventral lobes.

The mandible (Figure 130g) resembles that of

M. antheliae but the scalelike area is not as protruded and has longer spinules and the posterior

surficial striations seen in that species are absent.

The paragnath (Figure 130/) is a hairy lobe. The

first maxilla (Figure 130/i) has four slender elements. The second maxilla (Figure 130i) resembles

that of M. antheliae but the large teeth on the lash

are acutely pointed rather than obtuse and the

inner (dorsal) distal spine has unilateral spinules.

The relatively short maxilliped (Figure 130;") has

an unornamented first segment. The second segment bears two unequal barbed setae and a group

of erect spines. The third segment has a large naked

spine and a small naked seta and terminates in an

unarticulated, spiniform, naked process much

shorter than the spine.

The ventral area between the maxillipeds and

the first pair of legs (Figure 130ft) is slightly protuberant; a sclerotized line connects the bases of

the maxillipeds.

Legs 1-4 have the same segmentation and spine

and setal formula as in M. antheliae. On leg 1

(Figure 130/) the barbs on the outer spines of the

exopod are stronger than in M. antheliae. Leg 2

shows only subtle differences from M. antheliae

in the last segment of the exopod (Figure 131a).

Leg 3 is identical with that of M. antheliae. In leg

4 (Figure 131fr) the inner coxal element is a long,

feathered seta 55 n (nearly as long as in leg 3

where it is 78 \*), and the inner margin of the basis

is smooth instead of haired as in legs 1-3. The

exopod of leg 4 is 159 n long. The endopod has

outer hairs on both segments. The first segment

is 43 x 33 n (not including the spiniform processes) and its inner distal feathered seta is 90 n.

The second segment is 105 n long (including the

terminal spiniform processes), 26 n in greatest

width proximally, and 19 n in least width distally;

the outer terminal spine is 32 n, the inner 67 n,

both finely barbed.

Leg 5 (Figure 131c) has a free segment 73 x 30 n

in greatest dimensions. The two naked terminal

setae are 42 n and 58 n. The naked seta on the body
FIGURE 130.—Metaxymolgus cinctus, new species. Female: a, dorsal (A) ; b, urosome, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, second antenna, posterior (E) ; /, labrum, with paragnaths indicated by broken lines, ventral (F) ; g,

mandible, posterior (F) ; h, first maxilla, posterior (D) ; i, second maxilla, posterior (F) ; /.

maxilliped, anterior (F) ; k, area between maxillipeds and first pair of legs, ventral (E) ; /, leg 1

and intercoxal plate, anterior (E) . Scale: A, 0.5 mm; B, 02 mm; c, 0.02 mm; D, 0.05 mm;

E, r, 0.1 mm.
NUMBER 127

FIGURE 131.— Metaxymolgus cinetus, new species. Female: a, endopod of leg 2, anterior (E) ; b,

leg 4 and intercoxal plate, anterior (E) ; c, leg 5, dorsal and slightly inner (F) . Male: d, dorsal

(c); e, urosome, dorsal (B) ; /, second antenna, anterior (F) ; g, maxilliped, inner (E) ; h,

endopod of leg 1, anterior (F) ; i, spermatophores, attached to female, ventral (B) . Scale:

B, 0.2 mm; E, F, 0.1 mm; c, 0.5 mm.
236

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

near the insertion of the segment is 33 n. The segment is ornamented with small spines along its

outer surface and with larger spines on its ventroinner surface.

Leg 6 is probably represented by the two setae

near the areas of attachment of the egg sacs (Figure

130c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.

MALE.—The body (Figure 131d) has a form

similar to that of the female. The length is 0.93

mm (0.86-0.96 mm) and the greatest width 0.35

mm (0.33-0.36 mm), based on ten specimens in

lactic acid. The ratio of the length to the width of

the prosome is 1.49:1. The ratio of the length of

the prosome to that of the urosome is 1.77:1.

The segment of leg 5 (Figure 13 \e) is 32 x 87 v.

There is no ventral intersegmental sclerite. The

genital segment is 164 x 152 n. The four postgenital

segments are 25 x 63 n, 26 x 58 n. 20 x 55 n, and

22 x 58 n from anterior to posterior.

The caudal ramus resembles that of the female

but is smaller, 30 x 26 n.

The surface of the body is ornamented with hairs

as in the female.

The rostrum is similar to that of the female.

The first antenna is segmented and armed as in

the female but there are two additional aesthetes

on the second segment and one additional aesthete

on the fourth segment (Figure 131d). The second

antenna (Figure 131/) resembles that of the female

except for the fine ornamentation consisting of

numerous long inner spinules on segments 1 and 2

and a row of minute spinules along the outer edge

of segment 2 and the inner edge of segment 4. The

fourth segment is 77 n along the outer edge, 55 n

along the inner edge, and 18 n wide, the two claws

being 37 n and 32 n.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those in the female.

The maxilliped (Figure 131g) resembles in general

form that of M. antheliae. The claw is 228 n along

its axis, proximally with the usual two unequal

setae and without surficial striations.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female except for the last segment of the endopod

of leg 1 (Figure 131/i) where the formula is 1,1,4,

the two spines being very unequal. Between the

two spines there is a pointed process a little longer

than the shorter spine and covered with small

spines.

Leg 5 (Figure 13\e) has a rectangular free segment, 27 x 9 i», ornamented with a few minute

outer spinules. The two terminal setae are 29 it

and 35 n and the adjacent seta on the body 24 n,

all three setae naked.

Leg 6 (Figure 13If) consists of a posteroventral

flap on the genital segment, bearing two naked

setae 29 n and 42 n.

The spermatophore (Figure 1311), attached to

the female in pairs, is oval, 116 x 65 v- (not including the neck).

The color in life in transmitted light is similar

to that of the female.

ETYMOLOGY.—The specific name, cinctus (Latin,

girded or tied about) refers to the constriction of

the genital segment of the female, as if the segment were tied with a string.

REMARKS.—Two features of the new species serve

to distinguish it from all others in the genus: the

constriction of the genital segment in the female

is unique; and the very unequal nature of the two

spines on the last segment of the endopod of leg I

of the male and the spinose process between them

are characteristic.

Metaxymolgus claudus, new species

FIGURES 132-134

TYPE MATERIAL.—17 ? $ , 3 o" o", and 2 copepodids from the stomach of a basket star, Euryale

aspera Lamarck, at Palembang, Indonesia, collected

by MacGillivray. One female is made the holotype,

one male the allotype, and the remaining specimens

paratypes (ZMA Co. 102.287).

FEMALE.—The total length (excluding the setae

on the caudal rami) is 0.88 mm (0.85-0.90 mm)

and the greatest width of the cephalosome is 0.30

(0.29-0.31 mm), based on ten specimens.

The body (Figure 132a) is slender. The first

pedigerous segment is completely incorporated in

the cephalosome. The first metastomal segment is

about as wide as the cephalosome, the second and

third segments less wide. The first urosomal segment is longer than wide and carries leg 5. The

genital segment (Figure 1326) is 136 x 72 n, thus

very slender; there is one setule near each genital

opening. The three postgenital segments are un-
NUMBER 127 237

FIGURE 132.—Metaxymolgus claudus, new species. Female: a, dorsal (A) ; b, urosome (B) ; c,

caudal ramus (c); d, first antenna (c); e, second antenna (c); /, labrum (c); g, mandible

(D) ; h, first maxilla (D) ; i, second maxilla (D) ; /, maxilliped (D) . Scale: A, 02 mm; B, C,

0.1 mm; o, 0.02 mm.
238 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ornamented except for the anal segment (52 x 44

n) which has a row of minute spinules along its

posteroventral border (Figure 132c). The caudal

ramus (Figure 132c) is 82-83 M. long (mean 82.4 »i)

and 19-24 n wide (mean 23.1 n), thus nearly four

times as long as wide. The armature of the caudal

ramus consists of a smooth, lateral seta at almost

half its length, four plumose terminal setae, and a

smooth dorsal seta implanted near the base of the

innermost terminal seta.

The first antenna (Figure I32d) is 7-segmented.

The third segment is partially subdivided. The

lengths of the segments (measured along their posterior margins) are 15, 48, 15, 30, 31, 26, and 35 n

respectively. The formula for the armature is 4,

14, 6, 3, 4 + 1 aesthete, 2 -f- 1 aesthete, and 7 + 1

aesthete.

The second antenna (Figure 132c) is 4-segmented. The armature is 1, 1, 3, and 5 -j- two

jointed claws. The second and fourth segments

bear inner spinules.

The labrum (Figure 132/) is incised medially.

The mandible (Figure 132g) has an enlarged

basal portion and a slender lash. The medial (convex) margin is armed with twelve triangular teeth.

The first maxilla (Figure 132/J) has four setae—

two terminal, one subterminal, and one lateral.

The second maxilla (Figure 132i) has a very stout

basal segment. The distal segment attenuates into

a short lash armed with about twelve teeth. The

auxiliary lash is about as long as the main lash

and armed with smaller teeth. The maxilliped

(Figure 132;) consists of an unarmed basal segment, a slender second segment with two short

spines, and a rounded terminal segment ending in

a slender claw and a spine.

Legs 1-4 (Figures 133a-d) are biramous, with the

exopods 3-segmented and the endopods of legs 1-3

being 3-segmented. The endopod of leg 4 (Figure

133d) is short and functionally 1-segmented, being

provided with an incomplete line of articulation.

An intercoxal plate, lateral basipod seta, and median coxopod seta are present on all legs (the latter small and glabrous in leg 4). The chaetotaxis

formula is usual for the genus.

Leg 5 (Figure 133c) has a free segment which is

tapered distally, its greatest length 37 n and greatest

width 16 |i. The medial margin has a proximal

swelling and a distal row of fine cilia. A long,

smooth seta is borne near the implantation of the

free segment. Terminally the free segment is armed

with a smooth seta (46 n) and a spine (32 »).

MALE.—The total length (without the ramal

setae) is 0.77 mm (0.76-0.81 mm) and the greatest

width of the cephalosome 0.24 mm (0.23-0.26 mm),

based on three specimens. The urosome (Figure

134a) is 6-segmented. The segment of leg 5 is wider

than long. The genital segment (104 x 100 n) bears

posterolaterally two long setae (representing leg

6).

The caudal ramus is 62 x 22 n, thus nearly three

times as long as wide.

Secondary sexual differences occur only in the

following appendages: the strongly prehensile maxilliped (Figure 1346), leg 1 (Figure 134c) with

the third endopod segment having 1,1,4 instead of

1,5 as in the female, and leg 5 (Figure 134d) where

the free segment is less slender (16 x 11 o, not

tapering, with the medial margin not ciliated.

ETYMOLOGY.—The specific name, claudus (Latin,

cripple), alludes to the incomplete segmentation of

the inner ramus of leg 4 in both sexes.

REMARKS.—In most species in this genus the caudal ramus is distinctly shorter than in Af. claudus.

Af. an.theliae, Af. botulosus, Af. brevifurcatus, Af.

chylamydis, A/, cinctus, Af. commodus, Af. congoensis, Af. cuspis, Af. gracilipes, Af. hetaericus, Af. inflatiseta, Af. myorae, Af. patulus, Af. sensilis, Af.

singularipes, and Af. venustus) have, at least in the

female, a short caudal ramus of about equal length

and width. In Af. inaequalis and Af. aculeatus the

length of the caudal ramus is about twice the

width. In M. spinulifer, Af. simplex, and Af. hirsutipes the caudal ramus is 3-4 times as long as

wide, resembling that in the new species. These

three species may be distinguished from Af. claudus,

however, on other characters.

Metaxymolgus claudus differs from Af. spinulifer

in having a much shorter exopod in leg 4, in the

longer terminal claws on the second antenna, in

the much shorter terminal lash on the second maxilla, and in having short elements only on the second segment of the maxilliped of the female. Also,

it differs from the incompletely known Af. simplex

in having a short terminal lash on the second maxilla, in a much more complex terminal armature

on the second antenna (two claws and no seta in

Af. simplex), and in the more slender genital segment in the female. Finally, Af. claudus differs

from Af. hirsutipes in the shape of leg 5 in the fe-
NUMBER 127 259

e

FIGURE 133.—Metaxymolgus claudus, new species. Female: a, leg 1 and part of intercoxal plate;

b, leg 2 and part of intercoxal plate; c, third segment of endopod of leg S; d, leg 4 and intercoxal plate; e, leg 5. Scale: E, 0.05 mm, applies to all drawings.
240

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 134.—Metaxymolgus claudus, new species. Male: a, urosome (n); b, maxilliped (c); e,

leg 1 (E) ; d, leg 5 (D) . Scale: B, C, 0.1 mm; i>. 0.02 mm; r, 0.05 mm.

male (longer and curved in Af. hirsutipes), in the

more proximal position of the lateral seta on the

caudal ramus, in the shorter endopod of leg 4, and

in the different shape of the maxilliped of the

female.

The most remarkable character of Metaxymolgus

claudus, and by which it differs from all known

species of the genus, is the incomplete nature of

the articulation between the first and second segments of the endopod of leg 4 in both sexes. In

this feature M. claudus resembles the genera Kelleria and Octopicola. These genera are, however,

otherwise (especially in the structure of the various

cephalosomal appendages) very different. In the

totality of its morphological characters Af. claudus

fits best in Metaxymolgus. The somewhat reduced

nature of the fourth endopod may be related to

the endoparasitic mode of life of Af. claudus,

whereas all other species in this genus are ectoassociates.

Metaxymolgus comtnodus (Humes, 1964)

Lichomolgus commodus Humes, 1964, pp. 71-74, figs. 139-

165 [from the nudibranchs Hexabranchus orbicularis Abraham and Doridopsis ruber Kelaart, region of Nosy B£,

Madagascar].

NEW HOST.—From Hexabranchus sp. (cf. H. marginatus (Quoy and Gaimard): 19 ? ? , 21 rfc?.

and 39 copepodids, from single host, intertidal,

Antsakoabe, Nosy Be", Madagascar, 9 August 1964;

71 $ $, 75 tfef. and 44 copepodids, from five

hosts, intertidal, Nosy N'Tangam, Nosy Be\ 15

April 1964. Both collections by AGH.

Metaxymolgus congoensis (T. Scott, 1894)

Lichomolgus congoensis T. Scott, 1894a, pp. 120, 121, pi. 13.

figs. 39-48 [in surface tow-net, Bananah Creek, Congo

River, Africa].

Metaxymolgus cuspis (Humes, 1964)

Lichomolgus cuspis Humes, 1964, pp. 65-68, figs. 74-108

[from the actiniarians Radianthus ritteri (Kwietniewskt)
NUMBER 127 241

and Stoichaclis giganteum (Forskal), region of Nosy Be1-

Madagascar].-Bouligand, 1966, p. 269.

Palythoa liscia Haddon and Duerden, region of Nosy Be\

Madagascar].

Metaxymolgus gracilipes (A. Scott, 1909)

Lichomolgus gracilipes A. Scott, 1909, pp. 265, 266, pi. 68,

figs. 11-20 [from a nudibranch,5 Pepla Bay, east coast of

Rotti Island, near Timor, 10°28' S, 123°24' E.

Lichomolgus (Macrochiron) gracilipes.—Monod and Dollfus,

1932a, p. 139.

}Epimolgus gracilipes.—Slock, 1957, p. 381.

Macrochiron gracilipes.—Stock, 1957, p. 381.

Metaxymolgus hetaericus (Humes and Ho, 1968)

Lichomolgus hetaericus Humes and Ho, 1968c, pp. 663-668,

figs. 89-106 [from the alcyonaceans Cladiella pachy dados

(Klunzinger) and Cladiella krempfi Hickson, region of

Nosy Be, Madagascar].

NEW HOST.—204 $ ? , 112 cf cf. and 72 copepodids from single colony of the alcyonacean Cladiella laciniosa (Tixier-Durivault), in 2 m, west of

Pte. de Tafondro, Nosy Be, Madagascar, SO May

1967, collected by AGH.

Metaxymolgus hirsutipes (T. Scott, 1893)

Lichomolgus hirsutipes T. Scott, 1893, pp. 206, 207, pi. 4,

figs. 1-12 [from Firth of Forth, Scotland]; 1894b, pp. 234,

258; 1902, p. 470 [from North Craig, Firth of Forth, Scotland]; 1905, pp. 202, 203 [in tubes of the polychaete

Sabella, Scotland]; 1906, p. 352 [on tubes of Sabella, Bass

Rock and North Craig, Firth of Forth, Scotland].—Thompson and A. Scott, 1900a, p. 335 [from Zostera beds, Liverpool Bay, England]; 1900b, p. 143 [from Zostera beds,

Liverpool Bay, England].—d'Arcy Thompson, 1901, p. 47.—

Herdman, 1901, p. 27.-T. Scott and A. Scott, 1901, p. 352

[from Kolguev Island, near south end of Novaya Zemlia,

Arctic].-Stock, 1957, p. 381.

Macrocheiron hirsutipes.—G. O. Sars, 1917a, pp. 155-166, pi.

92 [from Norway].

^Macrochiron (Macrochiron) hirsutipes.—Sewell, 1949, pp.

100-104, figs. 24a-g, 25a-h [from weed-washings, Addu

Atoll, Maldive Archipelago].

Macrochiron hirsutipes.—Stock, 1957, p. 381.

Metaxymolgus inaequalis (Humes and Ho, 1966)

Lichomolgus inaequalis Humes and Ho, 1966a, pp. 3-6, figs.

1-28 [from the zoanthids Palythoa tuberculosa (Esper) and

5

 The host is one of the following three nudibranchs

collected at Siboga Station 301: Hexabranchus marginatus

(Quoy and Gaimard), Glossodoris annae (Bergh) (=Chromodoris quadricolor Riippel and Leuckart, var. annae Bergh),

and Trevelyana rubromaculata Bergh. See Monod and Dollfus, 1932a, p. 139 and table.

Metaxymolgus inflatiseta, new species

FIGURES 135-137

TYPE MATERIAL.—5 ? $, 5 cf cf> and 1 copepodid from one euthyneuran gastropod, Onchidium}

sp., intertidal under rock, Pte. Mahatsinjo, Nosy

Be, Madagascar, 22 June 1963, collected by AGH.

Holotype $, allotype, and 5 paratypes (3 $ $, 2

cf cf) deposited in USNM, the remaining paratypes

(dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED (all probably from

Onchidium sp.).—1 $, 2 cf cf» and 1 copepodid

from two hosts, intertidal, Pte. Lokobe, Nosy Be,

Madagascar, 5 August 1960; 1 Q* from one host,

intertidal, Pte. Mahatsinjo, Nosy Be, 21 August

1960; 4 $ 9« ^ cf cf. and 3 copepodids from two

hosts, intertidal, Ambanoro, Nosy Be, 23 August

1960; 2 $ $, 1 cf from one host, intertidal, Pte.

Ambarionaomby, Nosy Komba, near Nosy B£, 27

August 1960; 1 copepodid from two hosts, intertidal, Ambariotelo, a small island nearly between

Nosy Komba and Nosy Be, 9 September 1960; and

2 $ 9, 5 cT cf from four hosts, Tany Kely, a small

island south of Nosy Be, 9 August 1960. All collected by AGH.

FEMALE.—The body (Figure 135a) has a moderately expanded prosome. The length (not including the setae on the caudal rami) is 1.04 mm (1.02-

1.07 mm) and the greatest width 0.53 mm (0.51-

0.54 mm), based on five specimens in lactic acid.

The segment of leg 1 is incompletely set off from

the cephalosome. The epimeral areas of the segments of legs 2 and 3 are slightly truncated, those

of the segment of leg 4 rounded. The ratio of the

length to the width of the prosome is 1.28:1. The

ratio of the length of the prosome to that of the

urosome is 2.04:1.

The segment of leg 4 (Figure 135&) is 70 x 153 n.

There is no ventral intersegmental sclerite between

this segment and the genital segment. The genital

segment in dorsal view is 161 x 164 n, rather globular, but constricted posteriorly. The areas of attachment of the egg sacs are located dorsolaterally

just behind the middle of the segment. Each area

(Figure 135c) bears two naked setae, 10 \i and 8 n,

and a small spiniform process. The three postgenital segments are 54 x 96 n, 39 x 83 n, and 49 x
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 135.— Metaxymolgus inftatiseta, new species. Female: a, dorsal (A) ; b, urosomt, dorsal

(B) ; c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum, ventral (E) ; /, first antenna, anterodorsal (E) ; g, second antenna, postero-inner (E) ; h, labrum,

ventral (F) ; i, mandible, posterior (D) . Scale: A, 0.5 mm; B, 02 mm; c, 0.0S mm; D, F, 0.05

mm; E, 0.1 mm.
NUMBER 127 24S

83 n from anterior to posterior. The anal segment bears on its posteroventral margin a row of

minute spinules on both sides.

The caudal ramus (Figure 135d) is quadrate,

30 x 30 n. The outer lateral seta is 60 n, the dorsal

seta 50 u. and the outermost terminal seta 68 n,

all three of them naked. The innermost terminal

seta is 143 n with proximal hairs. The two long

median terminal setae are 300 v (outer) and 495 n

(inner), both with very fine barbules along their

midregions.

The body surface bears very few hairs (sensilla).

The egg sac (Figure 135a) is elongated, 760 x

240 n. reaching to the ends of the longest ramal

setae. There are many eggs, each about 60 n in

diameter.

The rostrum (Figure \S5e) has an incomplete,

broadly rounded posteroventral margin.

The first antenna (Figure 135/) is slender, 360 n

long, and 7-segmented. The lengths of the segments (measured along their posterior nonsetiferous margins) are 28 (55 n along the anterior

margin), 105, 26, 59, 52, 39, and 24 n respectively.

The formula for the armature is 4, 13, 6, 3, 4 -|- 1

aesthete, 2 -f 1 aesthete, and 7 -f 1 aesthete. All

the setae are naked.

The second antenna (Figure 135g) is 4-segmented. Each of the first two segments bears a

slender seta. The third segment bears two slender

setae and a spiniform recurved seta. The fourth

segment, 74 \i along its outer edge, 50 n along its

inner edge, and 31 n wide, bears terminally two

very unequal claws, 55 n and 32 n, and five setiform

elements. All the elements are naked.

The labrum (Figure 135/i) has two broad posteroventral lobes separated by a shallow median

incision. The mandible (Figure 135i) has on the

convex side of its basal region beyond the indentation a st-a lei ike area with strong spinules followed

by a row of shorter spinules and on the concave

side a row of slender spinules; the lash is long and

sparsely barbed. The paragnath (Figure 135/i) is a

small hairy lobe. The first maxilla (Figure 136a)

has four elements, one of them with short spinules.

The second maxilla (Figure 1366) has an unornamented first segment. The second segment bears

on its convex (ventral) side a small proximal spinule, on its posterior surface a seta barbed along

one edge, and on its concave (dorsal) side a spine

with strong lateral spinules. The lash is elongated

with graduated spines along one edge and with

both edges barbed distally. The maxilliped (Figure 136c) is 3-segmented. The first segment is unarmed. The second segment bears two very unequal

setae, one short and naked, the other more than

ten times as long and barbed. The third segment

bears a naked spine and a small naked setule and

terminates in a spiniform barbed process with a

slender tip.

The ventral area between the maxillipeds and

the first pair of legs (Figure 136d) is not protuberant. A sclerotized line connects the bases of

the maxillipeds.

Legs 1-4 (Figures \S6e-h) have 3-segmented rami

except for the endopod of leg 4 which is 2-segmented. The armature is typically lichomolgid.

The inner seta on the coxa of legs 1-3 is large and

plumose, but in leg 4 is minute (9 n long) and

naked. The inner margin of the basis is haired

in legs 1-3 but naked in leg 4. The exopod of leg

4 is 159 n long, the third segment having 11,1,5.

The endopod of leg 4 has hairs along the outer

margins of both segments. The first segment is 31

x 18 n, with its inner distal plumose seta 37 I*. The

second segment is 58 x 18 n in greatest dimensions

(including the terminal spiniform processes); the

outer terminal spine is 30 \i and the inner spine

45 |i, both fringed.

Leg 5 (Figure 137a) has an elongated free segment, 77 x 21 n, without a distinct proximal inner

expansion, though a crease in the cuticula suggests

such a development. (In an outer view of the segment, as in Figure 1376, neither a crease nor an

expansion is evident.) The segment bears two

terminal naked setae, 66 n and 72 n, and is ornamented along the outer dorsal side with minute

spines. The adjacent seta on the body is naked.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 135c).

The color in life in transmitted light is translucid, the ovary gray, the eye red.

MALE.—The body (Figure 137c) resembles in

general form that of the female, though the

epimeral areas of the segment of leg 4 are truncated rather than rounded. The length (without

the ramal setae) is 0.89 mm (0.84-0.94 mm) and

the greatest width 0.37 mm (0.36-0.38), based on

five specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.41:1. The
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 136.—Metaxymolgus inflatiseta, new species. Female: a, first maxilla, anterior (D) ; b,

second maxilla, posterior (D) ; c, maxilliped, posterior (D) ; d, area between maxillipeds and

first pair of legs, ventral (E) ; e, leg 1 and intercoxal plate, anterior (E) ; /, leg 2 and intercoxal plate, anterior (E) ; g, endopod of leg 3, anterior (E) ; h, leg 4 and intercoxal plate,

anterior (E) . Scale: D, 0.05 mm; E, 0.1 mm.
NUMBER 127

FIGURE 137.—Metaxymolgus inflatiseta, new species. Female: a, leg 5, dorsal (F) ; b, free segment of leg 5, outer (F) . Male: c, dorsal (c); d, urosome, dorsal (B) ; e, second antenna,

postero inner (F) ; /, maxilliped, inner (E) ; g, endopod of leg 1, anterior (F) ; h, third segment of endopod of leg 2, anterior (D) ; i, third segment of endopod of leg 3, anterior (D) ; ;,

leg 5, dorsal (c). Scale: B, 0.2 mm; c, 0.03 mm; D, F, 0.05 mm; E, 0.1 mm; c, 0.3 mm.
246 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

ratio of the length of the prosome to that of the

urosorae is 1.46:1.

The segment of leg 5 (Figure 137d) is 44 x % ji.

There is no ventral intersegmental sclerite. The

genital segment is 180 x 195 n, a little wider than

long. The four postgenital segments are 38 x 65 n,

36 x 60 n, 22 x 58 n, and 31 x 61 n from anterior

to posterior.

The caudal ramus is quadrate as in the female,

but smaller, 25 x 25 i*.

The body surface is largely unornamented as in

the female.

The rostrum resembles that of the female.

The first antenna is segmented and armed as in

the female, but there are two additional aesthetes

on the second segment and another additional aesthete on the fourth segment (Figure 137c) so that

the formula is 4, 13 + 2 aesthetes, 6, 3 -\- 1 aesthete, 4 -f 1 aesthete, 2 + 1 aesthete, and 7 -f 1

aesthete.

The second antenna (Figure 137?) shows wxu.il

dimorphism in certain details. The setae on the

first and second segments and one seta on the third

segment are inflated proximally. The spiniform

seta on the third segment is less recurved than in

the female and bears small barbules. The inner

surface of the second segment bears minute spinules.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The slender maxilliped (Figure 137/) is 4-segmented (assuming that the proximal part of the

claw represents a fourth segment). The first and

third segments are unornamented. The second segment bears two setae, barbed along one edge, and

a row of spines. The claw is 140 u along its axis

including the terminal lamella, is weakly divided

about midway, and bears proximally two small

setae, one of them barbed along one edge.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 have the same segmentation and armature as in the female except for the formula of the

third segment of the endopod of leg 1 (Figure

137g) which is 1,1,4 instead of 1,5 as in that sex.

In leg 2 the third segment of the endopod (Figure

137/J) is 53 n long (without the longest spiniform

process of 12 u) and the three spines from outer to

inner are 23, 26, and 36 n. (In the female this segment is 65 ti, without the process of 7 n, and the

spines are 23, 26, and 34 n.) In leg 3 the third

segment of the endopod (Figure 137/) is 52 n (without the spiniform process of 18 ») and the three

spines from outer to inner are 23, 23, and 35 p.

(In the female this segment is 62 n, without the

process of 7 n, and the spines are 23, 26, and 34 n.)

The endopod of leg 4 is similar to that in the

female.

Leg 5 (Figure 137/) has a slender, elongated unornamented free segment, 33 x 8 n, with nearly

parallel sides. The two terminal elements comprise

an inner spine, 15 M, with a unilateral lamella and

a long naked seta, 46 n.

Leg 6 consists of a posteroventral flap on the

genital segment bearing two naked setae 39 n and

50 n (Figure 137d).

The spermatophore was not observed.

The color in life resembles that of the female.

ETYMOLOGY.—The specific name inflatiseta

(Latin, inftatus, swollen, and seta) alludes to the

three swollen setae on the second antenna of the

male.

REMARKS.—In the form of the genital segment

of the female, Af. inflatiseta resembles four known

species: Af. securiger (Humes), Af. botulosus (Stock

and Kleeton), Af. sensilis (Humes), and Af. hetaericus (Humes and Ho). In the first three of these,

however, the free segment of leg 5 in the female

has a proximal inner expansion. In Af. hetaericus

the free segment of leg 5 is unornamented and

short, the ratio being 2.3:1.

The male of Af. inflatiseta may be separated from

all others in the genus by the peculiar inflated setae

on the second antenna. The only other species

with swollen setae on this appendage is Af. sensilis,

but here the swelling has a different form (compare Humes, 1964, fig. 192, with our Figure 137<r).

The free segment of leg 5 in Af. sensilis has a few

small spinules, whereas in Af. inflatiseta it is unornamented.

Certain appendages of Af. inflatiseta and Af. securiger are sufficiently similar to suggest a rather

close relationship between these two species, both

from opisthobranchs. Such similarities are seen in

the second antenna, mandible, second maxilla,

maxilliped, and legs 1—4.

Metaxymolgiis myorae (Greenwood, 1971)

Lichomoljpts myorae Greenwood, 1971, pp. 298-306, figs.

la h, 2a-h, 3a i, table 1 [from the actiniarian Stoichactis

haddoni (Sav.-Kent), Moreton Bay, Queensland, Australia].
NUMBER 127 247

Metaxymolgus patulus (Humes, 1959)

Lichomolgus patulus Humes, 1959, pp. 298-303, figs. 65-96

[from the nudibranch Phyllidia trilineata Cuvier, region

of Nosy Be, Madagascar].-Gotto, 1962, p. 104.

NEW HOSTS.—From the alcyonacean Sinularia

mayi Liittschwager: 2 $ ? , 1 cf from two colonies,

in 20 m, Tany Kely, a small island south of Nosy

Be, Madagascar, 20 December 1963, collected by

JHS.

From the crinoid Cenometra emendatrix (Bell):

1 $» 6 cf cf from five hosts, in 20 m, Tany Kely,

near Nosy Be, Madagascar, 10 September 1963, collected by AGH.

Metaxymolgus sensilis (Humes, 1964)

Lichomolgus sensilis Humes, 1964, pp. 74-78, figs. 166-197

[from the nudibranch Trevelyana rubromaculata Bergh,

region of Noty Be, Madagascar].

Metaxymolgus simplex (Thompson and A. Scott,

1903)

Lichomolgus simplex Thompson and A. Scott, 190S, p. 279,

pi. 15, figs. 27-34 [from washings of sponges. Gulf of Mannar, Ceylon].-Stock, 1957, p. Ml.

Macrochiron simplex.—Stock, 1957, p. 381.

Metaxymolgus singularipes (Humes and Ho, 1968)

Lichomolgus singularipes Humes and Ho, 1968c, pp. 685-

689, figs. 168-187 [from the alcyonacean Parerythropodium

rubiginosum Verseveldt, region of Nosy Be, Madagascar].

NEW RECORDS.—2 $ $, 6 cf cf, and 2 copepodids

from pieces of single colony of Parerythropodium

rubiginosum Verseveldt, in 1 m, Pte. Mahatsinjo,

Nosy Be, Madagascar, 7 June 1967; 10 9 9 , 4 cf cf»

and 4 copepodids from pieces of single colony of

Parerythropodium sp., in 1 m, Pte. a la Fievre, Nosy

Be, Madagascar, 15 June 1967. Both collections by

AGH.

Metaxymolgus spinulifer (Humes and Frost, 1964)

Lichomolgus spinulifer Humes and Frost, 1964, pp. 142-145,

figs. 134-166 [from the alcyonacean Lemnalia sp., Nosy

Be, Madagascar].—Bouligand, 1966, p. 269.—Humes and

Ho, 1968b, p. 23 [from the alcyonaceans Lemnalia flava

May, Lemnalia elegans (May), Lemnalia amabilis TixierDurivault, Lemnalia africana (May), and Paralemnalia

thyrsoides (Hemprich and Ehrenberg), region of Nosy

Be, Madagascar].-Humes, 1970c, p. 160.

NEW ALCYONACEAN HOSTS (all collected by AGH).

—From Lemnalia longiramus Verseveldt: 104 9 9 (

186 cf cf. and 173 copepodids from single colony,

in 12 m, west of the harbor at Hellville, Nosy B6,

Madagascar, 4 August 1967.

From Lemnalia digitata (May): 25 9 $» ^ o* o*»

and 67 copepodids from single colony, in 17 m,

near the black buoy between Pte. Ambarionaomby,

Nosy Komba, and Pte. de Tafondro, Nosy B£,

Madagascar, 5 August 1967; 15 9 9 , 16 cfef. and

1 copepodid from single colony, in 2 m, Ambariobe,

a small island almost between Nosy Komba and

Nosy Be, Madagascar, 22 August 1967.

From Lemnalia cervicornis (May): 99 9 $» 70

cf cT, and 48 copepodids from single colony, in

20 m, Bane de Cinq Metres, near Nosy Be, Madagascar, 6 August 1967; 1 9» $ 6* cf from two small

colonies, in 20 m, Bane de Dzamandzar, near Nosy

Be, Madagascar, 30 August 1967.

From Lemnalia crassicaulis Verseveldt: 62 9 $»

41 cf cf > and 2 copepodids from single colony, in

20 m, Bane de Cinq Metres, near Nosy Be, Madagascar, 6 August 1967.

From Lemnalia tenuis Verseveldt: 23 9 9» H

cf cf, and 2 copepodids from single colony, in 50 m,

Bane de Cinq Metres, near Nosy B£, Madagascar,

3 September 1967.

From Lemnalia madagascarensis Verseveldt: 51

9 ?» 73 cf cf, and 38 copepodids from single colony, in 24 m, Bane des Freres, Isles Mitsio, near

Nosy Be, Madagascar, 12°58' S, 48°28' E, 17 August

1967.

From Paralemnalia clavata Verseveldt: 93 9 ?»

95 cf cf, and 19 copepodids from single colony, in

2 m, Ambariobe, near Nosy B£, Madagascar, 25

May 1967; 52 9 9 , 44 0" cf. and 10 copepodids

from single colony, in 1 m, Ambariobe, 25 June

1967.

From Sinularia polydactyla (Hemprich and

Ehrenberg): 2 9 ? from single colony, in 2 m, Ambariobe, near Nosy B£, Madagascar, 22 August

1967.

NEW RECORDS FROM ALCYONACEANS (collected by

AGH).—From Lemnalia flava (May): 186 99 , 153

cf cf, and 14 copepodids from single colony, in 1 m,

Pte. Ambarionaomby, Nosy Komba, near Nosy Be,

Madagascar, 8 June 1967; 2 9 9 , 2 cf cf from single

colony, in 1 m, Pte. Ambarionaomby, 8 June 1967;

2 9 9, 2 cf cf from single colony, in 1 m, Pte.

Ambarionaomby, 8 June 1967; 136 9 9 , 66 cf cf,
248 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

and 9 copepodids from single colony, in 2 m, east

of Ambariotelo, a small island almost between Nosy

Komba and Nosy Be, Madagascar, 20 July 1967;

and 25 ? 9 , 38 tf tf, and 6 copepodids from single

colony, in 1.5 m, Ambariobe, near Nosy Be, Madagascar, 22 August 1967.

From Lemnalia africana (May): 8 $ $ , 16 o* o"»

and 21 copepodids from single colony, in 12 m,

west of the harbor at Hellville, Nosy B6, Madagascar, 4 August 1967.

From Paralemnalia thyrsoides (Hemprich and

Ehrenberg): 36 $ $, 68 tf tf, and 13 copepodids

from single colony, in 18 m, Bane de Touareg, Bay

of Ampasindava, near Nosy B6, Madagascar, 11

July 1967; 31 $ $, 37 0*0*. and 41 copepodids

from two colonies, in 12 m, west of the harbor at

Hellville, Nosy B£, Madagascar, 4 August 1967; and

38 9 9 , 33 o* cf, and 1 copepodid from single colony, in 20 m, west of Andilana, Nosy B£, Madagascar, 13° 18' S, 48°07' E, 24 August 1967.

Metaxymolgus venustus (Humes, 1959)

Lichomolgus venustus Humes, 1959, pp. 290-298, figs. 36-64.

From the nudibranch Phyllidia trilineata Cuvier, region

of Nosy Be, Madagascar.—Gotto, 1962, p. 104.

NEW HOST.—16 $9, 8 <? c? from five crinoids,

Cenometra emendatrix (Bell), in 20 m, Tany Kely,

near Nosy B£, Madagascar, 10 September 1963, collected by AGH.

Genus Monomolgus Humes and Frost, 1964

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six moderately long setae. Rostrum either not well developed or rounded posteroventrally. First antenna 7-segmented, in the female

with the formula 4, 13, 6, 3, 4 -f 1 aesthete, 2 + 1

aesthete, and 7 -+- 1 aesthete; in the male with 4,

13-J-2 aesthetes, 6, 3 + 1 aesthete, 4 -f 1 aesthete,

2 -+- 1 aesthete, and 7 -|- 1 aesthete. Second antenna 4-segmented, with the armature 1, 1, 3, and

I -f- a few minute elements, there being only a

single terminal claw.

Labrum indented medially. Mandible with the

basal region distal to the indentation with its convex side lacking a scalelike area with spinules but

having distally a serrated fringe and with its concave side bearing spinules; lash either short or long.

Paragnath a small lobe with setules or hairs. First

maxilla with three or four elements. Second maxilla of the usual lichomolgid type. Maxilliped 3 -

segmented in the female with a pointed tip; in the

male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having either 11,1,5 or 111,1,5.

Leg 4 endopod with the formula 0-0;I. Leg 1 endopod in the male with the last segment either 1,5 as

in the female or 1,1,4. Leg 5 with a free segment

bearing two setae. Leg 6 represented in the female

by the two small elements near the area of attachment of each egg sac; in the male by the ventral

posterolateral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Monomolgus unihastatus Humes

and Frost.

REMARKS.—The two species placed in the genus

Monomolgus show certain differences which may be

of generic value. These may be noted particularly

Key to Species of the Genus Monomolgus

FEMALES

Caudal ramus 2.63:1; third segment of leg 3 with formula 1,1,2; third segment of exopod of

leg 4 with 11,1,5 M . ptammocorae

Caudal ramus 3.9:1; third segment of leg 3 with 1,11,2; third segment of exopod of leg 4

with 111,1,5 M. unihastatus

MALES

Caudal ramus 2.6:1; without sexual dimorphism in formula of leg 1 endopod M. psammocorae

Caudal ramus 5.7:1; with sexual dimorphism in formula of leg 1 endopod (third segment

W4 M. unihastatus
NUMBER 127 249

in the mandible, the third segment of leg 4 exopod,

and the third segment of leg 1 endopod in the male.

In spite of these it seems best to regard the two

species as congeneric until more related species are

discovered to provide more information on the

weight of such characters.

Monomolgus unihastatus Humes and Frost, 1964

FIGURE 138

Monomolgus unihastatus Humes and Frost, 1964, pp. 132—

135, figs. 1-36 [from the hard coral Porites sp. cf. andrewsi Vaughan, region of Nosy Be4

, Madagascar].-Bouli

gand, 1966, p. 269-Humes and Ho, 1968a, p. 371 [from

Porites sp. cf. nigrescens Dana, Nosy Be\ Madagascar].

NEW HOST.—2 ? 9 , 4 cf cf from the alcyonacean

Parerythropodium fulvum (Forskll), in 1 m, Ambariobe, near Nosy B£, Madagascar, 15 January

1964, collected by AGH. (This is perhaps an accidental host.)

NEW RECORDS (collected by AGH).-^6 $ ? , 28

cf cf, and 20 copepodids from Porites sp., in 2 ra,

off Ampombilava, Nosy B£, Madagascar, 5 June

1967; 25 ? ° , 3 c?cf>

 an d

 1 copepodid from Porites sp., in 2 m, Ambariobe, near Nosy B£, Madagascar, 22 August 1967.

FIGURE 1$8.—Monomolgus unihastatus Humes and Frost. Female: a, dorsal; b, mandible; c,

leg 4. Male: d, dorsal. (From Humes and Frost, 1964, figs. 1, 12, 22, 25.) Length of female

1.30 mm, of male 1.24 mm.
250

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Monomolgus psammocorae Humes and Ho, 1967

Monomolgus psammocorae Humes and Ho, 1967b, pp. 9-14,

figs. 52-79 [from the hard coral Psammocora contigua

(Esper), Ambariobe, near Nosy Be, Madagascar].

Genus Nasomolgus Sewell, 1949

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum not well

developed. First antenna 7-segmented, in both

sexes with the formula 4, 13, 6, 3, 4 -f- 1 aesthete,

2 -f- 1 aesthete, and 7 -f 1 aesthete. Second antenna 4-segmented, the formula being 1, 1, 3, and

II -f- 5 small elements, there being two terminal

claws.

Labrum with the two lobes separated by an indentation and with two prominent, ventrally directed setae on its anterolateral areas. Mandible

with a slender, unornamented base carrying distally and at an angle to it a bipectinate blade.

Paragnath a small hairy lobe. First maxilla with

four elements. Second maxilla with the usual

lichomolgid form but with a very short lash.

Maxilliped in the female 3-segmented, slender,

with a greatly elongated and very slender third

segment; in the male 4-segmented (taking the

proximal part of the claw as a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod with

the formula 0-1 ;2, the seta on the first segment

leathered, those on the second segment naked or

with very short barbules. Leg 1 endopod of the

male with the same armature as in the female. Leg

5 with a minute free segment bearing two setae.

Leg 6 represented in the female by the two setae

near the area of attachment of each egg sac; in

the male by the ventral posterolateral flap on the

genital segment bearing two setae.

Other features as in the species below.

Associated with polychaete annelids.

TYPE-SPECIES.—Nasomolgus cristatus Sewell.

REMARKS.—The five s|>ecies of Nasomolgus constitute a well-unified group in respect to both their

morphological characters and their host preferences.

The males of N. cristatus and N. parvvhis are

unknown.

Key to Species of the Genus Nasomolgus

FEMALES

1. Third segment of endopod of leg 3 with formula 1,11,2 2

Third segment of endopod of leg 3 with formula 11,2 N. firmus

2. Caudal ramus 14.6:1 N. Uptut

Caudal ramus much shorter, not exceeding 3:1 3

3. Caudal ramus about 2.5:1 N. cristatus

Caudal ramus less than 1.5:1 4

4. Length (without ramal setae) 0.87 mm; greatest dimensions of genital segment 99 x 112 /x

N. rudis

Length (without ramal setae) 0.57 mm; greatest dimensions of genital segment 72 x 83 fi

N. parvutus

KNOWN MALES

1. Third segment of endopod of leg 3 with formula 1,115 2

Third segment of endopod of leg 3 with formula 115 N. firmus

2. Caudal ramus 8.4:1 N. leptus

Caudal ramus 1.3:1 ... N. rudis

Nasomolgus cristatus Sewell, 1949

Nasomolgus cristatus Sewell, 1949, pp. 126, 127, fig. 35a-e

[from coast of southern Arabia].—Humes and Ho, 1967c,

pp. 386, 397.

REMARKS.—The male is unknown.

Nasomolgus firmus Humes and Ho, 1967

FIGURE 139«-e

Nasomolgus firmus Humes and Ho, 1967c, pp. 386-390, 6gs.

60-86 [from the polychaete Sabellastarte magnified (Shaw),

region of Nosy Be\ Madagascar].
NUMBER 127 251

PIGURE \S9.—Nasomolgus firmus Humes and Ho. Female: a, dorsal; b, mandible; c, maxilliped;

d, leg 4. Male: e, dorsal. Nasomolgus leptus Humes and Ho. Female: /, rostral area and

labrum. (From Humes and Ho, 1967c. figs. 60, 68, 72. 76, 78, 91.) Length of N. firmus female

0.81 mm. male 0.58 mm.

Nasomolgus leptus Humes and Ho, 1967

FICURE 139/

Nasomolgus leptus Humes and Ho, 1967c, pp. 390-393, figs.

87-109 [from Sabellastarte magnifica, Nosy Be\ Madagascar].

Nasomolgus parvulus Humes and Ho, 1967

Nasomolgus parvulus Humes and Ho, 1967c, pp. 395-397,

figs. 136-142 [from Sabellastarte magnifica, region of Nosy

Be. Madagascar].

REMARKS.—The male is unknown.

Nasomolgus rudis Humes and Ho, 1967

Nasomolgus rudis Humes and Ho, 1967c, pp. 393-395, figs.

110-135 [from Sabellastarte magnifica, region of Nosy Be\

Madagascar].

Genus Octopicola Humes, 1957

DIAGNOSIS.—Body cyclopiform, but elongated and

slender. Urosome in both sexes 6-segmented.

Caudal ramus with six setae. Rostrum sub triangular

with a rounded tip. First antenna 7-segmented, in

both sexes with the armature 4, 13, 6, 3, 4 + 1

aesthete, 2 -f- 1 aesthete, and 7 -f 1 aesthete.

Second antenna 1-segmented, the formula being

1, 1, 3 (some of these elements being spines), and

IV -f- 3, there being four terminal claws.

Labrum with two long, slender lobes separated by

an indentation. Mandible with a strong, distally

directed tooth on the convex side of the base; blade

broad and attenuated, placed at an angle to the

base, with its concave edge spinulose or denticulated. Paragnath a small obscure lobe. First maxilla

with three elements. Second maxilla of the usual

lichomolgid type. Maxilliped in the female 3-segmented with a pointed tip; in the male 4-segmented

(assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 in both sexes with 3-segmented rami

except leg 4 endopod which is 1-segmented, armed

with three terminal setae (two with lateral spinules,

the third plumose). Leg 4 exopod with the third

segment having 11,1,5. No sexual dimorphism in the

formula of the legs. Leg 5 with a minute lobe

bearing two terminal elements and a seta nearby.

Leg 6 in the female represented by a spinule and a

seta near the genital openings; in the male by the

two seta on the genital flap.

Other features as in the species below.

Associated with octopuses.

TYPE-SPECIES.—Octopicola superbus Humes.
252 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Key to Species of the Genus Octopicola

FEMALES

1. Caudal rani us 7.6:1; seta adjacent to lobe of leg 5 inserted on a distinct, subrectangular

lobe O. itocki

Caudal ramus 9-10:1; seta adjacent to lobe of leg 5 inserted directly on body surface rather

than on a lobe 2

2. Fifth pedigerous segment with small but distinct tergal plates; longer of the two setae of

leg 6 short, reaching only slightly beyond genital segment O. superbus antiUnuU

Fifth pedigerous segment without tergal plates; longer of the two setae of leg 6 very long,

reaching to the end of the first postgcnital segment or beyond O. tuperbus superbus

MALES

1. Length 1.30 mm; outermost seta of leg 4 endopod showing sexual dimorphism, being

sinuous and spiniform O. ilotki

Length 156-1.90 mm; outermost seta of leg 4 endopod not sinuous and spiniform 2

2. Longer of the two setae of leg 6 short, less than one-fifth the length of the first postgenital

segment; third segment of second antenna with a prominent, densely spinuloae projection

... O. superbus antUUnsis

Longer of the two setae of leg 6 very long, reaching nearly to the end of the first postgenital

segment; third segment of second antenna with a short, finely denticulated triangular

process O. superbus superbus

Octopicola superbus superbus Humes, 1957

Octopicola superba Humes, 1957, pp. 2-7, pi. 1, figs. 1-12,

pi. 2, figs. 13-22 [from the eggs and skin of the cephalopod

Octopus vulgaris Lamarck, Banyuls, Mediterranean coast

of France].—Delamare Deboutteville, Humes, and Paris,

1957, pp. 504-506.

Octopicola superbus.-Boucquet and Stock, 1960, pp. 1-6,

figs. 1-4 [from Octopus vulgaris, Roscoff, Channel coast of

France; Banyuls sur-mcr, Mediterranean coast of France].—

Gotto, 1962, p. 101,-Laubier, 1966, p. 234 [from Gulf of

Marseille, France].

Octopicola superbus antillensis Stock, Humes, and

Gooding, 1963

Octopicola superbus antillensis Stock, Humes, and Gooding,

1963a, pp. 5-9, figs. 1, 2 [from Octopus vulgaris, Barbados,

Curasao].

NEW HOST—3 $ ?, 1 rf from Octopus briareus

Robson, Key Largo, Florida, 11 May 1969, collected

by E. S. McSweeny, University of Miami.

NEW RECORD.—4 9 $, 1 copepodid from Octopus

vulgaris, off Alligator Point, Franklin County,

Florida, 18 December 1965, collected by J. Rudloe.

Octopicola stocki Humes, 1963

FIGURE 140

Octopicola stocki Humes, 1963, pp. 271-279, figs. 1-30 [from

a cephalopod-originally identified as Octopus (Tritaxeopus) cornutus Owen but later determined by Dr. G. L.

Voss to be Octopus cyaneus Gray—at Noiy Re, Madagascar].

Genus Odontomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum without a

distinct posteroventral margin. First antenna 7-

segmented, with the armature in the female 4, 13,

6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 -|- 1

aesthete; in the male 4, 13 -|- 2, 6, 3 -f- 1 aesthete,

4 -f- 1 aesthete, 2 -(- 1 aesthete, and 7 -\- 1 aesthete.

Second antenna 4-segmented, with the formula

1, 1, 3, and I, there being a single terminal claw.

Labrum with the two lobes separated by a median

indentation. Mandible with the basal part having

on its convex side a pointed, proximally directed

tooth followed by a series of serrations and on its

concave side beyond the indentation two lobes with

spinules; lash long. Paragnath a small lobe. First

maxilla with three elements. Second maxilla of the

usual lichomolgid type. Maxilliped of the female

3-segmented with a pointed tip; in the male 4—

segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3 segmented rami except for leg

4 endopod which is 2-segmented, and with the
NUMBER 127 253

FIGURE 140.—Octopicola stocki Humes. Female: a, dorsal; b, second antenna; c, labrum; d,

mandible; e, leg S; f, leg 4; g, leg 5. Male: h, dorsal. (From Humes, 1963, figs. 1, 8-10, 20-22,

24.) Length of female 1.72 mm, of male 1.30 mm.

Key to Species of the Genus Odontomolgus

FEMALES

1. Second segment of second maxilla with a setiferous sphere; longest seta on caudal ramus

shorter than ramus O. actinophorus

Second segment of second maxilla without a setiferous sphere; longest seta on caudal ramus

much longer than ramus 2

2. Genital segment abruptly constricted in its posterior fourth; free segment of leg 5

elongated (90 p long) with a slight inner proximal expansion O. rhadmus

Genital segment pyriform, without a sharp constriction; free segment of leg 5 short (34 ft

long) without an inner proximal expansion O. camptdus

MALES

1. Longest seta on caudal ramus shorter than ramus; ornamentation of second segment of

second antenna consisting of only a few (2-4) conical protuberances O. actmopihonu

Longest seta on caudal ramus much longer than ramus; ornamentation of second segment

of second antenna composed of many spinules or spinelike bosses 2

2. Second segment of second antenna with an inner row of spinules; two setae on free segment

of leg 5 in the ratio of 3:1; genital segment longer than wide O. rhadmus

Second segment of second antenna with many short spinelike bosses; two setae on free

segment of leg 5 in the ratio of about 3:2; genital segment wider than long ... O. campulus
254

usual lichomolgidiform armature. Leg 4 exopod

with the last segment 11,1,5. Leg 4 endopod 0-1 ;II,

the seta on the first segment being feathered. In

the male of O. actinophorus and of O. rhadinus,

leg 1 endopod with the third segment having the

formula 1,1,4 rather than 1,5 as in the female; in

O. campulus no sexual dimorphism in the formula

of leg 1. Leg 5 in both sexes with a free segment

bearing two terminal setae. Leg 6 represented in

the female by the two small setae near the area of

attachment of each egg sac, in the male by a

posteroventral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES. — Odontomolgus actinophorus

(Humes and Frost).

ETYMOLOGY.—The name is a combination of the

Greek words ooou? (a tooth), alluding to the toothSMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

like process on the mandible, and . Gender

masculine.

Odontomolgus actinophorus (Humes and Frost,

1964)

FIGURE 141

Lichomolgus actinophorus Humes and Frost. 1964, pp. 135-

137, figs. 37-65 [from the hard corals Pavona angulata

Klunzinger and Pavona cactus (Forskil), region of Nosy

\U, Madagascar].—Bouligand, 1966, p. 269.—Humes and

Ho, 1968a, p. 371 [from Pavona danai (Milne Edward*

and Haime), Pavona danai or Pavona angularis (Klunzinger), and Pavona ? venusta (Dana), region of Nosy B6,

Madagascar].

Odontomolgus campulus (Humes and Ho, 1968)

Lichomolgus campulus Humes and Ho, 1968a, pp. 354—357,

figs. 1-24 [from the hard coral Alvtopora »p., Nosy lie.

Madagascar].

FIGURE 141.-Odontomolgus actinophorus (Humes and Frost). Female: a, dorsal; b, second antenna; c, mandible; d, leg 4 and intercoxal plate. Male: e, dorsal. (From Humes and Frost,

1964, figs. 37, 44, 46, 56, 58.) Length of female 1.42 mm, of male 128 mm.
NUMBER 127 255

NEW HOST.—9 9 9, 3 cT cf from the hard coral

Goniopora sp., in 3 m, Nosy N'Tangam, Nosy Be",

Madagascar, 24 June 1967, collected by AGH.

Odontomolgus rhadinus (Humes and Ho, 1967)

Lichomolgus rhadinus Humes and Ho, 1967b, pp. 5-9, figs.

25-51 [from the madreporarian Psammocora contigua

(Esper), Nosy Be, Madagascar].

NEW HOST.—Many specimens from Pavona sp.,

intertidal to 10 cm, Nosy Iranja, southwest of Nosy

Be, Madagascar, 9 August 1967, collected by AGH.

Genus Panjakus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, in the

female with the formula 4, 13, 6, 3, 4 -f- 1 aesthete,

2 -f 1 aesthete, and 7 -f 1 aesthete; in the male

with 4, 13 + 2 aesthetes, 6, 3 -f 1 aesthete, 4 -j- 1

aesthete, 2 + 1 aesthete, and 7 -f- 1 aesthete. Second

antenna 4-segmented, with the armature 1, 1, 3,

and I, there being a single terminal claw.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal area distal to the

indentation having on the convex side a short,

distally directed digitiform process followed by a

serrated fringe and on its concave side two lobes

each with a row of spinules; lash moderately long.

Paragnath a small hairy lobe. First maxilla with

four elements. Second maxilla of the usual licho

molgid type, but with a large stout digitiform

process on the first segment. Maxilliped of the female 3-segmented, with a pointed tip; in the male

4-segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform type. Leg 4 exopod with the

third segment having 11,1,5 or 111,1,5. Leg 4 endopod with 0-1 ;II, the seta on the first segment being

feathered. In the male the third segment of leg 1

endopod with 1,1,4 instead of 1,5 as in the female.

Leg 5 in both sexes with a free segment bearing

two terminal setae. Leg 6 in the female represented

by two setae and a small spiniform process, in the

male by a posteroventral flap on the genital segment bearing two setae.

Other features as in the species described below.

Associated with madreporarian corals.

TVPE-SPECIES. — Panjakus hydnophorae, new

species.

ETYMOLOGY.—The generic name is a Latinized

form of the Malgache word "mpanjaka," meaning

chief or king. Gender masculine.

REMARKS.—The genus Panjakus bears a certain

relationship to the genera Andrianellus and Rakotoa mentioned elsewhere in this work, both of

which have strong similarities (the 4-segmented

second antenna with a single terminal claw, the

mandible with two spinulose lobes on the concave

side, and the second maxilla with a large, prominent toothlike process on the first segment). Panjakus differs from Andrianellus, which has the

formula 0-1 ;I on the endopod of leg 4 and which

shows no sexual dimorphism in leg 1. Panjakus may

be distinguished from Rakotoa where the endopod

of leg 4 is reduced to an unarmed lobe, where the

armature of the third segment of the exopod of leg

4 is 1,3, and where in the male the second antenna

and leg 1 do not show sexual dimorphism.

Key to Species of the Genus Panjakus

FEMALES

Free segment of leg 5 long, more than half as long as the genital segment; leg 4 exopod with

third segment having IIIJ3 **• hydnophorae

Free segment of leg 5 short, only about one-ninth the length of the genital segment; leg 4

exopod with third segment having 11.1,5 P. platygyrae

MALES

Second segment of second antenna ornamented with knobs; leg 4 endopod with third segment

having 111,1,5 P- hydnophorae

Second segment of second antenna smooth; leg 4 endopod with third segment having 11,1,5

P. platygyrae
256

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Panjakus hydnophorae, new species

FIGURES 142-144

TYPE MATERIAL.—59 9 9, 30 cf cf from the

madreporarian coral Hydnophora sp., in 2 m, Ambariotelo, a small island almost between Nosy

Komba and Nosy Be, northwestern Madagascar,

20 July 1967, collected by AGH. Holotype $, allotype, and 83 paratypes (56 $ $, 27 cf cf) deposited

in USNM, the remaining paratypes (dissected) in

the collection of AGH.

OTHER MATERIAL EXAMINED.—From Hydnophora

sp.: 18 ? ?, 7 d1 cf. in 2 m, Tany Kely, a small

island south of Nosy Be, 21 May 1967; and 24 9 $,

14 cf cf. in 2 m, off Ampombilava, Nosy B£, 5 June

1967. All collected by AGH.

From Hydnophora ? exesa (Pallas): 4 9 9, 3

cf cf» in 1 m, west of Pte. de Tafondro, Nosy Be, 2

October 1963.

From Hydnophora tenella Quelch: 8 9 9, 14

cf cf # and 3 copepodids, in 3 m, opposite Ambariotsimaramara, southern shore of Nosy Bi, 18 October

1963.

FEMALE.—The body (Figure 142a,6) is moderately slender, with the prosome a little thickened

in lateral view. The length (excluding the setae

on the caudal rami) is 1.34 mm (1.23-1.42 mm) and

the greatest width 0.51 mm (0.47-0.55 mm), based

on ten specimens in lactic acid. The segment of

leg 1 is separated from the cephalosome by a dorsal

transverse furrow. The epimeral areas of the pedigerous segments are formed as illustrated in the

figures. The ratio of the length to the width of the

prosome is 1.50:1. The ratio of the length of the

prosome to that of the urosome is 1.14:1.

The segment of leg 5 (Figure 142c) is 57 x 260 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 156 x 208 n, wider than long, broadest

in its anterior half and tapering posteriorly. The

areas of attachment of the egg sacs are located

dorsolaterally, just in front of the middle of the

segment. Each area (Figure 142d) bears two small

setae about 7 n (one sharply pointed and the other

with a hyaline tip often broken off) and a small

spiniform process. The three postgenital segments

are 94 x 107 n, 83 x 94 n, and 88 x 91 M. from anterior to posterior. The posterior border of the

anal segment bears on each side a row of small

spinules.

The caudal ramus (Figure 142e) is greatly elongated, 212-234 x 26 n, the width taken at the middle, about 8.58 times longer than wide. The longest

terminal seta is not much longer than the ramus.

The outer lateral seta is 91 n, the dorsal seta 55 n

and displaced toward the inner margin of the

ramus, the outermost terminal seta 72 »i, the innermost terminal seta 88 n, and the two median

terminal setae 185 n (outer) and 255 n (inner),

both inserted a little ventrally on the ramus. All

six setae are naked.

The body surface bears a few hairs (sensilla) as

indicated in Figure 142aA

The egg sac (Figure 142a) is oval, 330 x 230 i*.

contains a few large eggs (eight in the specimen

drawn), and reaches only to the last postgenital

segment.

The rostrum (Figure 142/) is a linguiform lobe.

Posterior to the rostrum there is a ventrally raised

area (shown also in Figure 1426).

The first antenna (Figure 142g) is slender, 7-

segmented, and 390 n long. The lengths of the

segments (measured along their posterior nonsetiferous margins) are 31 (55 n along the anterior

margin), 96, 33, 58, 64, 45, and 39 n respectively.

The formula for the armature is 4, IS, 6, 3, 4 -|- 1

aesthete, 2 -f 1 aesthete, and 7 -f 1 aesthete. All

the setae are naked.

The second antenna (Figure \42h) is 4—segmented. The first and second segments bear a small

inner seta. The third segment bears three setae, one

much shorter than the other two. The fourth segment, 52 (i along its outer edge, 20 n along its inner

edge, and 23 n wide, has a single terminal claw

33 n along its axis.

The labrum (Figure 143a) has two broad posteroventral lobes with wide, hyaline membranous

margins.

The mandible (Figure 1436) has on its concave

margin a shallow indentation followed by two

lobes, each with a marginal row of spinules, and

on its convex margin a short, distally directed digitiform process followed by a serrated hyaline fringe.

The lash is moderately long with short lateral

spinules. The paragnath (Figure 143c) is a small,

elongated lobe with distal hairs. The first maxilla

(Figure 143c) has four elements. The second maxilla

(Figure 143d) is 2-segmented. The first segment is

unarmed but has a large stout digitiform process

extending ventrally (see Figure 1426). The second
NUMBER 1X7

Ficuu 142.—Panjakus hydnophorae, new species. Female: a, dorsal (A) ; b, lateral (B) ; c,

urosome, dorsal (c); d, area of attachment of egg sac, dorsal (D) ; e, caudal ramus, dorsal (E) ;

/, rostrum, ventral (E) ; g, first antenna, with arrows indicating positions of additional aesthetes

in male, anterodorsal (E) ; h, second antenna, inner (F) . Scale: A, B, 0.5 mm; c, 0.2 mm; D,

0.05 mm; E, T, 0.1 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

b d

g

FIGURE 143.—Panjakus hydnophorae, new species. Female: a, labrum, ventral (F) ; b, mandible,

posterior (c); c, first maxilla and paragnath, posterior (c); d, second maxilla, posterior (F) ;

e, maxilliped, anterior (c); /, area between maxillipeds and first pair of legs, ventral (E) ; g,

leg 1 and intercoxal plate, anterior (E) ; h, leg 2 and intercoxal plate, anterior (E) ; i, third

segment of endopod of leg 3, anterior (E) . Scale: E, F, 0.1 mm; c, 0.05 mm.
NUMBER 127 259

segment bears two setae (one naked, the other

with a pectinate fringe) and a slender setule; the

terminal lash bears a row of spines. The maxilliped

(Figure 143e) is 3-segmented. The first segment is

unarmed. The second segment bears two short,

naked setae. The third segment bears a naked seta

and a naked spine and terminates in a spiniform

process with a few spinules.

The ventral area between the maxillipeds and

the first pair of legs (Figure 143/) protrudes a little

ventrally (Figure 1426). A line connects the bases

of the maxillipeds.

Legs 1-4 (Figures 143g-i, 144a) have 3-segmented

rami except for the endopod of leg 4 which is 2-

segmented. The spinal and setal formula is as

follows:

l\ coxa 0-1 batii 1-0 exp 1-0; 1-1; 111,1,4

enp 0-1; 0-1; 1.5

P. coxa 0-1 baits 1-0 cxp 1-0; 1-1; III.U

enp 0-1; 0-2; 1,11,3

P. coxa 0-1 bam 1-0 cxp Hi; 1-1; 111,13

enp 0-1; 0-2; 1.11.2

P« coxa 0-1 basis 1-0 exp 1-0; 1-1; 111,1,5

enp 0-1; II

The inner coxal seta of legs 1-3 is long and

plumose, but in leg 4 that seta is minute (7n) and

naked. The inner margin of the basis is haired in

all four legs. The exopod of leg 4 is 146 n long.

The first segment of the endopod is 33 x 31 n (the

length not including the spinous processes), with

its inner distal plumose seta 102 n. The second segment is 54 x 23 n in greatest dimensions, including

the terminal spiniform processes; its two terminal

spines are 28 \t (outer) and 77 n (inner). Both endopod segments have outer marginal hairs.

Leg 5 (Figure 144&,c) has a free segment 96 x 34

n in greatest dimensions when seen in lateral

(outer) view. The two terminal setae are 50 n and

60 n, and the adjacent seta on the body is 105 n. All

three setae are naked. The dorsal and outer surfaces of the distal half of the ramus are ornamented

with minute spinules.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure I42d).

The color in life in transmitted light is brownish

opaque, with a few small reddish brown dots scattered on the dorsal surfaces of the prosome, the

eye red, the egg sacs gray.

MALE.—The body (Figure 144d) is slender. The

length is 1.29 mm (1.24-1.33 mm) and the greatest

width 0.42 mm (0.40-0.44 mm), based on ten specimens in lactic acid. The line of separation between the cephalosome and the segment of leg 1 is

more weakly developed than in the female. The

ratio of the length to the width of the prosome is

1.55:1. The ratio of the length of the prosome to

that of the urosome is 1:1.06, the urosome being

slightly longer than the prosome.

The segment of leg 5 (Figure 144*) is 42 x 195 i*.

There is no ventral intersegmental sclerite. The

genital segment is 234 x 234 n, in dorsal view with

its lateral edges subparallel. The four postgenital

segments are 52 x 95 n, 62 x 83 n, 60 x 74 n, and 70

x 78 n from anterior to posterior.

The caudal ramus resembles that of the female

but is a little more slender, 208 x 21 n, or 9.91 times

longer than wide.

The body surface has a few hairs as in the female.

The rostrum is like that of the female.

The first antenna is similar to that of the female

but has three additional long aesthetes (at points

indicated by arrows in Figure 142g), so that the

formula is 4, 13 -{- 2 aesthetes, 6, 3 -|- 1 aesthete,

4 -f 1 aesthete, 2 -j- 1 aesthete, and 7 -f 1 aesthete.

The second antenna (Figure 144/) resembles that

of the female but the inner surface of the second

segment has numerous small, rounded knobs.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 144g) is 4-segmented (assuming that the proximal part of the claw represents a fourth segment). The first segment is

unarmed but has a small postero-inner distal digitiform lobe. The second segment bears two naked

setae and two rows of spines. The small third segment is unarmed. The claw is 268 n along its axis,

including the large terminal lamella, and is only

slightly recurved. It shows an incomplete division

about midway and bears two proximal unequal

naked setae.

The ventral area between the maxillipeds and

the first pair of legs is like that in the female.

Legs 1-4 are segmented as in the female and have

the same armature as in that sex except for the

third endopod segment of leg 1 (Figure 144/i) which

has the formula 1,1,4. These two spines are 27 n

(outer) and 57 >* (inner).

Leg 5 (Figure 144i) has an unornamented free

segment 42 x 13 n, with the terminal setae 35 n

and 50 n.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 144.—Panjakus hydnophorae, new species. Female: a, leg 4 and intercoxal plate, anterior

(E) ; b, leg 5, dorsal (F) ; c, free segment of leg 5, lateral (outer) (F) . Male: d, dorsal (A) ; e,

urosome, dorsal (c); /, second antenna, outer (F) ; g, maxilliped, inner (H) ; h, endopod of

leg 1, anterior (E) ; i, leg 5, dorsal (c); ;, leg 6, ventral (E) . Scale: A, OS mm; c, H, 02 mm;

E, T, 0.1 mm; c, 0.05 mm.
NUMBER 187 261

Leg 6 (Figure 144;") is a posteroventral flap on

the genital segment bearing two slender naked

setae 50 n and 60 n.

The spermatophore was not observed.

The color in life in transmitted light resembles

that of the female but the body tends to be less

brownish and the prosomal dots are smaller.

ETYMOLOGY.—The specific name hydnophorae is

formed from the generic name of the host coral.

Panjakus platygyrae, new species

Ficuus 145. 146

TYPE MATERIAL.—6 99 , 4 0*0* from single

colony of the madreporarian coral Platygyra ?

lamellina (Ehrenberg), in 2 m, Ambariotelo, a small

island almost between Nosy Komba and Nosy B£,

northwestern Madagascar, 23 May 1964, collected

by AGH. Holotype 9 . allotype, and 4 paratypes (3

9 9 . 1 o*) deposited in USNM, the remaining four

paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED (all collected by

AGH).—From Platygyra daedala (Ellis and Solander): 2 9 $. 2 0*0" from single colony, in 4 m,

Pte. Lokobe, Nosy Be, 24 December 1963; and 2

cf o*, 1 copepodid, from single colony, in 2 m, Pte.

a la Fievre, Nosy Be, 26 December 1963.

From Platygyra sp. cf. daedala (Ellis and Solander): 4 9 $» 1 cf from single colony, in 1 m,

Ambariobe, a small island almost between Nosy

Komba and Nosy Be\ 16 January 1964.

FEMALE.—The body (Figure 145a) is moderately

slender. The prosome in lateral view is a little

thickened as in Panjakus hydnophorae. The length

(not including the setae on the caudal rami) is

1.45 mm (1.37-1.53 mm) and the greatest width

0.57 mm (0.55-0.59 mm), based on six specimens

in lactic acid. The segment of leg 1 is completely

fused with the cephalosome. The epimeral regions

of the pedigerous segments are formed as shown in

the figure. The ratio of the length to the width of

the prosome is 1.29:1. The ratio of the length of

the prosome to that of the urosome is 1.04:1.

The segment of leg 5 (Figure 1456) is 65 x 210 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 208 x 215 n, broadest just in front of

the middle of the segment and tapering posteriorly.

The areas of attachment of the egg sacs are situated

dorsally near the middle of the segment. Each area

(Figure 145c) bears two naked setae 10 n long (the

posterior one peculiar in having a sclerotized proximal part and a hyaline, blunt distal part) with a

small spinous process between them. The three

postgenital segments are 78 x 96 n, 57 x 78 \i, and

101 x 68 n from anterior to posterior. The posteroventral border of the anal segment bears a row of

minute spinules on each side.

The caudal ramus (Figure 145d) is greatly

elongated, 140 x 18 n, or 7.8 times longer than

wide. The longest terminal seta is shorter than the

ramus. The outer lateral seta is 44 n, the dorsal

seta 27 n, the outermost terminal seta 52 n, the

innermost terminal seta 63 n, and the two median

terminal setae 88 v- (outer) and 97 n (inner). All

six setae are naked.

The body surface bears a few small hairs (sensilla) as indicated in the figures.

The egg sac is unknown. One female, however,

had three large eggs attached to the genital segment, each egg about 130 »i in diameter.

The rostrum is similar to that of P. hydnophorae.

The first antenna (Figure 145^), 389 n long,

resembles that of P. hydnophorae. The lengths of

the seven segments (measured along their posterior

nonsetiferous margins) are 24 (55 \i along the

anterior margin), 108, 44, 54, 58, 42, and 28 »

respectively. The formula for the armature is the

same as in P. hydnophorae. All the setae are naked.

The second antenna (Figure 145/) is similar to

that of P. hydnophorae. The fourth segment, 38 n

along its outer edge, 21 n along its inner edge, and

22 i* wide, bears a single claw 33 n along its axis.

The labrum is like that of P. hydnophorae.

The mandible resembles closely that of P. hyrnophorae, the only apparent difference being that the

short digitiform process on the convex side is

a little more slender than in that species. The

paragnath is similar to that in P. hydnophorae. The

first maxilla (Figure 145g) bears four elements. The

second maxilla (Figure 145/i) has a large digitiform

process on the first segment, and the dorsal (inner)

seta on the second segment has a broad spinulose

flap. The maxilliped (Figure 145») is more elongated than in P. hydnophorae but has the same

number of elements as in that species.

The ventral area between the maxillipeds and

the first pair of legs (Figure 145;) protrudes ventrally. A line connects the bases of the maxillipeds.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 145.—Panjakus platygyrae, new species. Female: a, dorsal (A) ; 6, urosome, dorsal (•);

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, first antenna,

anterodorsal (E) ; /, second antenna, postero-inner (D) ; g, first maxilla, posterior (E) ; h, second

maxilla, anterior (F) ; i, maxilliped, posterior (F) ; ;, area between maxillipeds and first pair

of legs, ventral (E) . Scale: A, 03 mm; B, 0.2 mm; c, F, 0.05 mm; D, E, 0.1 mm.
NUMBER 127

FrcutE 146.—Panjakus platygyrae, new species. Female: a, leg 1 and intercoxal plate, anterior

(E) ; b, leg 2 and intercoxal plate, anterior (E) ; c, third segment of endopod of leg 3, anterior

(D) ; d, leg 4 and intercoxal plate, anterior (E) ; e, leg 5, dorsal (c). Male: /, dorsal (A) ; g,

urosorae, dorsal (•); h, maxilliped, outer (E) ; i, endopod of leg 1, anterior (D) ; j , leg 6,

ventral (o). Scale: A, 0.5 mm; B, 0.2 mm; c, 0.05 mm; D, E, 0.1 mm.
264 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Legs 1-4 (Figure 146a-d) are segmented and

armed as in P. hydnophorae, except that the third

segment of the exopod of leg 4 has the armature

11,1,5. The inner coxal seta is naked and much

smaller (12 n) in leg 4 than in legs 1-3. The inner

margin of the basis bears a row of hairs in all

four legs. The exopod of leg 4 is 125 n long. The

first segment of the endopod is 32 x 20 v- (the length

not including the spinous processes), with its inner

distal plumose seta 55 n. The second segment is

57 x 19 ^ (including the terminal spiniform processes); its two terminal spines are 17 n (outer) and

23 n (inner). Both endopod segments have outer

marginal hairs.

Leg 5 (Figure \46e) has a small, unornamented

free segment 22 x 14 n. The two terminal setae are

31 n and 25 n, and the adjacent seta on the body

is 25 i*. All three setae are naked.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 145c).

The color in life in transmitted light is opaque,

the intestine slightly yellowish, the eye dark red.

MALE.—The body (Figure 146/) resembles in

general form that of the female. The length is 1.31

mm (1.26-1.36 mm) and the greatest width 0.47

mm (0.46-0.47 mm), based on four specimens in

lactic acid. The ratio of the length to the width of

the prosome is 1.38:1. The ratio of the length of the

prosome to that of the urosome is 1:1.04, the

urosome being slightly longer than the prosome.

The segment of leg 5 (Figure 146g) is 44 x 166 n.

There is no ventral intersegmental sclerite. The

genital segment is 234 x 237 n, in dorsal view with

slightly rounded lateral edges. The four postgenital

segments are 49 x 88 n, 55 x 74 n, 14 x 62 n, and

87 x 60 n from anterior to posterior.

The caudal ramus is like that of the female.

The rostrum resembles that of the female.

The first antenna (350 n long) is similar to that

of the female but there are three additional long

aesthetes (the longest one 200 v) as in the male

of P. hydnophorae, so that the formula is the same

as in that species.

The second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla resemble

those of the female. The maxilliped (Figure I46h)

has a small postero-inner distal knob on the first

segment. The second segment bears two naked setae,

two rows of spines, and a small spinulose crest.

The small third segment is unarmed. The daw is

200 n along its axis, including the terminal lamella,

and is only slightly recurved. It is partially divided

about midway and bears proximally two unequal

setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented as in the female and have

the same spine and setal formula except for the

third endopod segment of leg 1 (Figure 146i) which

has the formula 1,1,4. These two spines are 20 n

(outer) and 31 n (inner).

Leg 5 resembles that of the female, with its free

segment being 18 x 10 n.

Leg 6 (Figure 146;) is a posterovrntral flap on

the genital segment bearing two slender naked setae,

17 n and 24 n.

The spermatophore was not observed.

The color in life resembles that of the female.

ETYMOLOCY.—The specific name platygyrae is

derived from the generic name of the host coral.

REMARKS.—Panjakus platygyrae differs from P.

hydnophorae chiefly in the much smaller leg 5 in

the female, the formula 11,1,5 on the last exopod

segment of leg 4, the longest seta on the caudal

ramus being shorter than the ramus, the male genital segment in dorsal view having slightly rounded

lateral edges, and the male second antenna not

showing sexual dimorphism.

Genus Paradoridicola Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum rounded.

First antenna 7-segmented, in the female with the

armature 4, 13, 6, 3, 4 -}- 1 aesthete, 2 + 1 aesthete,

and 7 -j- 1 aesthete (so far as known); in the male

4, 13 -f 2 aesthetes, 6, 3 + 1 aesthete, 4 -j- 1

aesthete, 2+ 1 aesthete, and 7 -f- 1 aesthete. Second

antenna 4-segmented, the formula 1, 1, 3, and one

terminal claw -j- several small elements.

Labrum incised medially. Mandible with the

basal region beyond the indentation having on its

convex side a scalelike area with spinules followed

by a serrated fringe and on its concave side a row

of spinules; lash long. Paragnath a small hairy

lobe. First maxilla with three or four elements.

Second maxilla of the usual lichomolgid type.

Maxilliped in the female 3-segmented with a

pointed tip, in the male 4-segmented (assuming
NUMBER 127 265

that the proximal part of the daw represents a

foUrth segment).

Legs 1-4 with 3-segmented rami except leg 4

endopod which is 2-segmented. Armature of the

typical lichomolgidiform type. Leg 4 exopod with

the third segment having 111,1,5. Leg 4 endopod

with the formula 0-1; 11, the seta being feathered

or with short spinules. Leg 1 endopod of the male

with the third segment having the formula 1,1,4

instead of 1,5 as in the female (male unknown in

P. robustus). Leg 5 with a free segment bearing two

terminal elements. Leg 6 represented in both sexes

by the two setae near the genital openings.

Other features as in the species below.

Associated with octocorals.

TYPE-SPECIES.—Paradoridicola squamiger (Humes

and Frost).

ETYMOLOGY.—The generic name is formed from

«aoa (beside) and Doridicola. Gender masculine.

REMARKS.—A key to the males of Paradoridicola

has not been prepared because the subtle interspecific differences are virtually impossible to use

satisfactorily in a key. Reference should be made

to the original descriptions and figures. The male

of P. robustus is unknown.

Key to Species of the Genus Paradoridicola

FEMALES

1. Caudal ramui with ratio 3.4:1 P. glabrip**

Caudal ramus with ratio about 1.5:1 or quadrate 2

2. Caudal ramui with ratio about 13:1 S

Caudal ramus quadrate 4

3. Genital segment in dorsal view gradually constricted posterior to genital openings

P. robustus

Genital segment in dorsal view abruptly constricted behind genital openings where sides

are parallel P. adelphus

4. Free segment of leg 5 broad and subtriangular P. triquetrut

Free segment of leg 5 not broad and triangular 5

5. Free segment of leg 5 with ratio 2.4:1. distinctly shorter than genital segment ... P. squamiger

Free segment of leg 5 with ratio 6.7:1, much longer than genital segment P. timtlariae

Paradoridicola squamiger (Humes and Frost,

1964)

IAchomolgus squamiger Humes and Frost, 1964, pp. 145-148,

figs. 167-202 [from the alcyonacean Sinularia polydactyla

(Ehrenberg), region of Nosy Be, Madagascar].—Humes and

Ho, 1968c, pp. 689-690 [from Sinularia whiteleggei Liittschwager, Nosy Be, Madagascar].-Bouligand, 1966, p. 269.

NEW RECORD.—From Sinularia polydactyla

(Ehrenberg): 9 9 9 , 26 tf tf, and 22 copepodids, in

15 m, Bane de la Lanterne, Bay of Tsimipaika, near

Nosy B£, Madagascar, 26 July 1967, collected by

AGH.

Paradoridicola adelphus (Humes and Ho, 1968)

IAchomolgus adelphus Humes and Ho, 1968c, pp. 650-655,

figs. 51-66 [from the alcyonaceans Sinularia whiteleggei

Liittschwager, Sinularia pedunculata Tixier-Durivault, and

Sinularia polydactyla (Ehrenberg), region of Nosy Be,

Madagascar].

Paradoridicola glabripes (Humes and Ho, 1968)

IAchomolgus glabripes Humes and Ho, 1968d, pp. 707-713,

figs. 49-68 [from the alcyonacean Xenia umbellata Lamarck, Nosy Be, Madagascar].

NEW HOSTS.—From Xenia macrospiculata Gohar:

7 9 9. 7 cf o">

 an d 12 copepodids, in 20 m, Bane

de Cinq Metres, near Nosy Be, Madagascar, 6

August 1967, collected by AGH.

From Xenia viridis Schenk: 73 9 9 , 76 <$ cf» * nd

58 copepodids, intertidal, Navetsy, Nosy B6, Madagascar, 7 August 1967, collected by AGH.

Paradoridicola robustus (Thompson and A. Scott,

1903)

IAchomolgus robustus Thompson and A. Scott, 1903, p. 280,

pi. 16, figs. 14-20 [from washings of dredged invertebrates,

Ceylon].

REMARKS.—The male is unknown.
266

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Paradoridicola similariae, new species

FICURES 147-149

TYPE MATERIAL.—23 $ ? , 12 dd> an d 22 copepodids from ten colonies of the alcyonacean Sinularia arborea Verseveldt, in 13 m, opposite Antsiabe,

on the southern shore of Nosy Komba, near Nosy

Be\ northwestern Madagascar, 2 September 1967,

collected by AGH. Holotype 9 , allotype, and 29

paratypes (20 9 9, 9 <$cf) deposited in USNM,

the remaining paratypes (dissected) and the copepodids in the collection of AGH.

OTHER MATERIAL EXAMINED (all from Sinularia

arborea).—14 9 9> 9 cf cf • an<^ H copepodids from

single colony, in 2 m, Pte. Lokobe, Nosy Be, 3 June

1967; 30 9 9 , 31 cf o \ and 29 copepodids from

single colony, in 23 m, Tany Kely, a small island

south of Nosy Be, 30 June 1967; 3 9 9, 3 cf cf

from single colony, in 2 m, off Ampombilava, Nosy

Be, 7 July 1967; 12 9 9, 14 <$ <?, and 6 copepodids

from single colony, in 12 m, west of harbor at

Hellville, Nosy Be, 4 August 1967; and 3 9 9 ,

11 cT, and 6 copepodids from six small colonies,

in 13 m, opposite Antsiabe, Nosy Komba, 2 September 1967. All collected by AGH.

FEMALE.—The body (Figure 147a) has a moderately broad prosome. The length (not including the

ramal setae) is 1.33 mm (1.23—1.46 mm) and the

greatest width is 0.56 mm (0.52-0.62 mm), based

on ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.62:1. The

ratio of the length of the prosome to that of the

urosome is 2.12:1. The segment of leg 1 is almost

completely fused with the cephalosome. The epimera of the segments of legs 2-4 are extended

laterally as illustrated in the figure.

The segment of leg 5 (Figure 1476) is 104 x 205 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 179 x 161 ji, in dorsal view broadest in

its anterior two-thirds with rounded margins. The

areas of attachment of the egg sacs are situated

dorsolaterally just in front of the middle of the

segment. Each area (Figure 147c) bears two naked

setae, 19 n and 7 n, with a small spiniform process

between them. The three postgenital segments are

81 x 95 n, 65 x 84 n, and 57 x 78 n from anterior

to posterior. The posteroventral borders of the

genital segment and the first two postgenital segments bear broad hyaline spinules without articulations. The posteroventral border of the anal

segment is smooth.

The caudal ramus (Figure 147d) is quadrate,

34 x 34 \i. The outer lateral seta is 96 »*, the dorsal

seta 66 u, the outermost terminal seta 130 n, the

innermost terminal seta 330 n, and the two long

median terminal setae 400 n (outer) and 550 n

(inner), both inserted between dorsal (unornamented) and ventral (with a row of minute spinules) flaps. All six setae are naked. The dorsal

surface of the ramus bears a few hairs.

The body surface has a few hairs (sensilla) as

indicated in Figure 1470,6.

In all ovigerous females collected the egg sacs

were incomplete. The general shape, however, is

elongated and oval, with numerous eggs about 47 n

in diameter.

The rostrum (Figure 147r) is broadly rounded

posteroventrally.

The first antenna (Figure 147/) is slender, 506 u

long, with the elongated second and fourth segments about equal in length. The lengths of the

seven segments (measured along their posterior

nonsetiferous margins) are 50 (88 u along the

anterior margin), 141, 38, 139, 44, 30, and 26 H.

The proximal part of the fourth segment is much

more slender (18 M> wide) than the corresponding

part of the second segment (30 |i wide). The

formula for the armature is 4, 13, 6, 3, 4 -f- 1

aesthete, 2 -f 1 aesthete, and 7 -+- 1 aesthete. All the

setae are naked.

The second antenna (Figure 147g) is elongated.

The first two segments bear a small inner seta; the

third segment bears three inner setae. The elongated fourth segment, 107 n along its outer edge, 78

n along its inner edge, and 17 n wide, bears terminally a slender claw 60 n along its axis and six small

hyaline elements. All the setae are naked.

The labrum (Figure 147A), mandible (Figure

147i), paragnath (Figure 147A), first maxilla (Figure 148a), second maxilla (Figure 1486), and maxilliped (Figure 148c) resemble closely those of P.

squamiger (Humes and Frost), differing only in

very small details.

The ventral area between the maxillipeds and

the first pair of legs (Figure 148d) is only slightly

protuberant, with a fine line connecting the bases

of the maxillipeds.

Legs 1-4 (Figure \4Se-h) are similar to those of

P. squamiger, with the same segmentation and
FIGURE 147.—Paradoridicola sinulariae, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ;

c, area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum, ventral

(B) ; /, first antenna, with arrows indicating positions of additional aesthetes in male, anterodorsal (B) ; g, second antenna, postero-inner (E) ; h, labrum, with paragnaths indicated by

broken lines, ventral (F) ; i. mandible, posterior (c). Scale: A, 1.0 mm; B, 02 mm; c, D, 0.05

mm; E, F, 0.1 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE \49>.—Paradoridicola sinulariae, new species. Female: A, first maxilla, anterior (c); b,

second maxilla, posterior (F) ; c, maxilliped, anterior (F) ; d, area between maxillipeds and first

pair of legs, ventral (c); e, leg 1 and intercoxal plate, anterior (c); /, leg 2, anterior (c); g,

endopod of leg 3, anterior (c); h, leg 4 and intercoxal plate, anterior (c) ; i, leg 5, dorsal (c).

Scale: c, 0.05 mm; r, 0.1 mm; c, 02 mm.
NUMBER 127 269

armature. The inner coxal seta of leg 4 is 65 n

and plumose. The exopod of leg 4 is 185 n, with the

third segment having the formula 111,1,5. The first

segment of the endopod is 52 x 31 n (not including

the spinous processes), with its inner distal plumose

seta 112 i*. The long second segment is 159 x 15.5 |i

(least width), including the spiniform processes.

The two terminal barbed spines are 39 n (outer)

and 78 p (inner). Both segments have an outer

row of hairs.

Leg 5 (Figure 148i) has an elongated arched

free segment, 240 x 36 n in greatest dimensions and

reaching almost to the middle of the first postgenital segment. The two terminal setae are 47 n

and 83 n, and the adjacent seta of the body is 33 I*.

All three setae are naked. The free segment is conspicuously ornamented with broad, scalelike spines

on its outer surface.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 147c).

The color in life in transmitted light is opaque,

with a few small red globules, the eye red, the egg

sacs opaque gray.

MALE.—The body (Figure 149a) has proportions

similar to those in the female. The length (not

including the setae on the caudal rami) is 0.97 mm

(0.09-1.05 mm) and the greatest width 0.34 mm

(0.32-0.36 mm), based on ten specimens in lactic

acid. The ratio of the length to the width of the

prosome is 1.69:1. The ratio of the length of the

prosome to that of the urosome is 1.67:1. The

epimera of the segments of legs 2—4 are formed

differently than in the female.

The segment of leg 5 (Figure 1496) is 39 x 100 |i.

There is no ventral intersegmental sclerite. The

genital segment is 220 x 186 n. The four postgenital

segments are 36 x 66 v, 39 x 64 n, 35 x 58 n, and 33 x

57 »* from anterior to posterior.

The caudal ramus resembles that of the female

but is smaller, 20 x 24.5 n, and slightly wider than

long.

The rostrum is like that of the female.

The first antenna is similar to that of the female

but there are two additional aesthetes on the

second segment and one additional aesthete on the

fourth segment (at locations indicated by arrows

in Figure 147/) so that the formula is 4, 13 + 2

aesthetes, 6, 3 -f- 1 aesthete, 4 -j- 1 aesthete, 2 -f- 1

aesthete, and 7 + 1 aesthete.

The second antenna (Figure 149c) has a general

form like that of the female but shows sexual

dimorphism in having on the first segment a few

obtuse spines near the seta, on the second segment

a row of slender spinules and an interrupted row

of obtuse spines, and on the fourth segment a few

small inner spinules. The fourth segment is 76 n

along its outer edge, 57 n along its inner edge, and

13 n wide, with the claw 51 n along its axis.

The labrum, mandible, paragnath, first maxilla,

and second maxilla resemble those of the female.

The slender maxilliped (Figure 149d) is very similar to that of P. squamiger. The claw is 265 n

along its axis, with very small rugosities along its

distal concave edge.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented as in the female and have

the same spine and setal formula except for the

endopod of leg 1 (Figure 149e) where the last

segment is 1,1,4. Sexual dimorphism also occurs

on the last segment of the endopod of leg 2 (Figure

149/) where the three spines are much shorter than

in the female and the shape of the segment is

different. Legs 3 and 4 are like those of the female.

Leg 5 (Figure 149g) has a slender, straight free

segment, 55 x 11 n, and its two terminal setae are

31 n and 39 n. The segment is ornamented with a

few very small outer spinules.

Leg 6 is much like that of P. squamiger; its two

naked setae are about 23 n.

The spermatophore (Figure 149h), attached to

the female in pairs, is elongated, 156 x 65 n not including the neck.

The color in life is similar to that of the female

but with fewer red globules.

ETYMOLOGY.—The specific name, sinulariae, is

based on the name of the alcyonacean genus serving

as host.

REMARKS.—Paradoridicola sinulariae is similar in

many respects to P. squamiger (Humes and Frost),

a species associated with Sinularia polydactyla

(Ehrenberg). It may be distinguished from P. squamiger, however, by the fourth segment of the first

antenna being about as long as the second segment,

by the relatively longer and more slender fourth

segment of the second antenna, by the longer fifth

legs and the shape of the genital segment in the

female, and by the nature of the sexual dimorphism

in the endopod of leg 1 of the male.
270

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 149.—Paradoridicola sinulariae, new species. Male: a, dorsal (H) ; b, urosome, dorsal (c);

c, second antenna, postero-inner (F) ; d, maxilliped, inner (E) ; e, endopod of leg 1, anterior

(F) ; /, endopod of leg 2, anterior (F) ; g, leg 5, dorsal (c); h, spermatophore, one of a pair

attached to female, dorsal (£). Scale: c, 0.05 mm; E, F, 0.1 mm; c, H, 0.2 mm.
NUMBER 127

Paradoridicola triquetrus (Humes and Ho, 1968)

Lichomolgus triquetrus Humes and Ho, 1968d, pp. 702-707,

figs. 27-48 [from the alcyonacean Anthelia gracilis (May),

Nosy Be, Madagascar].

Genus Paramacrochiron Sewell, 1949

DIAGNOSIS.—Body cyclopifonn. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. First antenna 7-

segmented. Second antenna 4-segmented, with a

single terminal claw.

Mandible with the basal region distal to the

indentation having on its convex side a setiform

spinulose process and on its concave side a row of

spinules; lash long. Paragnath a small hairy lobe.

First maxilla with three or four elements. Second

maxilla of the usual lichomolgid type. Maxilliped

in the female 3-segmented with a pointed tip, in

271

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except leg 4

endopod which is 1-segmented. Armature of the

lichomolgidiform pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

two terminal elements; in the female the segment

nearly as long as the exopod, in the male relatively

shorter, about equal to the first two exopod segments. Leg 5 with a free segment bearing two

terminal elements. Leg 6 represented by two elements near the genital openings.

Other features as in the species described below.

Associated with scyphozoan medusae.

TYPE-SPECIES. — Paramacrochiron maximum

(Thompson and A. Scott).

REMARKS.—Males are insufficiently known for

the preparation of a key for that sex.

Key to Species of the Genus Paramacrochiron

FEMALES

1. Endopod of leg 4 with an outer marginal notch P. maximum

Endopod of leg 4 without an outer marginal notch 2

2. Second segment of first antenna with a prominent outer protuberance; body length less

than 2.7 mm P. sewelli

Second segment of first antenna without a prominent outer protuberance; body length

more than 2.8 mm S

S. Second segment of second antenna with a short outer marginal process; outer spines on

exopod of leg 4 adculate P. pacificum

Second segment of second antenna without an outer process; outer spines on exopod of

leg 4 with serrated fringes 4

4. Maxilliped with terminal process bent at a right angle P. rhixostomae

Maxilliped with terminal process not bent at a right angle 5

5. Egg sac reaching only a little beyond second postgenital segment; free segment of leg 5

with ratio 43:1, two terminal elements unarmed ..P. etmorense

Egg sac reaching a little beyond caudal ramus; free segment of leg 5 with ratio 4.0:1,

two terminal elements with barbules P. japonUmn

Paramacrochiron maximum (Thompson and A.

Scott, 1903)

Pseudanthessius maximus Thompson and A. Scott, 1903, p.

276, pi. 14, figs. 1-11 [in surface tow-net, Ceylon].—Krishnaswamy. 1950, pp. 52, 53. figs. 60-63 [in plankton, Madras,

India].

Lichomolgus (Macrochiron) maximus.—Monod and Dollfus,

1932a, p. 139.

Macrochiron (Paramacrochiron) maximum.—Sewell, 1949, pp.

108, 109 [in surface tow-net, Andaman Islands].—Krishnaswamy, 1953, pp. 67, 68, fig. 9 [in plankton, Madras,

India].

Macrochiron maximus.—Stock, 1957, p. 381.

Paramacrochiron maximum.—Stock, 1957, p. 381.—Humes,

1969e, p. 180.

Paramacrochiron ennorense Reddiah, 1968

Paramacrochiron ennorense Reddiah, 1968a, pp. 195-201,

figs. 1-3 [from unidentified medusae, near Madras, India].—

Humes. 1969e, p. 182.

Paramacrochiron japonicum Humes, 1970

FIGURE 150

Paramacrochiron japonicum Humes, 1970d, pp. 223-232, figs.

1-31 [from the medusa Thysanostoma thysanura Haeckel,

Sirahama, Wakayama-ken, Japan].
272

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE \50.—Paramacrochiron japonicum Humes. Female: A, dorsal; b, second antenna; e, mandible; d, leg 4 and intercoxal plate. Male: e, dorsal; /, leg 4 and intercoxal plate. (From Humes,

1970d, figs. 1, 9, 11, 20, 23, 27.) Length of female 3.28 mm, of male 2.69 mm.

Paramacrochiron pacificutn (C. B. Wilson, 1950)

Pseudanthessius pacificus C. B. Wilson, 1950, pp. 315, 316,

pi. 32, figs. 481-489 [in plankton, off Luzon, Philippine

Islands].

REMARKS.—Although the original description

lacks certain information, on the basis of the large

size and the relatively long endopod of leg 4 it

seems justified to place this species in Paramacrochiron.

Paramacrochiron rhizostomae Reddiah, 1968

Paramacrochiron rhizostomae Reddiah, 1968a, pp. 205-208,

figs. 5-6 [from the medusa Rhizostoma sp., Gulf of Mannar, India].-Humes 1969e, p. 182.

Paramacrochiron sewelli Reddiah, 1968

Paramacrochiron sewelli Reddiah, 1968a, pp. 201-205, fig. 4

[from the medusa Lychnorhiza malayensis Stiasny, near

Madras, India] .-Humes, 1969e, p. 182.

Genus Paramolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmentetl. Caudal ramus with six setae, in P. clavatus very short.

Rostrum rounded or incomplete posteroventrally.

First antenna 7-segmented, in the female with the

formula 4, 13, 6, 3, 4 -}- 1 aesthete, 2 + 1 aesthete,

and 7 -f- 1 aesthete; in the male with 4, 13 -|- 2

aesthetes, 6, 3 -f 1 aesthete, 4+ 1 aesthete, 2+ 1

aesthete, and 7+ 1 aesthete (except that in P.

constrictus and P. insectus the second segment is

13+ 1 aesthete). Second antenna 4-segmented

with the formula 1,1,3 and I + several small elements, there being only one terminal claw.

Labrum with the two lobes separated by a cleft

or indentation. Mandible with the basal region

distal to the indentation having on its convex side

a scalelike area with spinules followed by a serrated

fringe (except that in P. constrictus the scalelike

area is naked and the first few serrations in the
NUMBER 127 273

fringe are directed proximally) and on its concave

side a row of spinules. Paragnath a small hairy

lobe. First maxilla with four elements (three in

P. constrictus and P. insectus). Second maxilla of

the usual lichomolgid form. Maxilliped in the female 3-segmented with a pointed tip, in the male

4-segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1; II, the seta feathered. Leg 1

endopod in the male with the third segment 1,1,4

instead of 1,5 as in the female (except without

such sexual dimorphism in P. clavatus). Leg 5

with a free segment bearing two elements. Leg 6

represented in the female by the two setae and the

process near the area of attachment of each egg

sac, in the male by the ventral posterolateral flap

on the genital segment bearing two setae.

Other features as in the species below.

Associated with actinarians and octocorals.

TYPE-SPECIES.—Paramolgus politus (Humes and

Ho).

ETYMOLOGY.—The name is a combination of the

Greek naoa (near), alluding to the lichomolgid affinities of this genus, and »u>Xyo;. Gender masculine.

REMARKS.—The six species of Paramolgus form

a fairly cohesive group. P. anomalus is placed here

with reservations since we have not seen actual

Key to Species of the Genus Paramolgus

FEMALES (except P. anomalus)

1. Area of attachment of egg sacs bearing a large pointed bladelike process in addition to the

two setae p. spathophorus

Area of attachment of egg sacs with only a small spinous process in addition to the

two setae 2

2. Body length 2.27 mm; longest seta on caudal ramus much shorter than ramus; lash of

second maxilla short, bearing only two large spines and two spikelike spinules P. clavatus

Body length less than 2 mm; longest seta on caudal ramus much longer than ramus; lash

of second maxilla longer, with a graded row of spinules or teeth S

S. Lash of second maxilla with a graded row of teeth; two elements on second segment of

maxilliped very unequal in length 4

Lash of second maxilla with a graded row of spinules; two elements on second segment

of maxilliped nearly equal in length 5

4. Caudal ramus 3.8:1; free segment of leg 5 without an inner proximal expansion P. politus

Caudal ramus 1.5:1; free segment of leg 5 with an inner proximal expansion P. simulans

5. Caudal ramus 1.9:1; third segment of second antenna with an unusually long seta (reaching

to more than halfway on the claw) P. constrictus

Caudal ramus 2.65:1; all three setae on third segment of second antenna short (less than

half the length of segment) P. insectus

MALES (except P. anomalus)

1. Lash of second maxilla short, bearing only two large spines and two spikelike elements;

inner seta on coxa of leg 4 long and plumose P. clavatus

Lash of second maxilla longer, with a graded row of spinules or teeth; inner seta on coxa

small and either naked or with a few barbules 2

2. Rostrum with two small knoblike protuberances on posteroventral margin P. insectus

Rostrum without such knobs 3

3. Free segment of leg 5 unornamented; third segment of second antenna with an unusually

long seta (reaching to more than halfway on the claw) P. constrictus

Free segment of leg 5 with very small spinules; all three setae on third segment of second

antenna shorter, reaching at most to half way on the fourth segment 4

4. Caudal ramus quadrate P. spathophorus

Caudal ramus distinctly longer than wide 5

5. Caudal ramus 3.8:1; with refractile knobs on inner surface of first two segments of second

antenna P. poUtus

Caudal ramus 1.5:1; with spinules on inner surface of first two segments of second

antenna P. simulant
274 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

specimens and since A. Scott's original description

is incomplete. P. constrictus and P. insectus, while

possessing many of the generic features, have certain characters in common which are somewhat at

variance with the other members of the genus; for

example, only one aesthete on the second segment

of the first antenna of the male, only three elements on the first maxilla, and the laterally notched

or insected genital segment in the female. P. clavatus similarly differs in having very short setae on

the caudal ramus and in lacking sexually dimorphic

armature on the endopod of leg 1 in the male.

Nevertheless, we feel that, for the present at least,

all six species should be grouped in the one genus.

Paramolgus anomalus has been omitted from the

following keys because of inadequate information

in the available description and figures.

Paramolgus politus (Humes and Ho, 1967)

FIGURE 151

Lichomolgus politus Humes and Ho, 1967d, pp. 5-6, figs.

1-28 [from the actinarian Rhodactis rhodostoma Ehrenberg. Nosy Komba, near Nosy Be, Madagascar].

Paramolgus anomalus (A. Scott, 1909)

Lichomolgus anomalus A. Scott, 1909, pp. 264-265, pi. 67,

figs. 6-17 [from Banda Sea, south of Amboina, 5*26.7' S,

127-6.5' EJ.-Sewell. 1949, p. 97 [in debris from a jar of

Alcyonacea, southern coast of Arabia].

}Lichomolgus (Macrochiron) albus.—Monod and Dollfus,

1932a. p. 139.

Paramolgus clavatus (Humes and Ho, 1968)

Lichomolgus clavatus Humes and Ho, 1968d, pp. 750-735,

figs. 128-148 [from the telestacean Coelogorgia palmosa

Milne Edwards and Haime, region of Nosy Be, Madagascar].

NEW HOSTS (collected by AGH).—2 <J ef from

Lemnalia longiramus Verseveldt, in 12 m, west of

harbor at Hellville, Nosy Be\ Madagascar, 4 August

FIGURE 151-Paramolgus politus (Humes and Ho). Female: a, dorsal; b, second antenna; c,

mandible; d, leg 4 and intercoxal plate. Male: e, dorsal. (From Humes and Ho, 1967d, figs. 1,

8, 10, 19, 21.) Length of female 1.78 mm, of male 1.33 mm.
NUMBER 127 275

1967; 1 cT. 1 ? copepodid from Lemnalia cervicornis (May), in 20 m, Bane de Cinq Metres, near

Nosy Be\ 6 August 1967; 1 9 from Lemnalia crassicaulis Verseveldt, in 20 m, Bane de Cinq Metres,

near Nosy Be\ 6 August 1967.

Paramolgus constrict us (Humes, 1969)

Lichomolgus constrictus Humes 1969b, pp. 2-10, figs. 1-31

[from the antipatharian Antipathes ericoides Pallas, region of Nosy Be, Madagascar].

Paramolgus insect us (Humes, 1969)

Lichomolgus imeclus Humes. 1969b, pp. 10-17, figs. 32-63

[from the antipatharians Antipathes sp. cf. spinescem

Gray, Antipathes myriophylla Pallas, and Antipathes abies

(Linnaeus), region of Nosy Re, Madagascar].

Paramolgus simulans (Humes and Ho, 1967)

Lichomolgus simulans Humes and Ho, 1967d, pp. 6-9, figs.

29-53 [from the actiniarian Rhodactis rhodostoma (Ehrenberg). Nosy Komba, near Nosy Be, Madagascar].

Paramolgus spathophorus (Humes and Ho, 1968)

Lichomolgus spathophorus Humes and Ho, 1968c, pp. 674-

680, figs. 128-147 [from the alcyonacean Sarcophyton glaucum (Quoy and Gaimard), Tany K.ely, near Nosy Be,

Madagascar. (Dr. Verseveldt has recently changed this

host identification to Sarcophyton trocheliophorum Marenzeller.)]

NEW RECORD.—2 $ $ from single colony of Sarcophyton glaucum, in 1 m, Ankify, on the mainland of Madagascar, near Nosy B6, 23 August 1967,

collected by AGH.

NEW HOST.—5 9 9 from single colony of the

alcyonacean Sarcophyton acutangulum (Marenzeller), in 25 m, Tany Kely, a small island south of

Nosy B£, Madagascar, 14 August 1967, collected by

AGH.

Genus Paraphiloconcha Yamaguti, 1936

DIAGNOSIS.—Body transformed, elongated, the

prosome in the female with the segments of legs

2-4 having long, blunt, digitiform lateral processes,

but in the male with similar processes only on the

segments of legs 2 and 3. External segmentation

of both prosome and urosome obscure. Caudal ramus with setae shorter than the ramus. Rostrum

present. First antenna 7-segmented. Second antenna 4-segmented with a single terminal claw.

Labrum incised medially. Mandible with a long,

tapered blade and lash. First maxilla with two elements. Second maxilla of the usual lichomolgid

type. Maxilliped in the female 3-segmented, in

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having two outer spines and one

terminal spine. Leg 1 endopod with the same formula in both sexes. Leg 5 with a free segment

bearing two terminal elements.

Other features as in the species below.

Associated with bivalve mollusks.

TYPE-SPECIES.—Paraphiloconcha meretricis Yamaguti.

Paraphiloconcha meretricis Yamaguti, 1936

FIGURE 152

Paraphiloconcha meretricis Yamaguti, 1936, pp. 121-123, pi.

12, figs. 63-75 [from the pericardium of Meretrix lamarcki

Deshayes, Sea of Japan].

Genus Paredromolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum not well

developed. First antenna 7-segmented, with the

armature 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete, and 7 -j- 1 aesthete; in the male 4, 13 -j- 2

aesthetes, 6, 3 + 1 aesthete, 4 + 1 aesthete, 2 + 1

aesthete, and 7 -f- 1 aesthete. Second antenna 4-

segmented, with the formula 1,1,3 and II + a

mucronate seta and several smaller elements, there

being two unequal terminal claws, one slender and

unjointed.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal area beyond the

indentation having on its convex side a distally

directed tooth and a striated fringe and on its concave side a row of spinules; lash long. Paragnath a

hairy lobe. First maxilla with three elements. Second maxilla of the usual lichomolgid type. Maxilliped of the female 3-segmented, with a pointed

tip, in the male 4-segmented (assuming the prox-
276

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE \52.—Paraphiloconcha meretricis Yamaguti. Female: a, donal; b, second antenna; c,

mouthpans, including mandible, first maxilla, and second maxilla; d, leg 4. Male: e, dorsal. (From

Yamaguti, 1936, pi. 12, figs. 63, 64, 67, 68, 74.) Length of female 3.1-5.5 mm. of male 2.4-2.75

imal part of the claw to represent a fourth segment).

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Leg 4 exopod

with the third segment 111,1,5- Leg 4 endopod with

0-1 ;II; the seta naked in the female, with extremely

small barbules in the male. In the male, leg 1

endopod with the third segment 1,1,4 instead of 1,5

as in the female. Leg 5 in both sexes with a free

segment bearing two terminal setae. Leg 6 in the

female represented by two small spines near the

area of attachment of each egg sac, in the male by

a posteroventral flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with alcyonaceans.

TYPE-SPECIES.—Paredromolgus decorus (Humes

and Frost).

ETYMOLOGY.—The name is a combination of the

Greek words jraoeooog (beside), referring to its affinity with Lichomolgus, and

Paredomolgus decorus (Humes and Frost, 1964)

FICUSE 153

Lichomolgus decorus Humes and Frost, 1964, pp. 140-142,

figs. 99-133 [from the alcyonacean Cladiella laciniosa

Tixicr-Durivault, Nosy H<. Madagascar].—Bouligand, 1966,

p. 269.

NEW HOST RECORDS (specimens collected by

AGH).—From Cladiella latissima (Tixier-Durivault): 65 9 9 , 62 o* cf. and 3 copepodids in 1 m,

Pte. de Tafondro, Nosy Be1, Madagascar, 12 July

1967; 43 9 9 , 16 tfd1. and 12 copepodids in 18 m,

Bane du Touareg, Bay of Ampasindava, near Nosy

B£, 11 July 1967. From Cladiella sphaerophora

(Hemprich and Ehrenberg): 2 99,3 c Tcf» nl

m

»

Nosy N'Tangam, near Nosy Be\ 21 July 1967.

Genus Pennatulicola Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform, with broad, flattened prosome and short urosome. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. First antenna 7-segmented. Second antenna 4-segmented, with two

terminal claws.

Mandible with the basal region beyond the indentation having on its convex side a prominent,

spinose, scalelike area followed by numerous spinules and on its concave side rows of spinules; lash

long. First maxilla with two or three elements. Second maxilla of the lichomolgid type. Maxilliped
NUMBER 127 277

FIGURE l5S.—Paredromolgus decorus (Humes and Frost). Female: a, dorsal; b, second antenna;

c, mandible; d, leg 4. Male: e, dorsal; /, endopod of leg 4. (From Humes and Frost, 1964, figs.

99, 106, 108, 118, 120, 131 in part.) Length of female 0.93 mm, of male 0.94 mm.

in the female 3-segmented with a pointed tip, in

the male 1-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

the formula 0-1 ;II, the seta feathered. Leg 1 endopod with the same formula in both sexes. Leg 5

with a free segment bearing two terminal elements

and having a process on its inner margin.

Other features as in the species below.

Associated with pennatulaceans.

TYPE-SPECIES.—Pennatulicola pteroidis (Delia

Valle, 1880).

ETYMOLOGY.—The generic name is formed from

Pennatulacea and the Latin colo, to inhabit. Gender masculine.

REMARKS.—A satisfactory key to the males of

Pennatulicola cannot be prepared from the existing descriptions of that sex.

Key to Species of the Genus Pennatulicola

FEMALES

1. Second segment of the endopod of legs 1-3 with two outer spiniform projections (in a

beaklike arrangement) P. pterophiku

Second segment of the endopod of legs 1-3 with a single spiniform projection 2

2. Free segment of leg 5 broadest in its distal half, with a small toothlike process on its inner

margin P. pteroidis

Free segment of leg 5 broadest in its proximal half, with a prominent toothlike process

on its inner margin P. serratipcs
278 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pennatulicola pteroidis (Delia Valle, 1880)

Lichomolgus pteroidis.-Delia Valle, 1880a, pp. 99-100. pi. 6,

figs. 33-42 [from Pteroides spinulosus, Naples, Italy]; 1880b,

pp. 119, 120, pi. 2. figs. 33-42 [from the sea pen Pteroides

spinulosus Herkl., Naples, Italy].-Carus, 1885, p. 351.—

De Zulueta. 1912, pp. 10-11, 50.-Stock, 1959b, pp. 66-70,

figs. 5, 6 [from Pteroides spinulosus, Gulf of Naples. Italy];

1960c, p. 242 [from the pennatulacean Pteroides griseum

Bohadsch, Banyuls, Mediterranean coast of France].—

Bouligand, 1960. p. 258; 1966, p. 269.-Stock and Kleeton,

1963b, pp. 257, 258, fig. 7 [from Pteroides griseum, Banyuls, Mediterranean coast of France].

Stellicola kossmanniana Valle, 1880, pp. 52-53, figs. 1-6

[from the pennatulacean Pteroeides griseum longespinosum Klk., Gulf of Constantinople].

Pennatulicola pterophilus (Stock, 1962)

FIGURE 154

Lichomolgus pterophilus Stock, 1962, pp. 155-163. figs. 1-20

[from the pennatulacean Pteroeides lacazei Kdlliker (perhaps Pteroeides argenteum Ellis and Solander), Billiton

Island, Sunda Sea].-Bouligand, 1966, p. 269.

Pennatulicola serratipes (Ummerkutty, 1962)

Lichomolgus serratipes Ummerkutty. 1962, pp. 55-59, 69,

pi. 13, figs. 5-10, pi. 14, figs. 1-5. addendum fig. la-e

[from the sea pen Pteroeides esperi Herklots. Gulf of Mannar, southeastern India].

Genus Philoconcha Yamaguti, 1936

DIAGNOSIS.—Body modified, elongate. Urosome

in the female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. Rostrum

present. First antenna 7-segmented. Second antenna 4-segmented, with a single terminal claw.

Labrum incised medially. Mandible with a long,

slender bipectinate blade and lash. First maxilla

with two setae. Second maxilla of the usual lichomolgid type. Maxilliped in the female reduced,

swollen, tipped with a small digitiform process and

two setae; in the male 4-segmented (considering

the proximal part of the claw to represent a fourth

segment).

FIGURE 15*.-Pennatulicola pterophilus (Stock) . Female: a, dorsal; b, mandible; c, leg 4; d, leg 5.

Male: e, couple consisting of an adult male and a female copepodid; /, second antenna. (From

Stock. 1962, figs. 1, 2, 4, 13. 18, 19.) Length of female 1.87 mm. of male 135 mm.
NUMBER 127 279

Key to Species of the Genus Philoconcha

FEMALES

Length up to 6.6 mm; caudal ramus 0.65-0.66 x OJ21-O.23 mm. P. amygdalae

Length 7.5 mm; caudal ramus 1.1 x 025 mm P. paphiae

MALES

Length 2.7-2.8 mm; caudal ramus 0.45 x 0.12-0.15 mm P. amygdalae

Length 5.75-4.4 mm; caudal ramus 0.82-0.96 x 0.15-0.25 mm. P. paphiae

Legs 1-4 with 5-segmented rami except for leg 4

endopod which consists of two, usually unarmed,

nodular segments. Armature lichomolgidiform in

pattern. Leg 4 exopod with the third segment having two outer spines and one terminal spine. Leg 1

endopod with the same formula in both sexes. Leg

5 with a free segment bearing two terminal elements.

Other features as in the species below.

Associated with bivalve mollusks.

TYPE-SPECIES.—Philoconcha amygdalae Yamaguti.

Philoconcha amygdalae Yamaguti, 1936

FIGURE 155a-g

Philoconcha amygdalae Yamaguti, 1956, pp. 118, 119, pi.

10, figs. 59-48; pi. 11, figs. 49-51 [from the bivalve Venerupis (Amygdala) philippinarum (Adams and Reeve), Tiba

Prefecture, Japan].

FIGURE 155.—a-g, Philoconcha amygdalae Yamaguti. Female: a, dorsal; b, second antenna;

c, mouthparts, including mandible, first maxilla, and second maxilla; d, endopod of leg 4. Male:

e, dorsal; /, leg 4; g, leg 5. (From Yamaguti, 1936, pi. 10, figs. 39, 40, 42, 45, pi. 11, figs. 49-51.

On Yamaguti's pi. 10 the designation for figs. 42 and 43 should be reversed.) Length of female

up to 6.6 mm, of male 2.7-2.8 mm. h, i, Philoconcha paphiae Yamaguti. Male: h, leg 4; i,

endopod of leg 4. (From Yamaguti, 1936, pi. 11, figs. 60, 61.) Length of female 15 mm, of

male 3.75-4.4 mm.
280

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Philoconcha paphiae Yamaguti, 1936

FIGURE 155/J,»

Philoconcha paphiae Yamaguti, 1936, pp. 119, 120, pi. 11,

figs. 52-62 [from the pericardium of the bivalve Paphia

euglypta (Philippi), Inland Sea, Japan].

Genus Plesiomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum rounded

posteroventrally. First antenna 7-segmented, in the

female with the formula 4, 13, 6, 3, 4 + 1 aesthete,

2 + 1 aesthete, and 7+ 1 aesthete; in the male

with 4, 13 + 1 aesthete, 6, 3 -f 1 aesthete, 4 + 1

aesthete, 2 + 1 aesthete, and 7 -+- 1 aesthete. Second antenna 4-segmented, with the armature 1,1,3

and II -J- 5 setae, there being two terminal claws.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal part bearing a small

sclerotized area on its posterior surface, its convex

side with a proximally directed, toothlike process

followed by a striated fringe, its concave side with

a row of spinules; lash long. Paragnath a small

hairy lobe. First maxilla with two elements. Second maxilla of the usual lichomolgid type. Maxilliped of the female 3-segmented, with a pointed

tip; in the male 4-segmented (assuming that the

proximal part of the claw represents a fourth segment).

Legs 1^1 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform type. Leg 4 exopod with

the third segment 11,1,5. Leg 4 endopod 0-1 ;II,

the seta plumose. In the male, leg I endopod with

the third segment 1,1,4 instead of 1,5 as in the female. Leg 5 in both sexes with a free segment bearing two terminal setae. Leg 6 in the female

represented by the two setae near the area of attachment of each egg sac, in the male by the

posteroventral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated with stoloniferan octocorals.

TYPE-SPECIES.—Plesiomolgus organicus (Humes

and Ho).

ETYMOLOGY.—The name is a combination of the

Greek word nXumo; (near), referring to the affinity

of this genus to Lichomolgus, and noXyoc Gender

masculine.

Key to Species of the Genus Plesiomolgus

FEMALES

Genital segment with lateral indentations near middle; caudal ramus 36 x 24 fi (1.5:1;

P. organicui

Genital segment with lateral indentations further back; caudal ramus 28 x 23 ft (122:1)

P. conjunctus

MALES

Caudal ramus 31 x 21 n (1.48:1); free segment of leg 5 36 x 9 u. (4:1) P. organicTM

Caudal ramus 25 x 21 n (1.19:1); free segment of leg 5 28 x 8 p (35:1) P. conjunctus

Plesiomolgus organicTM (Humes and Ho, 1967)

FIGURE 156

Lichomolgus organicus Humes and Ho, 1967a, pp. 2-7, figs.

1-30 [from the stoloniferan Tubipora musica (Linnaeus),

region of Nosy B6, Madagascar].

Plesiomolgus conjunctus (Humes and Ho, 1967)

Lichomolgus conjunctus Humes and Ho, 1967a, pp. 7^-10,

figs. 31-36 [from Tubipora musica, region of Nosy B£,

Madagascar],

NEW RECORD.—4 9 $ , 1 o* from Tubipora musica, in 1 m, Tany Kely, near Nosy Be, Madagascar, 27 May 1967, collected by AGH.

Genus Prionomolgus Humes and Ho, 1968

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum weakly developed, without a definite posteroventral margin.

First antenna 7-segmented, in the female with the
NUMBER 1*7 281

a

FicuftE 156.—Plcsiomolgus organieus (Humes and Ho). Female: a, dorsal; b, second antenna; r,

mandible; d, second maxilla; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes and

Ho, 1967a, figs. 1, 8, 10, 20. 22.) Length of female 0.99 mm, of male 0.89 mm.

formula 4, 13, 6, 3, 4 -f- 1 aesthete, 2 + 1 aesthete,

and 7 + 1 aesthete; in the male with -1,13 + 2

aesthetes, 6, 3 + 1 aesthete, 4+ 1 aesthete, 2+ 1

aesthete, and 7 + 1 aesthete. Second antenna 3 -

segmented (the original third and fourth segments

fused), with the formula 1, 1, 3 + I + one small

element, there being a single terminal claw.

Labrum with a deep median cleft between two

lobes. Mandible with the base separated by a constriction into an unornamented proximal part and

a distal area whose concave edge has two lobes,

both with strongly serrated margins, and whose convex edge has a short row of spinules followed by

a triangular process and then by a finely spinulose

margin; lash relatively short. Paragnath a small

lobe, naked except for one setule. First maxilla

with three elements with serrated flanges. Second

maxilla of the usual lichomolgid type. Maxilliped

of the female 3-segmented, the third segment with

a spiniform tip. Maxilliped of the male 4-segmented (assuming that the proximal part of the

claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform pattern, except leg 4 endopod which has the formula 0-1 ;I. Leg 4 exopod

with the last segment having 11,1,5. Formula for

the male similar except leg 1 endopod, where the

third segment is 1,1,4 instead of 1,5 as in the female. Leg 5 in both sexes with a free segment bearing two terminal setae. Leg 6 in the female

represented by two setae near the area of attachment of each egg sac, in the male by a posteroventral flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Prionomolgus lanccolatus Humes

and Ho.

Prionomolgus lanceolatus Humes and Ho, 1968

FIGURE 157

Prionomolgus lanceolatus Humes and Ho, 1968a, pp. 372-

375, figs. 183-210 [associated with the hard coral Fachyseris speciosa (Dana), region of Nosy Be*, northwestern

Madagascar].

Genus Pseudomacrochiron Reddiah, 1969

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six setae. First antenna 7-segmented. Second antenna 4-segmented, the last

segment with two claws (except one clawlike spine

in P. fucicolum) and several setae. Mandible with

the basal region beyond the indentation having on

its concave side a row of setae and on its convex

side a spinulose scalelike area followed by a serrated fringe; lash long. First maxilla with three or

four elements. Second maxilla of the usual licho-
282

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 157.—Prionomolgus lanceolatus Humes and Ho. Female: a, dorsal; b, second antenna; c,

mandible; d, endopod of leg 4. Male: e, dorsal. (From Humes and Ho, 1968a. figs. 18S. 190.

192, 201, and 203.) Length of female 1.36 mm. of male 1.04 mm.

molgid type. Maxilliped in the female 3-segmented; in the male 4-segmented (assuming that

the proximal part of the claw represents a fourth

segment).6

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 1-segmented. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5 (formula unknown in

P. malayense). Leg 4 endopod in the female much

shorter than the exopod and bearing two terminal

spines. Leg 5 with a free segment bearing two

terminal elements.

Other features as in the species below.

Associated with scyphozoan medusae, as far as

known.

TYPE-SPECIES.—Pseudomacrochiron parvum (A.

Scott).

T he maxillipeds in P. stocki represented by Reddiah

(1969) as 2-segmented in the female and 3-segmented in the

male, are, judging from his figs. 2D and 3B, in all probability 3- and 4-segmented respectively.

REMARKS.—We find it impossible to prepare a

satisfactory key to the species of Pseudomacrochiron

on the basis of available information.

Pseudomacrochiron parvum (A. Scott, 1909)

FICURE 158

Pseudanthessius parvus A. Scott, 1909, pp. 269. 270, pi. 69,

figs. 1-7 [from Lawui, coast of Obi Major, 0°24' S, 127° 36'

E.]

Lichomolgus (Macrochiron) parvus.—Monod and Dollfus,

1932a, p. 139.

Macrochiron (Paramacrochiron) parvum.—Sev/eU, 1949, p.

109 [in weed-washings, Nankauri Harbor, Nicobar Islands].

-Canapati and Shanthakumari. 1962, pp. 10, 12, 16.

Macrochiron parvum.—Stock, 1957, p. 381.

Pseudomacrochiron parvum.—Reddiah, 1969, p. 44.

Pseudomacrochiron fucicolum (T. Scott, 1912)

Pseudanthessius fucicolus T. Scott, 1912, pp. 571, 572, pi. 12.

figs. 1-13 [from gulf-weed, St. Helena, 15°57' S, 5°40' W,

to Tuskar Rock, 51° IS' N, 7°2O' W].
NUMBER 127 283

FIGURE l&S.—Pseudomacrochiron parvum (A. Scott). Female: a, dorsal; b, second antenna; c,

mandible and first maxilla; d, leg 4. (Redrawn after A. Scott, 1909, pi. 69. figs. 1, 3, 4, 7.)

Length of female 0.9 mm; male unknown.

Pseudanthessius fucicolus.—non Kelleria, see Nicholls, 1944,

p. 54.

Macrochiron (Paramacrochiron) fucicolum.—Sewell, 1949, p.

108.

Kelleria /uricoto.-Bayly, 1971, p. 111.

Pseiidomacrochiron malayense (Sewell, 1949)

Macrochiron (Paramacrochiron) malayense Sewell, 1949, pp.

109-111, fig. 28A-F [in surface tow-net, Kurau River, Perak,

Federated Malay States; and off Viper Island, Port Blair,

Andaman Islands].

Macrochiron malayense.—Stock, 1957, p. 382.

Pseudomacrochiron malayense.—Reddiah, 1969, p. 44.

Pseiidomacrochiron ornatum (Krishnaswamy,

1952)

Macrochiron (Paramacrochiron) ornatum Krishnaswamy,

1952. pp. 330-333, fig. 4 [in plankton, Madras, India];

1953, p. 67 [in plankton, Madras, India].

Pseudomacrochiron omnium.-Reddiah. 1969, p. 44.

Pseudomacrochiron stocki Reddiah, 1969

Pseudomacrochiron stocki Reddiah, 1969, pp. 44-49, figs. 1-3

[from the medusa Dactylometra quinquicirrha L. Agassiz,

Madras, India].

Genus Rakotoa Humes and Stock, 1972

DIAGNOSIS.—Body modified. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six relatively short setae. Rostrum with a weakly defined posteroventral margin.

First antenna 7-segmented, with the formula in the

female 4, 13, 6, 3, 4 + 1 aesthete, 2 + 1 aesthete,

and 7 + 1 aesthete; in the male 4, 13 -{- 2 aesthetes, 6, 3 -f 1 aesthete, 4 -j- 1 aesthete, 2 + 1

aesthete, and 7 -j- 1 aesthete. Second antenna 4—

segmented, with the armature 1, 1, 3, and I -|- 1

setule, there being a single terminal claw.

Labrum with the two lobes separated by a broad

indentation. Mandible with the basal area distal

to the indentation having on its convex side a serrated fringe and on the concave side two lobes,

each with two rows of spinules; lash stout and moderately long. First maxilla with three elements.

Second maxilla of the usual lichomolgid type, but

the first segment bearing a long digitiform process

as in Andrianellus. Maxilliped in the female 3-

segmented, with a pointed tip, in the male 4-segmented (assuming the proximal part of the claw

to represent a fourth segment).
284

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is vestigial and represented only

by an unornamented lobe which may bear a

minute distal lobe suggesting a 2-segmented condition. Armature of legs 3, 4 reduced, leg 3 endopod

being 0-1; 0-2; 1 and leg 4 exopod 1-0; 0-1; 1,3.

Both sexes with similar armature. Leg 5 in both

sexes with a free segment bearing two terminal

setae. Leg 6 in the female represented by the two

setae and spiniform process near the area of attachment of each egg sac, in the male by a posteroventral flap on the genital segment bearing two

setae.

Other features as in the species described below.

Associated with madreporarian corals.

TYPE-SPECIES.—Rakotoa proteus, new species.

ETYMOLOGY.—The generic name is for Rakoto,

a prince who became King Radama II of Madagascar. Gender masculine.

Rakotoa proteus, new species

FIGURES 159-161

TYPE MATERIAL.—10 ? $ , 4 0*0* from seven

colonies of the madreporarian coral Favia sp., in

10 cm, Boloboxo, Nosy Faly, an island to the east

of Nosy B£, northwestern Madagascar, 10 August

1964, collected by AGH. Holotype $, allotype, and

8 paratypes (7 ? ? , 1 c?) deposited in USNM, the

remaining paratypes (dissected) in the collection of

AGH.

FEMALE.—The body (Figure 159a) has a somewhat thickened prosome as in Andrianellus exsertidens. The length (not including the setae on the

caudal rami) is 1.60 mm (1.58-1.64 mm) and the

greatest width 0.60 mm (0.55-0.63 mm), based on

ten specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.38:1. The ratio

of the length of the prosome to that of the urosome

is 1.26:1. The segment of leg 1 is separated from

the cephalosome by a transverse dorsal furrow. The

epimera of the segments of legs 1-4 are only slightly

developed and rounded.

The segment of leg 5 (Figure 1596) is 96 x 350 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 230 x 286 n in greatest dimensions,

broadest in its anterior fourth, and tapered posteriorly. The areas of attachment of the egg sacs are

located dorsolaterally just in front of the middle

of the segment. Each area (Figure 159c) bears two

very small setae about 5 M long and a minute spiniform process. The three postgenital segments are

109 x 156 », 83 x 138 »i, and 109 x 143 M from anterior to posterior. The posterior border of the anal

segment bears on each side dorsal and ventral rows

of minute spinules.

The caudal ramus (Figure 159d) is elongated,

125 x 47 »*, or 2.66 times longer than wide. All the

setae are shorter than the ramus and naked. The

outer lateral seta is 44 n, the dorsal seta (displaced

to the inner distal edge of the ramus) 40 n, the

outermost terminal seta 45 n, the innermost terminal seta 43 |i, and the two median terminal setae

68 n (outer) and lo n (inner).

The body surface has a few small hairs (sensilla)

and refractile points as shown in Figure 159a,6.

The egg sac is unknown.

The first antenna (Figure 159*) is 7-segmented

and 307 n long. The lengths of the seven segments

(measured along their posterior nonsetiferous margins) are 18 ('>'> n along the anterior margin), 87,

34, 48, 41, 24, and 18 n respectively. The formula

for the armature is 4, 13, 6, 3, 4 -f- 1 aesthete, 2 -f 1

aesthete, and 7 -f 1 aesthete. All the setae are

naked.

The second antenna (Figure 159/) is 4-segmented.

Each of the first two segments bears a small inner

seta. The third segment bears three setae. The

fourth segment, 50 n along its outer edge, 30 »i

along its inner edge, and 22 n wide, bears terminally a claw, 33 n along its axis, and a minute

setule.

The labrum (Figure 159g) has two broad, posteroventral lobes.

The mandible (Figure 159/i) resembles that of

Andianellus exsertidens, but each of the two spinulose lobes on the concave side has two rows of

spinules and the convex side of the base is smooth,

without a spiniform process. Paragnaths could not

be identified with certainty. The first maxilla (Figure 159i) has three elements. The second maxilla

(Figure 159;") resembles that of A. exsertidens, but

the toothlike process on the first segment is relatively shorter and more slender. The maxilliped

(Figure 159A) also is similar to that of A. exsertidens, but the two setae on the second segment are

more unequal; the third segment has a larger
FIGURE 159.—Rakotoa proteus, new species. Female: a, dorsal (A); b, urosome (B) ; c, area

of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, first antenna, with arrows

indicating positions of additional aesthetes in male, anterodorsal (D) ; /, second antenna, posterior (D) ; g, labrum. ventral (D) ; h, mandible, posterior (c); i, first maxilla, anterior (c); /,

second maxilla, anterior (E) ; k, maxilliped, inner (F.) ; /, area between maxillipeds and first

pair of legs, ventral (E) . Scale: A, B, 05 mm; c, 0.05 mm; D, E, 0.1 mm.
286

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

naked articulated spine and a small seta and

terminates in a naked spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 159/) protrudes ventrally. The bases of the maxillipeds are not connected by a line as in many lichomolgids.

Legs 1-4 (Figure 160a-c,e) have 3-segmented

rami except for the endopod of leg 4 which is

vestigial. The armature of these legs is as follows:

Pi coxa 0-1 basis 1-0 exp 1-0; 1-1; III.I.4

enp 0-1; 0-1; U

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 11,1,5

enp 0-1; 0-2; U U

P, coxa 0-1 basis 1-0 exp 1-0; 1-1; 1,1.4

enp 0-1; 0-2; 1

P. coxa 0-1 basis 1-0 exp 1-0; 0-1; 13

enp —

The coxa of legs 1-3 bears a long plumose inner

seta, but in leg 4 that seta is much smaller (17 n)

and naked. The inner margin of the basis bears a

row of hairs in legs 1-3 but is naked in leg 4. The

spines on the rami have their distal ends slightly

expanded as a smooth lamella surmounted by a

slender flagellum. In one female the endopod of

the right leg (Figure 160d) shows an incomplete

separation of the second and third segments (the

line of articulation being present only on the posterior surface). In another female the exopod of

leg 4 (Figure 160/) shows a similar incomplete

separation of these segments and has an extra

spine, so that the formula is 1-0; 0-1; 1,1,3. The

endopod of leg 4 is vestigial, represented in Figure

160<? by a small lobe only 5 n long. The form of this

endopod is variable, however, with occasionally

a minute distal lobe, suggesting a 2-segmented condition. Figure 160gJi illustrates the pairs of endopods in two females.

Leg 5 (Figure 160i,;) has an elongated unornamented free segment 133 x 47 f* in lateral (outer)

view. The two terminal setae are 18 n and 40 n, and

the adjacent seta on the body is 20 n. All three

setae are naked.

Leg 6 is represented by the two minute setae on

the area of attachment of each egg sac (Figure

159c).

The color in life in transmitted light is opaque

gray, the intestine reddish, the eye not visible.

MALE.—The body form (Figure 16 la) resembles

that of the female. The length (without the ramal

setae) is 1.51 mm (1.48-1.55 mm) and the greatest

width 0.54 mm (0.53-0.54 mm), based on four

specimens in lactic acid. The ratio of the length

to the width of the prosome is 1.42:1. The ratio

of the length of the prosome to that of the urosome

is about 1:1. The segmentation and the development of the epimera are like the female.

The segment of leg 5 (Figure 1616) is 70 x 275 I*.

There is no ventral imersegmental sclerite. The

genital segment is 234 x 319 i», wider than long.

The four postgenital segments are 91 x 138 n, 86

x 122 ii, 70 x 113 n, and 94 x 135 M- from anterior

to posterior.

The caudal ramus is similar to that of the female, 117 x 11 n.

The body surface has hairs and refractile points

as in the female.

The rostrum resembles that in the female.

The first antenna is like that of the female, but

there are three additional long aesthetes (at the locations indicated by arrows in Figure I Mr), so that

the formula is 4, 13 -f- 2 aesthetes, 6, 3 -}- 1 aesthete, 4 -f 1 aesthete, 2 -f 1 aesthete, and 7 -f- 1

aesthete.

The second antenna, labrum, mandible, first

maxilla, and second maxilla are like those of the

female. The maxilliped (Figure 161c) is 4-segmented (assuming that the proximal part of the

claw represents a fourth segment). There are two

setae and a row of short spinules on the second

segment. The claw is 139 n along its axis, including

the small terminal lamella, lacks a distinct division

but has a row of minute spinules, and has the usual

two unequal proximal setae, the larger with a hyaline, weakly sclerotized distal half.

The ventral area between the maxillipeds and

the first pair of legs is like that in the female.

Legs 1-4 are like those of the female. In one

male the left leg 4 (Figure 16Id) is abnormal in

that the second exopod segment bears three inner

setae and is rather indistinctly separated from the

third segment. In another male the left endopod

of this leg has two setae on the third segment

(Figure 16 \e).

Leg 5 (Figure 1616) has a small u nor n amen ted

free segment, 22 x 13 n, with the two terminal setae

36 n and 66 n.
FIGURE 160.—Rakotoa proteus, new species. Female: a, leg 1 and intercoxal plate, anterior (D) ;

b, leg 2 and intercoxal plate, anterior (D) ; c, leg 3 and intercoxal plate, anterior (D) ; d, abnormal endopod of leg S, anterior (E) ; e, leg 4, anterior (D) ; /, abnormal exopod of leg 4,

anterior (D) ; g, right and left endopods of leg 4 of a single female, anterior (F) ; h, right and

left endopods of leg 4 of a single female, anterior (F) ; i, leg 5, dorsal (D) ; /, free segment of

leg 5, lateral (outer) (D) . Scale: D, 0.1 mm; F, 0.02 mm.
288

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 161.—Rakotoa proteus, new species. Male: a, dorsal (A) ; b, urosome, dorsal (B) ; c, maxilliped, inner (E) ; d, abnormal leg 3, anterior (E) ; e, abnormal left endopod of leg 3, anterior (E) ;

/, spermatophores, attached to female, lateral (c). Scale: A, B, 05 mm; E, 0.1 mm, c, 0.2 mm.

Leg 6 (Figure 1616) consists of the usual posteroventral flap on the genital segment bearing two

naked setae, 31 n and 34 i*.

The spermatophore (Figure 161/), attached to

the female in pairs, is elongated oval, 229 x 108 n,

not including the neck.

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—The specific name proteus, from

Proteus, the name of a sea deity who could change

himself into all kinds of shapes, alludes to the

variability of the armature in legs 2-4.

REMARKS.—Rakotoa proteus is remarkable in the

armature of legs 2-4. The normal pattern is shown

in the formula given above. The armature of leg

1 is constant in each of the 28 legs of the 14 specimens (10 $ $, 4 cf c?) collected. The armature

of legs 2-4, however, varies as follows:
NUMBER 127 289

P,

i\

Exp

Enp

Exp

Enp

Exp

in 23 legs

in 5 legs

in 24 legs

in 4 legs

in 23 legs

in 3 legs

in 2 legs

in 19 legs

in 3 legs

in 1 leg

in 1 leg

in 1 leg

in 2 legs

in 1 leg

in 21 legs

in 1 leg

in 1 leg

in 3 legs

in 2 legs

82.15%

17.85%

85.71%

14.29%

82.15%

10.70%

7.15%

67.87%

10.70%

3.57%

5-57%

3.57%

7.15%

3.57%

75.01%

3.57%

337%

10.70%

7.15%

Genus Ravahina Humes and Ho, 1968

DIAGNOSIS.—Body transformed, with an expanded

and swollen prosome. Urosome 5-segmented, slender, and sometimes contractile. Caudal ramus with

six short setae. Rostral area weakly developed.

First antenna 7-segmented, the formula being S,

14, 5, 4, 5, 2 -f- 1 aesthete, and 7 -f 1 aesthete.

Second antenna 4-segmented, with the armature 1,

1, 3, and 6 -J- I, there being a single terminal

claw, but an adjacent seta jointed and spiniform.

Labrum with the two lobes separated by a median cleft. Mandible with the base divided into

two parts by an indentation, the part distal to the

indentation having on its convex side two small

surfifial lobes and a distal serrated fringe, and on

its concave side a row of spines; tip of mandible

forming a small spiniform process. First maxilla

with three elements. Second maxilla of the usual

lichomolgid type. Maxilliped 3-segmented, with

the third segment terminating in a spine or spiniform process.

Legs 1-4 lacking the inner coxal seta and all exopods 3-segmented but with reduced number of

elements on the third segment. Leg 1 endopod

0-0;0-0;I,2 and leg 2 endopod 0-0;0-l;II. Endopods of legs 3 and 4 unarmed and vestigial, consisting of a small knob without clear articulation

with the basis. Leg 5 with a free segment bearing

two terminal setae. Leg 6 represented by two setae

near the area of attachment of each egg sac.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Ravahina tumida Humes and

Ho.

REMARKS.—The male is unknown.

Ravahina tumida Humes and Ho, 1968

FICUKE 162

Ravahina tumida Humes and Ho, 1968a, pp. 378-380, figs.

241-259 [associated with Porites sp. cf. andrewsi Vaughan,

region of Nosy Be, northwestern Madagascar].

FICURE 162.-Ravahina tumida Humes and Ho. Female: a, dorsal; b, lateral; c, mandible; d, kg

4. (From Humes and Ho. 1968a. figs 241, 242, 251, 258.) Length of female, 1.67-1.97 mm.
290 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Schedomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented. Caudal ramus with six moderately short setae. Rostrum

not well developed. First antenna 7-segmented, in

the female with the armature 4, 13, 6, 3, 4 -J- 1

aesthete, 2 -f- 1 aesthete, and 7 + 1 aesthete; in

the male with the formula 4, 13 -(- 2 aesthetes, 6,

3 + 1 aesthete, 4 -f- 1 aesthete, 2 -f 1 aesthete,

and 7 + 1 aesthete. Second antenna 3-segmented

(apparently from a fusion of the original third and

fourth segments). First and second segments with

one seta. Third segment either without a trace of

the inner setae or, at most, with only minute spiniform projections, and with a single terminal claw.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal area distal to the

indentation having on its convex side four slender,

digitiform processes followed by a broadly triangular process and then a serrated fringe and on its

concave side two lobes bearing long spinules; lash

long. Paragnath a small hairy lobe. First maxilla

with three elements. Second maxilla of the usual

lichomolgid form. Maxilliped in the female 3-segmented and with a pointed tip; in the male 4-

segmented (recognizing the proximal part of the

claw as a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with 0-1 ;II, the seta being feathered. Armature

similar in both sexes except for the last segment

of leg 1 endopod in the male being 1,1,4 instead

of 1,5 as in the female. Leg 5 with a free segment

bearing two setae. Leg 6 represented in the female

by the two setae and the process near the area of

attachment of each egg sac, in the male by a ventral posterolateral flap on the genital segment bearing two setae.

Other features as in the species he low.

Associated with madreporarians.

TYPE-SPKCIES.—Schedomolgus arcuatipes (Humes

and Ho).

ETYMOLOGY.—The name is a combination of the

Greek words axeftou (near), alluding to the affinity

of this genus with lichomolgid genera, and

Gender masculine.

Key to Species of the Genus Schedomolgus

FEMALES

First postgenital segment with a prominent posteroventral lobe; second segment of first antenna

without sclerotizations on proximal dorsal surface S. lobophorut

First postgenital segment without a posteroventral lobe; second segment of first antenna with

small sclerotizations on proximal dorsal surface S. arcuatipet

MALES

Second postgenital segment with a prominent posteroventral lobe; free segment of leg 5,

68 x 19 p, with small spinules; second segment of first antenna without sclerotizations on

proximal dorsal surface S. lobophonu

Second postgenital segment without a posteroventral lobe; free segment of leg 5, 48 x 12 p.,

without ornamentation; second segment of first antenna with small sclerotizations on

proximal dorsal surface S. arcuatipes

Schedomolgus arcuatipes (Humes and Ho, 1968)

FIGURE 163

Lichomolgus arcuatipes Humes and Ho, 1968a, pp. 362-364

figs. 79-104 [from the hard coral Acropora palifera (Lamarck), region of Nosy Be\ northwestern Madagascar].

Schedomolgus lobophorus (Humes and Ho, 1968)

Lichomolgus lobophorus Humes and Ho, 1968a, pp. 364-366,

figs. 105-133 [from the hard corals Acropora scherzeriana

Brueggemann, Acropora sp., and Acropora cytherea Dana,

region of Nosy Be. Madagascar].

NEW HOST.—15 $ $, 13 cfcff and 1 copepodid

from the madreporarian Acropora florida (Dana),

in 2 m, Rigili Island, Eniwetok Atoll, Marshall

Islands, 3 July 1969; 3 $ ? , 2 <? cf from Acropora

florida, in 4 m, south end of Parry Island, Eniwetok

Atoll, 5 July 1969. Both collections by AGH.
NUMBER 127 291

FIGURE 16S.—Schedomolgus arcuatipes (Humes and Ho). Female: a, dorsal; b, second antenna;

c, mandible; d, second maxilla; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes and

Ho, 1968a. figs. 79, 85, 87. 90. 96. 98.) Length of female 1.06 mm, of male 1.23 mm.

Genus Seivellochiron Humes, 1969

DIAGNOSIS.—Body cyclopiform. Urosome 5-segmented in the female, 6-segmented in the male.

Caudal ramus with six setae. Rostrum broadly

triangular with a rounded tip. First antenna 7-

segmented, the armature in the female being 4,

13, 6, 3, 4 -|- 1 aesthete, 2 -+- 1 aesthete, and 7

-|- 1 aesthete; in the male, 4, 13 -f- 2 aesthetes, 6,

3 -\- 1 aesthete, 4 + 1 aesthete, 2 + 1 aesthete,

and 7 -f- 1 aesthete. Second antenna 3-segmented

(the original third and fourth segments fused),

with the formula 1,1,3 -f- I and 6 setae, there being one terminal claw (but one of the six setae

almost clawlike).

Labrum with a deep, median cleft between the

two lobes. Mandible with the concave side having

a deep incision followed by a row of spinules and

the convex side having a scalelike area with a row

of spinules, followed by a hyaline spike, and then

by a row of toothlike serrations. Paragnath a small

hairy lobe. First maxilla with four setae. Second

maxilla of the usual lichomolgid type. Maxilliped

of the female 3-segmented, the third segment indistinctly set off from the second and with a spiniform tip. Maxilliped of the male 4-segmented

(recognizing the proximal part of the claw as the

fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which consists of a single segment. Armature of the usual lichomolgidiform pattern, except leg 4 endopod which has two spines and is

distinctly shorter than the exopod in both sexes.

Formula for the male similar except for leg 1 endopod where the third segment is 1,1,5 instead of

1,5 as in the female. Leg 5 in both sexes with a

free segment bearing two terminal elements (a

spine and a seta). Leg 6 in the female represented

by the two setae and the spiniform process near

the area of attachment of each egg sac, in the male

by a posteroventral flap on the genital segment

bearing two setae and a minute spiniform process.

Other features as in the species below.

Associated with scyphozoan medusae.

TYPE-SPECIES.—Sewellochiron fidens Humes.
292 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Seuellochiron fidens Humes, 1969

FICURE 164

Sewellochiron fidens Humes, 1969e, pp. 172-182, figs. 1-27

[associated with the medusa Cassiopea xamachana R. P.

Bigelow, near La Parguera, southwestern Puerto Rico].

Genus Spaniomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform or modified. Urosome in the female 5-segmented, in the male 6-

segmented. Caudal ramus with six short setae, the

two median terminal ones expanded. Rostral area

without a posteroventral border but with long

setules. First antenna 7-segmented, in the female

with the formula 1, 13, 6, 3, 4 -f- 1 aesthete, 2 + 1

aesthete, and 7 -f 1 aesthete; in the male with the

armature 1, 13 -f 2 aesthetes, 6, 3 -f- 1 aesthete,

4 -}- 1 aesthete, 2 -f 1 aesthete, and 7 -f- 1 aesthete. Second antenna 3-segmented (apparently

the result of fusion of the original third and fourth

segments). First and second segments with one seta.

Third segment with three small vestigial elements

and terminally bearing a single small claw plus

several minute elements.

Labrum with the two lobes separated by a deep

cleft. Mandible with the basal area beyond the

indentation having on its convex side a large hyaline expansion followed by a serrated fringe and

on its concave side rows of spinules; lash long.

Paragnath a small hairy lobe. First maxilla with

four elements. Second maxilla of the usual lichomolgid form. Maxilliped in the female 3-*egFICUKE 164— Sewellochiron fidens Humes. Female: a, dorsal; b, rostrum; c, second antenna; d,

mandible; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes, 1969e, figs. 1, 7, 10, 12,

21, 23.) Length of female 1.84 mm, of male 1.43 mm.
NUMBER 127 293

merited with a pointed tip, in the male 4-segmented

(assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is 2-segmented. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 0-1 ;II, the seta being feathered.

Leg 1 endopod of the male with the third segment

1,1,4 or 1,5. Leg 5 with a free segment bearing

two setae. Leg 6 in the female represented by the

two setae and the process near the area of attachment of each egg sac, in the male by the ventral

posterolateral flap on the genital segment bearing

two setae.

Other features as in the species below.

Associated with madreporarians.

TYPE-SPECIES.—Spaniomolgus compositus (Humes

and Frost, 1964).

ETYMOLOGY.—The name is a combination of the

Greek words OJUXVIOS (few, or scanty), referring to

the presence of only one seta on the first segment

of the first antenna, and \u>\yo<;. Gender masculine.

Key to Species of the Genus Spaniomolgus

FEMALES

1. Prosomc unusually broad and thickened, ratio 1.07:1, urosome slender; caudal ramus

2.87:1; outer spines on leg 4 exopod with smooth lamellae S. crassus

Prosome not unusually broad, ratio 1.27:1 or 1.38:1; caudal ramus 5:1 or 9.1:1; outer

spines on leg 4 exopod with barbed lamellae 2

2. Caudal ramus 9.1:1; free segment of leg 5, 147 x 14 y. (10.5:1) S. geminus

Caudal ramus 5:1; free segment of leg 5, 1S5 x 17 fi (7.9:1) S. compositus

MALES

1. Third segment of leg 1 endopod with 1,5 as in female S. crassus

Third segment of leg 1 endopod with 1,1,4 2

2. Caudal ramus 7.4:1; middle spine on leg 2 endopod modified, with truncated tip S. geminus

Caudal ramus 4.7:1; middle spine on leg 2 endopod not modified S. compositus

Spaniomolgus compositus (Humes and Frost,

1964)

FICURE 165

Lichomolgus compositus Humes and Frost, 1964, pp. 1S7-139,

figs. 66-98 [from the hard coral Seriatopora subseriata

Ehrenberg, Nosy Bi'-. northwestern Madagascar].—Bouligand,

1966, p. 269-Humes and Ho, 1968a, p. 371 [from the

hard coral Seriatopora octoptera Ehrenberg and Seriatopora sp., region of Nosy Be, Madagascar].

N EW RECORD.—103 $ $ , 28 o*cf» and 6 copepodids from Seriatopora sp., in 2 m, Ambariobe, a

small island almost between Nosy Komha and Nosy

Be, Madagascar, 25 May 1967, collected by AGH.

Spaniomolgus crassus (Humes and Ho, 1968)

Lichomolgus crassus Humes and Ho, 1968a, pp. 368-371, figs.

151-182 [from the hard corals Stylophora pistillata (Esper),

Stylophora mordax (Dana), and Acropora sp., region of

Nosy Be, Madagascar].

Spaniomolgus geminus (Humes and Ho, 1968)

Lichomolgus geminus Humes and Ho, 1968a, pp. 366-368,

figs. 134-150 [from the hard corals Stylophora pistillata,

Stylophora mordax, and Acropora sp., region of Nosy Be,

Madagascar].

NEW RECORD.—26 $ $, 24 tf c?» and 16 copepodids from Stylophora sp., in 4 m, Antsamantsara,

Nosy Be, Madagascar, 6 June 1967, collected by

AGH.

Genus Stellicola Kossmann, 1877

DIAGNOSIS.—Body cydopiform, in the female

with a broad prosome and relatively short urosome.

Urosome in die female 5-segmented, in the male

fundamentally 6-segmented but with the segment

of leg 5 fused with the genital segment. Genital

segment in the female wider than long. Caudal

ramus with six setae. Rostrum broadly rounded or
294

SMITHSONIAN CONTRIBUTIONS TO ZOOLOT.v

FIGURE 165.—Spaniomolgus compositus (Hume* and Frost). Female: a, dorsal; b. caudal ramus;

c, second antenna; d, mandible; e, leg 4 and intercoxal plate; /, endopod of leg 4. Male: g, dorsal.

(From Humes and Frost, 1964, figs. 66, 70, 75, 77. 87, 88, 90.) Length of female 1.58 mm. of

male 1.21 mm.

truncated. First antenna 7-segmented, in both

sexes with the armature 4, 13, 6, 3, 4 -f- 1 aesthete,

2 -}- 1 aesthete, and 7+ 1 aesthete. Second antenna 3-segmented, the third segment being the

result of fusion of two original segments; formula

1,1, and 3 -f

 on e

 terminal claw -|- several small

elements.

Labrum deeply incised medially. Mandible with

the basal area distal to the constriction having on

its convex side a row of spinules or a serrated

fringe and on its concave side a row of spinules;

lash long. Paragnath a small hairy lobe. First maxilla with four elements. Second maxilla with the

initial few teeth on the lash enlarged and with an

unusually strong, spinose, auxiliary element. Maxilliped in the female 3-segmented and with pointed

tip; in the male 4-segmented (assuming that the

proximal part of the claw represents a fourth segment) except in L. femineus (see below).

Legs 1-4 with 3-segmented rami, except leg 4

endopod which is 2-segmented. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

0-1; 11,1 or, (in 5. pollex) 0-0;II. Basis of leg 4 in

both sexes laterally elongated. No sexual dimorphism in the formula of leg 1 endopod. Leg 5 with

a free segment with two terminal elements. Leg 6

represented by the two setae near the genital openings.

Other features as in the species below.

Associated mainly with asteroid echinoderms.

TYPE-SPECIES.—Stellicola thorelli Kossmann.

REMARKS.—The males of four species are unknown.
NUMBER 127 295

Key to Species of the Genus Stellicola

FEMALES

1. Endopod of leg 4 with formula O-O;II $. pollex

Endopod of leg 4 with formula O-1;II,1 2

2. Genital segment with two posterior dorsal sclerotized elevations S. caenHew

Genital segment without such elevations 3

3. Free segment of leg 5 a little more than 3:1 4

Free segment of leg 5 less than 3:1 5

4. Body length 1.41 mm; genital segment 135 x 239 /*, much wider than long; several setae on

first antenna with short hairs S. femineus

Body length 1.00 mm; genital segment ISO x 177 /*, wider than long; all setae on first

antenna naked s. illgi

5. Ratio of body length to width about 1.28:1 $. Umgkaudatus

Ratio of body length to width about 13:1 6

6. Two setae on segment of leg 5 about equal in length 7

Two setae on segment of leg 5 distinctly unequal in length 8

7. Basis of leg 4 greatly elongated, ratio of its length to length of exopod about 135:5

S. pUurobranchi

Basis of leg 4 not as greatly elongated, ratio of its length to length of exopod about 9:5

S. oreastHphOm

8. Postgenital segments and caudal rami together more than four times as long as genital

segment S. thoretti

Postgenital segments and caudal rami together only a little longer than genital segment or

shorter than that segment 9

9. Postgenital segments and caudal rami together a little longer than genital segment

S. cttrttcaudatus

Postgenital segments and caudal rami together a little shorter than genital segment

S. helothuriae

KNOWN MALES

1. With thumblike process on third segment of second antenna; formula of endopod of leg

4. O-0-.II S. p&Uex

Without such a thumblike process; formula of endopod of leg 4, 0-1 ;II,1 2

2. Maxilliped with last segment having a serrated truncated tip instead of being attenuated

and dawlike S. iOgi

Maxilliped with last segment being pointed and clawlike 3

3. Maxilliped female-like, with terminal spiniform process instead of a claw S. feminent

Maxilliped with a distinctly prehensile claw 4

4. Maxilliped elongated, 4-segmented, with long prehensile claw S. oremstriphihu

Maxilliped small, 2-segmented, with relatively small but prehensile daw 5

5. First segment of second maxilla with three peculiar sclerotized processes S. caendeus

First segment of second maxilla without such processes $. fcolothnrfae

Stellicola thorelli Kossmann, 1877

Stellicola thorelli Kossman, 1877, pp. 11-13, pi. 1, fig. 1

[from the asteroid Ophidiaster multiflorit M. and Tr., Red

Sea].-Stock, 1957, p. 382-Humes and Ho, 1967e. p. 222.

REMARKS.—The male is unknown.

Stellicola caendeus (Stebbing, 1900)

Linckiomolgus caeruleus Stebbing, 1900, pp. 664-666, pi.

74B [from the asteroid Linckia, Feather Island, China

straits, eastern end of New Guinea].—Clark, 1921, p. 6S6.

Stellicola semperi.-Monod and Dollfus, 1932a, p. 142.

Stellicola caeruleus.-Stock, 1957, p. 382.-Hume« and Ho,

1967e, pp. 219-221, pi. 15, figs. 121-128; pi. 16. figs. 129-

134; pi. 17, figs. 135-141.

Stellicola curticaudatus (Thompson and A. Scott,

1903)

Paralichomolgus curticaudatus.—Thompson and A. Scott,

1903, pp. 281, 282, pi. 16, figs. 1-7 [from washing of dredged

invertebrates, Ceylon].-A. Scott, 1909, p. 267 [Sibogp

Station 226, midway between the Ludpara and Schildpad

Islands].
2% SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Lichomolgus (Stellicota) curticaudatus.—Monod and Dollfus,

1932a, p. 139.

Stellicola curticaudatus.—Stock, 1957, p. 382.-Humes and

Ho, 1967e, p. 222.

REMARKS.—The male is unknown.

Stellicola femineus Humes and Ho, 1967

Stellicola femineus Humes and Ho, 1967c, pp. 213-215, pi.

8, fig. 67; pi. 9, figs. 68-75; pi. 10, figs. 76-84; pi. 11,

figs. 85-92 [from the asteroids Leiaster leachi Gray and

Leiaster speciosus von Martens, region of Nosy Be, Madagascar].

REMARKS.—The male of this species is unusual

in having a female-like maxilliped, without a long

terminal claw.

Stellicola holothuriae (Ummerkutty, 1962)

Lichomolgus holothuriae Umraerkutty, 1962, pp. 52-55, pi.

8, figs. 4-12, pi. 9, figs. 1-5 [from holothurian washings,

Gulf of Mannar, southeastern India].

Stellicola holothuriae.—Humes and Ho, 1967c. p. 222.

Stellicola illgi, new species

FIGURES 166-168

TYPE MATERIAL.—104 99 , 14 rfcf from the

sea star Linckia laevigata (Linnaeus), Urukthapel,

Palau Islands, 21 July 1955; USNM 206221. Holotype 9 , allotype, and 109 paratypes (99 99 ,

1 0

 c?cf) deposited in USNM, the remaining paratypes (dissected) in the collection of AGH.

OTHER MATERIAL EXAMINED.—226 9 9 , 262 <$ tf,

and 8 copepodids from 10 specimens of Linckia

laevigata, in 2 m, behind reef at Nukumbutho

Passage, southeast of Suva, Viti Levu, Fiji, 13 August 1971, collected by AGH. These copepods were

readily visible crawling over the body surface of

the sea stars.

FEMALE.—The body (Figure 166a) has a broad,

flattened prosome and relatively short urosome.

The length (not including the setae on the caudal

rami) is 1.00 mm (0.94-1.06 mm) and the greatest

width 0.74 mm (0.72-0.80 mm), based on ten specimens in lactic acid. The ratio of the length to the

width of the prosome is 1:1. The segment of leg

1 is fused with the cephalosome. The epimera of

the segments of legs 1-3 are characteristically

formed as illustrated in the figure. The segment

of leg 4 is small, without expanded epimera, and

partly covered in dorsal view by the tergum of the

preceding segment. The ratio of the length of the

prosome to that of the urosome is 2.18:1.

The segment of leg 5 (Figure 1666) is 112 x

195 n. There is no ventral intersegmental sclerite

between that segment and the genital segment.

The genital segment is 130 x 177 u,, wider than

long. The areas of attachment of the egg sacs are

situated ventrolaterally in the posterior part of the

segment. Each area (Figure 166c) bears a plumose

seta, 44 n, mounted on an elongated, well-sclerotized base (shown in detail in Figure 166a"), a

unilaterally spinulose seta, 27 n, and a small spinous

process. The neck of the egg sac enters the segment ventral to the two setae. The three postgenital segments are 42 x 78 u, 35 x 70 u, and

11 x (id (i (greatest width). The posterodorsal margin of the anal segment bears on each side a row

of very small spinules.

The caudal ramus (Figure 166e) is short, 29 x

23 u in greatest dimensions. The outer lateral seta

is 65 ii and naked. The dorsal seta is 50 n and

naked. The outermost terminal seta is 130 n with

lateral spinules, those on the proximal inner side

being especially strong. The innermost terminal

seta is 114 n with strong proximal outer spinules.

The two median terminal setae are 315 H (outer)

and 528 n (inner), both weakly haired and inserted dorsally to a small ventral flap with a row

of minute spinules. There is a prominent setule

on the proximal outer edge of the ramus.

The body surface bears a few small hairs (sensilla) as indicated in Figure 166a,6.

The egg sac (Figure 166a) is elongated, 485-500

x 230 n, broadest distally, and tapering proximally

near the attachment. The numerous eggs are about

56 11 in diameter.

The rostrum (Figure 166/) is broad with slightly

angular lateral margins.

The first antenna (Figure 167a) is 331 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 13 (47 n

along the anterior margin), 138, 42, 49, 26, 15, and

14 n respectively. The formula for the armature is

4, 13, 6, 3, 4 + 1 aesthete, 2 -+- 1 aesthete, and

7 -}- 1 aesthete. All the setae are naked.

The second antenna (Figure 1676) is 330 n long

and 3-segmented, without a trace of division of

the third segment. The first two segments bear an
NUMBER 127 297

FIGURE 166.—Stellicola illgi, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ; c, area of

attachment of egg sac, ventral (c) ; d, base of seta of leg 6, dorsal (c); e, caudal ramus, dorsal

(c); e, caudal ramus, dorsal (D) ; /, rostrum, ventral (B) . Scale: A, 0.5 mm; B, 0.2 mm; c, D,

0.05 mm.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

d

FICUKE 167.—Stellicola illgi, new species. Female: a, first antenna, dorsal (E) ; b, second antenna,

posterior (E) ; c, claw on second antenna, flat view and anterior (c); d, labrum, ventral (E); e,

mandible, posterior (F) ; /, first maxilla, posterior (c); g, second maxilla, anterior (F) ; h,

maxilliped, posterior (F) ; i, area between maxillipeds and first pair of legs, ventral (B) ; ;, leg

1 and intercoxal plate, anterior (E) . Scale: B, 02 mm; c, 0.05 mm; E, F, 0.1 mm.
NUMBER 127 299

unusually long seta. The last segment, 148 n along

its outer edge, 94 n along its inner edge, and 31 n

wide, bears three inner setae; terminally there is a

single claw, 70 n along its axis (turned ventrally

in Figure \67b but shown in flat view in Figure

167c), and five setae. All the setae are naked.

The labrum (Figure 167rf) has two divergent

posteroventral lobes.

The mandible (Figure I67e) bears a serrated

fringe along its convex margin; the concave margin

is not noticeably indented and bears a row of spinules which continues on the posterior surface of

the base. The lash is moderately long and bears

small lateral spinules. The paragnath is a small

lobe with a few hairs, located medially to the first

maxilla. The first maxilla (Figure 167/) bears four

elements. The second maxilla (Figure 167g) has a

relatively small, unarmed first segment. The second

segment bears a surficial posterior barbed seta and

a large inner spine with strong unilateral spinules.

The terminal lash is short (about as long as the

inner spine) and bears strong teeth and a posterior

row of spines. The maxilliped (Figure 167/t) is 3-

segmented. The second segment bears two small

setae. The third segment has a hyaline setule and

a small spine and terminates in a spiniform process. All the elements are naked.

The ventral surface between the maxillipeds and

the first pair of legs (Figure 167/) is not protuberant.

Legs 1-4 (Figures 167/, 168a-c) have 3-segmented

rami except for the endopod of leg 4 which is 2-

segmented. The spinal and setal formula is as

follows:

Pt coxa 0-1 basis 1-0 exp 1-0; 1-1

enp 0-1; 0-1

P, coxa 0-1 basis 1-0 exp 1-0; 1-1;

enp 0-1; 0-2;

P, coxa 0-1 basis 1-0 exp 1-0; 1-1;

enp 0-1; 0-2;

P4 coxa 0-1 basis 1-0 exp 1-0; 1-1;

enp 0-1; II.l

111,1,4

1,5

IH.I.5

I.II.3

IH.I3

I.H.2

11,1,5

The inner coxal seta of legs 1-3 is long and plumose but in leg 4 this seta is minute (6 n) and

naked. The basis becomes progressively more

elongated laterally until in leg 4 it is about 5.5

times longer than wide. The exopod of leg 4 is

58 fi long. The first segment of the endopod is

25 x 14 f*, with its inner distal naked seta 30 n.

The second segment is 31 x 13 n, with its two

barbed spines 22 n (outer) and 61 n (inner) and

its inner plumose seta 66 n. Hairs are present on

the outer margins of both segments and on the

inner margin of the second segment.

Leg 5 (Figure 168d) has an unornamented free

segment, 50 x 15 M- (greatest length, width at middle). The two terminal elements are 15 n (finely

barbed) and 127 n (naked). The adjacent seta on

the body is 55 n and plumose.

Leg 6 is represented by the two setae near the

attachment of each egg sac (Figure 166c).

The color in life is unknown.

MALE.—The body (Figure 168c) in general outline is similar to that of the female. The length

(without the ramal setae) is 0.60 mm (0.55-0.64

mm) and the greatest width 0.43 mm (0.41-0.43

mm), based on seven specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.02:1. The ratio of the length of the prosome to

that of the urosome is 1.64:1.

The segment of leg 5 is fused with the genital

segment (Figure 168/), the combined segment being

180 x 138 n. The four postgenital segments are 27

x 60 n, 24 x 54 n, 22 x 46 n, and 23 x 41 n

(width at the middle) from anterior to posterior.

The caudal ramus resembles that of the female

but is smaller, 16 x 15 n, and of slightly different

proportions.

The rostrum, first antenna (without added aesthetes), and second antenna are like those of the

female.

The labrum (Figure 168g) has more elaborately

outlined lobes than in the female.

The mandible, paragnath, first maxilla, and second maxilla are similar to those of the female.

The maxilliped (Figure 168/J) is small, 4-segmented, and somewhat twisted. The first segment

is unarmed. The second segment bears two small,

naked setae and a hyaline crest with a serrated

edge. The small third segment is unarmed. The

elongated fourth segment is not clawlike but bears

a long, naked seta and has a serrated tip.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are like those of the female.

Leg 5 is similar to that of the female but the

free segment is smaller, 22 x 8 n.

Leg 6 (Figure 168i) is a posteroventral flap on

the genital segment bearing two naked setae, 52 n

and 28 n.
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 168.—Stellicola illgi, new species. Female: a, leg 2, anterior (E) ; b, third segment of

endopod of leg 3, anterior (E) ; c, leg 4 and intercoxal plate, anterior (r) ; d, leg 5, ventral (F) .

Male: e, dorsal (A) ; /, urosome, dorsal (E) ; g, labrum, ventral (c); h, maxilliped, postcro outer

(c); i, leg 6, ventral (F) . Scale: A, 0.5 mm; c, 0.05 mm; E, F, 0.1 mm..
NUMBER 127 301

The spermatophore was not observed.

The color in life is unknown.

ETYMOLOGY.—This species is named for Dr. Paul

L. Illg, of the University of Washington, who most

generously sent us the specimens for study.

REMARKS.—Stellicola illgi may be distinguished

from other species in the genus by a combination

of the following characters: the short lash on the

second maxilla, the second endopod segment of

leg 4 with two spines and a plumose seta, the size

and form of leg 5, and the nature of the male

maxilliped (without a true claw).

Stellicola longicaudatus (Thompson and A. Scott,

1903)

Paralichomolgus longicaudatus Thompson and A. Scott, 1903,

p. 282, pi. 20, figs. &-8 [from washings of dredged invertebrates, Ceylon].

Stellicola longicaudatus.—Humes and Ho, 1967e, p. 222.

REMARKS.—The male is unknown.

Stellicola oreastriphilus Kossmann, 1877

Stellicola oreastriphilus Kossman, 1877, p. 13, pi. 2, fig. 1

[from the asteroid Asteropsis carinifera M. Tr., Red Sea].—

Humes and Ho, 1966b, pp. 100, 101 [from the asteroid

Choriaster granulatus (Lutken), region of Nosy Re, Madagascar]; I967e, pp. 218, 219, pi. 14, figs. 115-120 [from the

asteroid Protoreaster nodosus (Linnaeus), Nosy Be\ Madagascar].

Lichomolgus (Stellicola) oreastriphilus.—Humes and Cressey,

1961c. pp. 83-92, figs. 1-31 [from the asteroids Protoreaster

lincki (Blainville), Culcita schmideliana (Rctzius), Pentaceraster mammillatus (Audouin), and Poraster superbus

(Mobius), region of Nosy Be\ Madagascar].

Stellicola plewrobranchi Kosmann, 1877

Stellicola pleurobranchi Kossmann, 1877, pp. 15-16, pi. 3,

fig. 3 [from the tectibranch Pleurobranchus, Palau Archipelago].—Aurivillius, 1883, p. 3.—Stock, 1957, p. 382.—

Humes and Ho, 1967e, p. 222.

Lichomolgus (Stellicola) pleurobranchi.—Monod and Dollfus,

1932a, p. 143.

REMARKS.—The male is unknown.

Stellicola pollex Humes and Ho, 1967

StelUcola pollex Humes and Ho, 1967e, pp. 216-218, pi. 2,

fig. 93; pi. 12, figs. 94-101; pi. 13, figs. 102-109; pi. 14,

figs. 110-114 [from the asteroid Linchia laevigata (Linnaeus) , M'bV S, 47°46' 30" E, Nosy Ovy, southwest of Nosy

Be, Madagascar].

NEW HOSTS.—From three specimens of Linckia

diplax (Muller and Troschel): 2 $ ? , 1 cf;

9 ? o , 39 cf ^ ; and 6 9 9 , 2 o"o*. 2 copepodids; subtidal, Hookena, Hawaii, 2 January 1959,

collected by P. L. Illg.

From ten specimens of Linckia multiflora (Lamarck): 3 ? ? , 2 cfcT. Hawaiian Islands, 13 December 1958, collected by A. H. Banner.

From a single specimen of Linckia guildingi

Gray: 3 9 9, 5 o* cf» 2 copepods, subtidal, Hookena, Hawaii, 2 January 1959, collected by P. L.

Illg.

Genus Synstellicola Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform, the prosome not

greatly widened, die urosome relatively long. Urosome in the female 4-segmented, in the male 5-

segmented (though in S. kossmanni and 5.

acanthasteris the anal segment in both sexes has

an indication of division). Caudal ramus with six

setae. Rostrum rounded. First antenna 7-segmented, in both sexes with the armature 4, 13, 6,

3, 4 -f- 1 aesthete, 2 -j- 1 aesthete, and 7+ 1 aesthete. Second antenna 3-segmented, the third segment the result of fusion of two original segments

(this segment sometimes showing an incomplete

division line), the formula being 1, 1, and 3 —|— 1

terminal claw -\- several small elements.

Labrum deeply incised medially. Mandible with

the basal area distal to the indentation having on

its convex side a row of teeth and on its concave

side a row of spinules; lash long. Paragnath a

small hairy lobe. First maxilla with four elements.

Second maxilla with the initial few teeth on the

lash often enlarged. Maxilliped in the female 3 -

segmented and with a pointed tip, in the male 4—

segmented (considering the proximal part of the

claw to represent a fourth segment).

Legs 1-4 with 3-segmented rami, except leg 4

endopod which is 2-segmented. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

0-l;II,l. No sexual dimorphism in the formula of

leg 1 endopod. Leg 5 with a free segment bearing

two terminal elements. Leg 6 represented by the

two setae near the genital openings.
3 0 2 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Other features as in the species below. ETYMOLOGY.—The generic name is formed from

Associated mainly with asteroid echinoderms. the

 Gre<* word <TM (together) and Stellicola. GenTYPE-SPECIES.—Sym*ellicola affinis (Humes and

 de r

 masculine.

H . REMARKS.—The male of Synstellicola lankensis

is unknown.

Key to Species of the Genus Synstellicola

FEMALES

1. Areas of attachment of egg sacs near middle of genital segment 2

Areas of attachment of egg sacs well behind middle of genital segment 5

2. Caudal ramus short, nearly quadrate S. acanthasteris

Caudal ramus about 2 times longer than wide, or at least 4 times longer than wide 9

3. Caudal ramus at least 4 times longer than wide 4

Caudal ramus about 2 times longer than wide S. Umkeruh

4. Caudal ramus 6.3-7 times longer than wide S. affmit

Caudal ramus 4 times longer than wide S. gracilit

5. Seta on first segment of second antenna unusually long, reaching nearly to end of appendage;

free segment of leg 5 swollen on inner margin S. longlseUt

Seta on first segment of second antenna not ususually long; free segment of leg 5 with tidei

nearly parallel 6

6. Genital segment in dorsal view with lateral margins anterior to genital openings nearly

parallel and with anterior "shoulders"; shorter of two elements on free segment of leg 5

with delicately serrate lamellae S. koummnni

Genital segment in dorsal view with lateral margins recurved, without anterior "shoulders";

shorter of two elements on free segment of leg 5 with short spinules S. pichoni

KNOWN MALES

1. Anal segment with annular suture 2

Anal segment without an annular suture S

2. Genital segment in dorsal view not globose, with lateral margins only slightly rounded;

longer of two setae on free segment of leg 5 and on leg 6 very long (equal to about fourfifths the width of genital segment) S. houmamd

Genital segment in dorsal view globose, with lateral margins strongly rounded; longer of

two setae on free segment of leg 5 and on leg 6 short (less than one-fifth the width of

genital segment) S. acanthasteris

3. Caudal ramus distinctly elongated 4

Caudal ramus nearly quadrate 5

4. Caudal ramus 7.5 times longer than wide S. affinis

Caudal ramus about 4 times longer than wide S. gracilis

5. Seta on first segment of second antenna not unusually long; claw of maxilliped evenly

recurved; free segment of leg 5 nearly 4 times longer than wide S. pichoni

Seta of first segment of second antenna unusually long; claw of maxilliped with two slight

flexures; free segment of leg 5 about 2 times longer than wide S. longuela

Synstellicola affinis (Humes and Ho, 1967) Synstellicola acanthasteris (Humes, 1970)

FIGURE 169 Stellicola acanthasteris Humes, 1970b, pp. 329-337, figs. 1-30

c# .»• . a- . „ , „ ,_._ „„„ „„„ [from the asteroid Acanthaster planci (Linnaeus), Eniwetok

Stellicola affinis Humes and Ho, 1967e, pp. 206-209, pi. 4,

 L

A. „ . . . „ -, . ,

 r K '

«;«» 9t_»i i K c ,„ ,, , c I .„ .„ r,

 F

 , Atoll, Marshall Islands].

figs. 25-31, pi. 5, figs. 32-41, pi. 6, figs. 42-48 [from the

asteroids Maculaster maculata maculata (Miiller and Tros- T , ,

chel), Maculaster savingnyi (Audouin), Retaster cribrosus NE W RECORDS (*»

 fro m

 Acanthaster planet, colvon Martens, and Luidia maculata Muller and Troschel, lected by AGH).—390 $ °., 657 cfcf, and 29 COregion of Nosy Be, Madagascar]; 1970b, p. 337. pepodids, from eight hosts (diameter about 20 cm),
NUMBER 127 303

FICURE 169.-.s>mttllicola affinis (Humes and Ho). Female: a, dorsal; b, second antenna; c,

mandible; d, second maxilla; e, leg 4 and intercoxal plate. Male: /, dorsal. (From Humes and

Ho, 1967e. figs. 25. 32. 34. 36, 42, 44.) Length of female 1.01 mm, of male 0.78 mm.

in 3 in, behind reef at Nukumbutho Passage, southeast of Suva, Viti Levu, Fiji, 13 August 1971; 18

9 ?» •1 <S d\ from single host, in 4 m, west of I.

Ngou, northwest of Noumea, New Caledonia,

22°13'44" S, 166o23'01" E, 3 August 1971; 7 $ ?,

7 cf o*» from single host, in 2 m, Ricaudy Reef,

near Noumea, New Caledonia, 22° 19W S,

166°26'28" E, 20 July 1971.

Synstellicola gracilis (Thompson and A. Scott,

1903)

Lichomolgus gracilis Thompson and A. Scott, 1903, p. 278,

pi. 15, figs. 1-9 [from washings of dredged invertebrates,

Ceylon].

stellicota gracHis.-Stock, 1957, p. 382.—Humes and Ho,

1966b, p. 100; 1967c, pp. 223. 224.-Humes, 1970b, p. 337.

Synstellicola lankensis (Thompson and A. Scott,

1903)

Lichomolgus lankensis Thompson and A. Scott, 1903, p. 279,

pi. 15, figs. 25, 26 [from washings of dredged invertebrates,

Ceylon].

Stellicola lankensis.—Stock, 1957, p. 382.—Humes and Ho,

1966b, p. 100; 1967e, pp. 209. 223.-Humes, 1970b, p. 337.

REMARKS.—The male is unknown.

Synstellicola longiseta (Humes and Ho, 1967)

Stellicola longiseta Humes and Ho, 1967e, pp. 209-213, pi. 6,

figs. 49, 50. pi. 7, figs. 51-60, pi. 8, figs. 61-66 [from the

asteroid Mithrodia clavigera (Lamarck), region of Nosy Be\

Madagascar].

Synstellicola hossmanni (Humes and Ho, 1967) Synstellicola pichoni (Humes and Ho, 1966)

Stellicola kossmanni Humes and Ho, 1967c, pp. 202-205, pi.

1, figs. 1-6, pi. 2, figs. 7-15, pi. 3, figs. 16-21, pi. 4,

figs. 22-24 [from the asteroid Protoreaster lincki (Blainville), Nosy Be, Madagascar].

Stellicola pichoni Humes and Ho, 1966b, pp. 97-100, figs.

30-59 [from the asteroid Choriaster granulatus (Liitken),

region of Nosy B£, Madagascar]; 1967e, pp. 215. 223, 224.-

Humes, 1970b, p. 337.
304 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Genus Telestacicola Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented.

Caudal ramus with six setae. Rostrum linguiform.

First antenna 7-segmented, in the female with the

armature 4, 13, 6, 3, 4 -f 1 aesthete, 2 + 1 aesthete,

and 7 + 1 aesthete; in the male 4, 13 + 1 aesthete,

6, 3 -f 1 aesthete, 4 + 1 aesthete, 2 + 1 aesthete,

and 7 -f 1 aesthete. Second antenna 4-segmented,

with the formula 1, 1, 3 and two terminal dentate

claws -J- five long setae.

Labrum incised medially. Mandible with the

basal region beyond the indentation having on its

convex side a patch of spinules followed by a row

of broad teeth and on its concave side a row of

spinules; lash long. Paragnath a small hairy lobe.

First maxilla with four elements. Second maxilla

of the lichomolgid type, the auxiliary spine about

as long as the rather short lash which has evenly

graded teeth. Maxilliped in the female 3-segmented, with a pointed tip and two elements; in

the male 4-segmented (assuming that the proximal

part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except for leg 4

endopod which is a single segment. Armature lichomolgidiform in pattern. Leg 4 exopod with the

third segment having 11,1,5. Leg 4 endopod with

11,1, the seta being feathered. Leg 1 endopod in

the male with the formula 1,1,4 instead of 1,5 as

in the female. Leg 5 with a free segment bearing

two terminal elements. Leg 6 represented by two

setae near the genital openings.

Other features as in the species described below.

Associated with telestacean octocorals.

TYPE-SPECIES.—Telestacicola angoti, new species.

ETYMOLOGY.—The generic name, coined from

Telestacea and the Latin -cola (inhabiting), refers

to the association of this copepod with telestaceans.

Gender masculine.

REMARKS.—Several genera of the Lichomolgidae

have a 1-segmented endopod in leg 4. However,

only Kelleria Gurney, 1927, has, as in Telestacicola,

the combination of a 4-segmented second antenna

with two terminal claws and the formula 11,1 on

the fourth endopod, suggesting a close relationship between the two genera.

Telestacicola shows several significant differences

from Kelleria, however. In Gurney's genus the

male has two aesthetes on the second segment of

the first antenna; one of the elements on the third

segment of the second antenna is a slender claw,

and the two claws of this appendage are slender

and not toothed; the spinules on the concave side

of the mandibular base are stout; the lash on the

second maxilla forms an angle with the segment

and bears irregularly sized spines; and the third

segment of the female maxilliped bears four elements.

Although Telestacicola is strikingly similar to

Kelleria in the nature of the fourth leg, it cannot

be included in that genus without an unnatural

extension of the generic diagnosis.

Telestacicola angoti, new species

FICURES 170-172

TYPE MATERIAL.—23 9 9» 29 cf cf. and 5 copej »•)<I ids from single colony of the telestacean

Coelogorgia palmosa Milne Edwards and Haime,

in 3 m, Pte. Lokobe, Nosy R6, Madagascar, 3 June

1967, collected by AGH. Holotype 9 , allotype, and

46 paratypes (20 99 , 26 tf tf) deposited in

USNM, the remaining paratypes (dissected) in the

collection of AGH.

OTHER MATERIAL EXAMINED.—5 9 9» ^ cf cf»

and 7 copepodids from single colony of the gorgonacean Muricella rubra robusta J. A. Thompson

and J. Simpson, in 3 m, Pte. Lokobe, Nosy Be\

Madagascar, 3 June 1967, collected by AGH.

FEMALE.—The body (Figure 170a) is slender and

elongated. The length (not including the setae on

the caudal rami) is 0.92 mm (0.90-0.95 mm) and the

greatest width 0.31 mm (0.30-0.32 mm), based on

ten specimens in lactic acid. The segment of leg

1 is incompletely set off from the cephalosome. The

epimeral areas of the pedigerous segments are

varied, as illustrated in the figure. The ratio of the

length to the width of the prosome is 1.57:1. The

ratio of the length of the prosome to that of the

urosome is 1.08:1.

The segment of leg 5 (Figure 1706) is 60 x 84 |i.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is elongated and in dorsal view not much

expanded, 140 x 91 n. The areas of attachment of

the egg sacs are situated dorsolaterally in the anterior half of the segment. Each area (Figure 170c)

bears two naked setae, 10 n and 35 n, and a small
NUMBER 127 505

FIGURE nQ.—Telestacicola angoti, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ; c,

part of segment of leg 5 and genital segment, lateral (c); d, caudal raraus, dorsal (c); e,

rostrum (D) ; /, first antenna, with arrows indicating positions of additional aesthetes in male,

posterior (E) ; g, second antenna, posterior (E) ; h, terminal claw of second antenna, posterior

(F) . Scale: K, 0.3 mm; B, 0.2 mm; c, 0.05 mm; D, E, 0.1 mm; F, 0.03 mm.
306

SMITHSONIAN CONTRIBUTIONS TO ZOOUM.V

spiniform process. The three postgenital segments

are 49 x 49 n, 39 x 42 n, and 62 x 39 n from anterior to posterior. The posteroventral border of

the anal segment is smooth.

The caudal ramus (Figure 170d) is elongated,

96 x 20 n, or 4.8 times longer than wide. The outer

lateral seta is 52 n and the dorsal seta 25 n, both

of them naked. The outermost terminal seta is 86 n

and the innermost terminal seta 110 n, both haired.

The two long median terminal setae are 176 n

(outer) and 275 n (inner), both haired and inserted

between unornamented dorsal and ventral flaps.

The body surface has a few small hairs (sensilla) as indicated in the figures.

The egg sacs were incomplete in all females seen

(Figure 170a). Each egg is about 60 M. in diameter.

The rostrum (Figure 170e) is a slender linguiform lobe.

The first antenna (Figure 170/) is 231 n long and

7-segmented. The lengths of the segments (measured along their posterior nonsetiferous margins)

are 28 (42 n along the anterior margin), 53, 18, 31,

30, 28, and 29 v respectively. The formula for the

armature is 4, 13, 6, 3, 4 -f- 1 aesthete, 2 -f- 1

aesthete, and 7 -(- 1 aesthete. All the setae are

naked.

The second antenna (Figure 170g) is 4-segmented. Each of the first two segments has a barbed

seta, that on the first segment being slightly spiniform. The third segment has three barbed setae.

The long fourth segment is 77 ^ along its outer

edge, 51 n along its inner edge, and 18 n wide; it

bears two claws 42 n and 47 n along their axes.

Each claw (Figure 170//) is coarsely dentate along

its concave side and bears a small subterminal

process on its convex side. In addition to the two

claws the segment bears five setae, two barbed on

both sides, two barbed only on one side, and one

naked.

The labrum (Figure 171a) has two posterior

lobes separated by a deep median incision. The

mandible (Figure 1716) has on the convex side of

the basal region a group of slender spinules followed by a row of broad teeth; on its concave side

beyond the indentation there is a row of spinules.

The lash is long. The paragnath (see Figure 172c

of the male) is a small hairy lobe. The first maxilla

(see Figure 172c of the male) has four elements.

The second maxilla (Figure 171c) has an unornamented first segment. The second segment bears

a small posterior surficial seta barbed along one

edge and a large inner auxiliary spine with long

spinules. The terminal lash is relatively short, approximately the length of the auxiliary spine, and

bears a unilateral row of small teeth. The maxilliped (Figure 17Id) has an unornamented first segment and two setae on the second segment. The

small third segment bears a spinule and a naked

spine with indistinct articulation and it terminates

in a finely barbed spiniform process.

The ventral area between the maxillipeds and

the first pair of legs (Figure 171 c) is only slightly

protuberant. A scleroti/ed line connects the bases

of the maxillipeds.

Legs 1-4 (Figure 171/-/) have 3-segmented rami

except for the endopod of leg I which consists of a

single segment. The inner coxal seta in legs 1-3

is long and plumose, but in leg 1 this seta is minute

(5 n) and naked. The inner margin of the basis in

legs 2 and 3 is haired, but in legs 1 and 1 this

margin is naked. The spine and sctal formula is

typically lichomolgid in pattern. Leg 4 cxopod is

128 n long, with the third segment having the formula 11,1,5. Leg 1 endopod is 64 x 16 n in greatest

dimensions. The distribution along the outer edge

of long hairs proximally and short spinules distally,

in addition to the location of the short inner plumose seta (12 n), suggests a prior division into two

segments. There is, however, no trace of a division

of the segment. The two terminal fringed spines

are 41 n (outer) and 56 \i (inner).

Leg 5 (Figure 171;) has a free segment 55 x 20 n

in greatest dimensions, with a proximal inner (ventral) expansion. (The free segment of leg 5 shown

in Figure 170c is foreshortened in the particular

position drawn.) The two terminal elements are

26 n and 47 n, both naked. The adjacent seta on

the body is 22 n and naked. The free segment is

ornamented with two groups of spinules, one on

the expansion, the other along the outer (dorsal)

side.

Leg 6 is represented by the two setae and the

process near the area of attachment of each egg sac.

The color in life in transmitted light is translucid, the alimentary tract in the prosome orangered, but in the urosome bluish, the eye red, the

egg sacs pale lavender to brown.

MALE.—The body (Figure 172a) is more slender

than in the female. The length (excluding the

ramal setae) is 0.65 mm (0.59-0.74 mm) and the
NUMBER 127

FIGURE 111.—Telcstacicola angoti, new species. Female: a, labrum, ventral (F) ; b, mandible,

posterior (F) ; c, second maxilla, posterior (F) ; d, maxilliped, posterior (c); e, area between

maxillipeds and first pair of legs, ventral (E) ; /, leg 1 and intercoxal plate, anterior (c); g, leg

2, anterior (E) ; h, endopod of leg 3, anterior (E) ; i, leg 4 and intercoxal plate, anterior (E) ; ;',

leg 5, dorsal (c). Scale: c, 0.05 mm; E, 0.1 mm; F, 0.03 mm.
308 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

greatest width 0.18 mm (0.16-0.19 mm), based on

ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 2.07:1. The

ratio of the length of the prosome to that of the

urosome is 1.20:1.

The segment of leg 5 (Figure 1726) is 29 x 46 (i.

There is no ventral intersegmental sclerite. The

genital segment in dorsal view is 99 x 86 n, with

nearly parallel sides. The four postgenital segments

are 37 x 40 m 33 x 34 n, 24 x 31 n, and 35 x 32 i*

from anterior to posterior.

The caudal ramus resembles that of the female,

but is smaller, 51 x 16 n, and relatively shorter,

with the ratio 3.19:1.

The rostrum is like that in the female.

The first antenna resembles that of the female,

but there are two additional aesthetes, one on segment 2 and the other on segment 4 (their positions

indicated by arrows in Figure 170/), so that the formula is 4, 13 -f 1 aesthete, 6, 3 -f- 1 aesthete, 4

-j- 1 aesthete, 2 -j- 1 aesthete, and 7 + 1 aesthete.

The second antenna, labrum, mandible, paragnath

(Figure 172c), first maxilla (Figure 172c), and second maxilla are similar to those in the female. The

maxilliped (Figure 172d) is 4-segmented (the proximal part of the claw probably representing a

fourth segment). The first and third segments are

unarmed. The second segment bears two setae and

FICURE 172.-Telestacicolo,angoti, new species. Male: a, dorsal (c); b, urosome, dorsal (D) ; c,

mandible and paragnath, anterior (ventral) (F) ; d, maxilliped, inner (c); e, endopod of leg

1, anterior (c) . Scale: c, 0.05 mm; D, 0.1 mm; F, 0.03 mm; c, 0.2 mm.
NUMBER 127 309

two rows of spinules. The claw is 92 n along its

axis, with an incomplete line of division about

midway and with two unequal proximal setae.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the female except for leg 1 endopod (Figure 172c) where

the third segment has 1,1,4 instead of 1,5 as in the

opposite sex.

Leg 5 (Figure 1726) has an unornamented, subrectangular free segment, 22 x 6.5 n, with its two

terminal elements 14 n and 25 n.

Leg 6 consists of a posteroventral flap on the

genital segment bearing two naked setae, 14 n and

30 n.

The spermatophore was not observed.

The color in life is similar to that of the female.

ETYMOLOGY.—This species is named for Dr. Michel Angot, former director of the Centre de l'Office de la Recherche Scientifique et Technique

Outre-Mer (ORSTOM) at Nosy Be, Madagascar.

Genus Xenomolgus Humes and Stock, 1972

DIAGNOSIS.—Body transformed, elongated. Urosome 5-segmented in the female, 6-segmented in

the male. Caudal ramus with six relatively short

setae. First antenna 7-segmented, in the female

with 4, 13, 6, 3, 4 -f 1 aesthete, 2 -f 1 aesthete,

and 7 -f 1 aesthete; in the male with 4, 13 + 2

aesthetes, 6, 3 -(- 1 aesthete, 4 -f- 1 aesthete, 2 +

1 aesthete, and 7+ 1 aesthete. Second antenna

4-segmented, with the armature 1,1,3 and 1,1,5,

having terminally one claw and one long, almost

clawlike seta.

Labrum with the two lobes separated by a median indentation. Mandible with its basal area

having on its convex side a small, hyaline expansion followed by a serrated fringe; its concave side

deeply indented, distally with a transverse row of

long spinules; lash reduced to a very short, naked

spiniform process. Paragnath a small lobe. First

maxilla with three elements. Second maxilla of the

usual lichomolgid type. Maxilliped in the female

3-segmented, with a pointed tip; in the male 4—

segmented (assuming that the proximal part of the

claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Last segment of

exopod of legs 1-3 with 111,1,4; HI,I,4; and 11,1,2

respectively. Last segment of endopod of legs 1-3

with 1,5; 1,11,3; and 11,1. Leg 4 exopod with last

segment 111,1,1. Leg 4 endopod with 0-0;II. Considerable variation in armature of legs 1-4. Inner

coxal seta on all four legs usually absent. Leg 1

in the male without sexual dimorphism. Leg 5

with a small free segment bearing two terminal

setae. Leg 6 represented in the female by the two

minute setae on the genital area, in the male by

a posteroventral flap on the genital segment bearing two setae.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Xenomolgus varius, new species.

ETYMOLOGY.—The generic name is formed from

the Greek words tevos (a guest), referring to the

association with corals, and nofcvoc Gender masculine.

Xenomolgus varius, new species

FIGURES 173-175

TYPE MATERIAL.—291 $ $, 20 <? o", and 6 copepodids from the hard coral Porites sp., in 1-2 m,

lagoon at Pte. du Tamarin, Mauritius, 15 February

1964, collected by JHS. Holotype $, allotype, and

150 paratypes (140 $ <j>, 10 dd) deposited in

ZMA, 137 paratypes (130 ? ? , 7 rfcf) in USNM,

and the remaining paratypes and the copepodids in

the collection of A. G. Humes.

OTHER MATERIAL EXAMINED (collected by JHS).—

8 9 $ from Porites sp., in about 1 m, lagoon at

Baie du Tombeau, Mauritius, 12 February 1964;

4 cf d from Porites sp., Baie du Tombeau, 14

February 1964.

FEMALE.—The body (Figure I73a,b) is elongated

and transformed, with a swollen prosome and

slender urosome. The cephalosome is flattened

dorsally (Figure 173&). The length (not including

the setae on the caudal rami) is 1.69 mm (1.63-

1.76 mm) and the greatest width 0.54 mm (0.52-

0.55 mm), based on ten specimens in lactic acid.

The ratio of the length to the width of the prosome is 1.43:1. The ratio of the length of the

prosome to that of the urosome is 1:1.33, the urosome being longer than the prosome. The segment

of leg 1 is set off from the cephalosome by a weak

dorsal transverse furrow. The pedigerous segments

lack projecting epimera.
310

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 173.—Xenomolgus varius, new species. Female: a, dorsal (A) ; b, lateral (A) ; c, urosome,

dorsal (B) ; d, area of attachment of egg sac, dorsal (c) ; e, caudal ramus, dorsal (D) ; /, rostrum,

ventral (D) ; g, first antenna, dorsal (E) ; h, second antenna, posterior (E) . Scale: A, B, 05 mm;

c, 0.03 mm; D, E, 0.1 mm.
NUMBER 127 311

The segment of leg 5 (Figure 173c) is 104 x 213 n.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is elongated, 237 n x 185 n in greatest

dimensions. The areas of attachment of the egg

sacs are located dorsolaterally and anteriorly. Each

area (Figure 173d) bears two, very small (2-3 n

long) naked setae. The three postgenital segments

are 208 x 166 n, 169 x 148 », and 148 x 146 n from

anterior to posterior. The posteroventral border

of the anal segment is smooth. The genital segment and the first two postgenital segments have a

weakly sclerotized posterior part.

The caudal ramus (Figure 173e) is 164 x 52 n

(the width taken proximally to the lateral seta),

the ratio being 3.1:1. The width at the lateral

seta is 58 n, and distal to that seta it is 38 n. The

outer lateral seta is 42 n, the dorsal seta 52 n, the

outermost terminal seta 40 n, and the innermost

terminal seta 39 n. The two median terminal setae

(shorter than the ramus) are 67 n (outer) and 122 n

pinner;, -vu six setae are naked. Mne dorsal surface

oi i iic ramus bears two refracule knobs. IN ear the

base ot tne lateral seta there is a marginal spur,

conspicuous in lateral view (Figure 1736).

Tne fine ornamentation ot the body surface consists of a few very small hairs (sensilla) and refractiie knobs on the dorsal surface of the urosome

(Figure 173c), a dorsal transverse band of refractile knobs on the segment of leg 1 and another

band on the posterior part of the cephalosome, as

well as a few similar knobs scattered over the anterior extremity of the cephalosome (Figure 173a).

None of the collected females carried intact

egg sacs. One female had three irregular elliptical

eggs attached, each egg about 170 x 130 n.

The rostrum (Figure 173/) is a broad, linguiform

lobe.

The first antenna (Figure 173g) is 7-segmented

and 205 \i long. The lengths of the segments

(measured along their posterior nonsetiferous margins) are 22 (51 n along the anterior margin), 52,

20, 18, 27, 17, and 20 n respectively. The formula

for the armature is 4, 13, 6, 3, 4 -f 1 aesthete, 2 + 1

aesthete, and 7+ 1 aesthete. (One seta on the

third segment is displaced distally, but it is considered to belong to the third rather than the fourth

segment). All the setae are naked.

The second antenna n*i£ure 173h) is 4-segmented. The first two segments bear a seta. The

third seement bears three setae. The fourth segment, 44 fi along its outer side, 33 n along its inner

side and 29 n wide, bears terminally a short stout

claw, 33 ji, a long, recurved proximally inflated seta

(81 »i in length and almost unguiform) with a very

narrow lamella along its concave margin, and five

small setae. All the elements are naked.

The labrum (Figure 174a) has two broad posteroventral lobes separated medially by an oval,

sclerotized area.

The mandible (Figure 174&) has on the concave

side of the base a deep indentation, followed by

a transverse row of about seven long spinules; on

the convex side there is a slight hyaline expansion

followed distally by a serrated fringe; the lash is

reduced to a very short, naked spiniform process.

The paragnath (Figure 174a) is a small, smooth

lobe. The first maxilla (Figure 174c) bears three

elements. The second maxilla (Figure 174d) is 2-

segmented. The first segment is large and unornamented. The second segment is elongated and bears

a small outer (ventral) setule, a surficial posterior

hyaline smooth seta, and an inner (dorsal) seta

with prominent spinules; the lash has unilateral

toothlike spines. The maxilliped (Figure 17e) is

3-segmented; the large first segment is unarmed,

the second segment bears two naked setae, and the

small third segment has a finely barbed spine along

one side and terminates in a naked spiniform

process (possibly a spine, though the articulation

is obscure).

The ventral area between the maxillipeds and

the first pair of legs (Figure 174/) is not protuberant. The median sclerite seen in most lichomolgids

immediately in front of the intercoxal plate of

leg 1 appears to be absent.

Legs 1-4 (Figures 174g-i, 175&) have 3-segmented rami, except for the endopod of leg 4

which is 2-segmented. The formula for the armature of the legs is as follows:

Pj coxa 0-0(0-1) basis 1-0 exp 1-0; 1-1; 111,1,4

(111,1,3)

enp 0-1; 0-1; 13

P, coxa 0-0 basis 1-0 exp 1-0; 1-1

(1-2) ;

111,1,4

enp 0-1; 0-1; 1,11,3

(UU)

P, coxa 0-0(0-1) basis 1-0 exp 1-0; 1-1; 111,1,2

(IIU.3)

(IIU.l)

enp 0-0; 0-1

(0-2);

11.1(1.1)

P4 coxa 0-0(0-1) basis 1-0 exp 1-0; 1-1; IIU.l

enp 0-0; II
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 174.—Xenomolgus varius, new species. Female: a, labrum, with paragnaths indicated by

broken lines, ventral (F) ; b, mandible, anterior (F) ; c, first maxilla, dorsal (F) ; d, second

maxilla, inner (F) ; e, maxilliped, antero-inner (F) ; /, area between maxillipeds and first pair

of legs, ventral (F) ; g, leg 1 and intercoxal plate, anterior (E) ; h, leg 2, anterior (r.); i, leg S,

anterior (E) . Scale: E, 0.1 mm;.F, 0.05 mm.
NUMBER 127 313

FIGURE 175.—Xcnomolgus varius, new species. Female: a, abnormal endopod of leg 3, anterior

(E) ; b, leg 4 and intercoxal plate, anterior (E) ; c, leg 5, lateral (F) . Male: d, dorsal (B) ; e, lateral

(B) ; /, urosome, dorsal (c); g, first antenna, dorsal (F) ; h, maxilliped, inner (E) ; i, leg 6,

ventral (D) . Scale: B, 03 mm; D, F, 0.1 mm; F, 0.05 mm; c, 02 mm.
314 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

In the above formula the armature shown in

parentheses (observed in only a few instances)

is believed to be abnormal. An example of abnormal armature is shown in leg 3 endopod (Figure

175a). Whenever it occurred, such abnormal armature was present on only one leg of an individual,

not symmetrically on both legs of the pair (except

in the case of the inner coxal setae shown in leg

4, Figure 1756).

The inner coxal seta apparently is normally

absent in this species. Eleven females showed no

trace of such a seta. Three females possessed the

seta as follows: 1 $ with the seta on right leg 1,

1 $ with the seta on right leg 3, and 1 $ with the

seta on right leg 3 and on both right and left leg

4. The inner margin of the basis is naked and

weakly sclerified in all four legs.

The outer spines on the exopods of legs 1-4 have

well-formed subterminal flagella; the lateral spinules are more strongly developed in leg 1 than

in the following legs. The second segment of the

endopod in legs 1-3 has a slender, outer distal

spiniform process, in leg 1 recurved and longer

than in the other two legs. Leg 4 is 86 »i long.

The first segment of the endopod is 16 x 14 n, the

second segment 26 x 17 j*. with the outer spine

19 n, the inner 28 i*.

Leg 5 (Figure 175c) has a small, u norm men ted

free segment 27.5 x 13 n, with its two naked

terminal elements 32 n and 24 n. The seta on the

body near the free segment is 43 n and naked.

Leg 6 is represented by the two minute setae on

the area of attachment of each egg sac (Figure

173d).

The color in life is unknown.

MALE.—The body (Figure 175d^) is slender,

with the cephalosome not swollen or flattened

dorsally as in the female. The length (without

the ramal setae) is 1.12 mm (1.03-1.21 mm) and

the greatest width 0.33 mm (0.13-0.34 mm), based

on ten specimens in lactic acid. The ratio of the

length to the width of the prosome is 1.63:1. The

ratio of the length of the prosome to that of the

urosome is 1:1.28, the urosome being longer than

the prosome.

The segment of leg 5 (Figure 175/) is 39 x 211 n.

There is no ventral intersegmental sclerite. The

genital segment is 195 x 225 n, a little wider than

long. The four postgenital segments are 99 x 116 n,

104 x 104 n, 78 x 83 n, and 62 x 74 n from anterior to posterior.

The caudal ramus resembles that of the female,

but there is no spur near the lateral seta, the two

dorsal surficial refractile areas are absent, and the

dimensions are smaller, 89 x 29 n.

The rostrum is similar to that of the female.

The first antenna (Figure 175g) is like that of the

female, but there are three additional aesthetes,

so that the formula is 4, 13 -j- 2 aesthetes, 6, 3 -j- 1

aesthete, 4 -(- 1 aesthete, 2 -f 1 aesthete, and

7 -f 1 aesthete.

The second antenna, labrum, mandible, paragnath, first maxilla, and second maxilla are similar

to those in the female.

The maxilliped (Figure 175A) is 4-segmented

(assuming that the proximal part of the claw

represents a fourth segment). The first segment

is unarmed. The second segment bears two equal,

naked inner setae and is ornamented with small

spines on its proximal inner corner. The small

third segment is unarmed. The claw is relatively

short, % n along its axis, with slight indication of

division midway, bears proximally two unequal

naked setae, and has a row of minute denticulations

along its concave side. The terminal lamella is

very narrow.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are similar to those of the female, with

the same armature. In seven males there was no

inner coxal seta. Only one of these males showed

abnormal armature (the last segment of the right

leg 2 endopod having 1,11,2 instead of the usual

condition of 1,11,3).

Leg 5 resembles that of the female, but the free

segment is smaller, 19 x 8 n.

Leg 6 (Figure 175*) is a posteroventral flap on

the genital segment bearing two naked setae, 21 n

and 22 |i.

The spermatophore was not observed.

The color in life is unknown.

ETYMOLOGY.—The specific name varius (Latin,

diversified or varying) alludes to the variability of

the armature on legs 1-4.

Genus Zamolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome 5—segmented in the female, 6-segmented in the male.
NUMBER 127 S15

Caudal minus with six setae. Rostrum broadly

rounded posteroventrally. First antenna 7-segmented, with the formula in the female 4, 13, 6, 3,

4 -f 1 aesthete, 2 -f 1 aesthete, 2 + 1 aesthete,

and 7 -f- 1 aesthete; in the male 4, 13 -|- 2 aesthetes,

6, 3 + 1 aesthete, 4 -f- 1 aesthete, 2 -f 1 aesthete,

and 7 -f- ' aesthete. Second antenna 4-segmented,

with the armature 1, 1, 3, and 1 + 5 small elements, there being a single terminal claw.

Labrum with a deep cleft separating the two

lobes. Mandible with the basal part having on its

convex side a roughened area followed by a slender,

proximally directed digitiform process and then by

a serrated fringe, and on its concave side beyond

the indentation a row of spinules; lash long.

Paragnath a small hairy lobe. First maxilla with

four elements. Second maxilla of the usual lichomolgid type. Maxilliped in the female 3-segmented,

with a pointed tip; in the male 4-segmented (assuming that the proximal part of the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami, except for leg

4 endopod which is 2-segmented. Armature of the

usual lichomolgidiform type, with leg 4 endopod

having the formula 0-1 ;II, the seta on the first

segment being either feathered or naked. Leg 4

exopod with the third segment having 111,1,5.

In the male leg 1 endopod with the third segment

1,1,4 instead of 1,5 as in the female. Leg 5 in both

sexes with a free segment bearing two terminal

setae. Leg 6 represented in the female by the two

seta and process near the area of attachment of

each egg sac and in the male by a posteroventral

flap on the genital segment bearing two setae.

Other features as in the species described below.

Associated with alcyonaceans.

TYPE-SPECIES.—Zamolgus tridens, new species.

ETYMOLOGY.—The name is a combination of the

Greek words £a (very) and

Key to Species of the Genus Zamolgus

FEMALES

Free segment of leg 5 with numerous spines; first maxilla with three elements; second maxilla

with lash and adjacent seta nearly equal in length; tip of maxilliped with a single, pointed

process and a spine; leg 4 endopod with seta on first segment plumose Z. acanthodes

Free segment of leg 5 smooth, not ornamented with spines; first maxilla with four elements;

second maxilla with lash and adjacent seta very unequal in length; tip of maxilliped with

two pointed processes and a spine; leg 4 endopod with seta on first segment naked ... Z. tridens

MALES

Free segment of leg 5 with a few small spines; second segment of maxilliped with one row of

spines Z. acanthodes

Free segment of leg 5 without small spines; second segment of maxilliped with two rows of

spines Z. trident

Zamolgus tridens, new species

FIGURES 176-178

TYPE MATERIAL.—63 $ $, 85 o" d". and 82 copepodids from single colony of the alcyonacean

Cespitularia turgida Verseveldt, in 20 m, west of

Andilana, Nosy B£, northwestern Madagascar,

13° 18' S, 48°07' E, 24 August 1967, collected by

AGH. Holotype $, allotype, and 120 paratypes

(50 $ c., 70 tf tf) deposited in USNM and the

remaining paratypes in the collection of AGH.

FEMALE.—The body (Figure 176a) has a moderately slender prosome. The length (not including

the setae on the caudal rami) is 1.01 mm (0.92-1.10

mm) and the greatest width 0.47 mm (0.43-0.52

mm), based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome

is 1.48:1. The ratio of the length of the prosome

to that of the urosome is 1.94:1. The segment of

leg 1 is clearly set off from the cephalosome by a

dorsal transverse furrow. The epimeral areas of the

pedigerous segments are rounded.

The segment of leg 5 (Figure 1766) is 78 x 156 n.

There is no definite ventral intersegmental sderite

between that segment and the genital segment.

The genital segment is 153 x 133 n, only slightly
316

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 176.—la mo Igus tridens, new species. Female: a, dorsal (A) ; b, urosomc, dorsal (B) ; c,

area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (c); e, rostrum, ventral (B) ;

/, first antenna, dorsal (D) ; g, second antenna, posterior (E) ; h, labrum, with paragnaths indicated by broken lines, ventral (E) . Scale: A, B, 02 mm; c, 0.02 mm; D, 0.1 mm; E, 0.05 mm.
NUMBER 127 317

expanded. The areas of attachment of the egg sacs

are situated dorsolaterally in front of the middle

of the segment. Each area (Figure 176c) bears two

small, naked setae 6 n long, with a small process

between them. The three postgenital segments are

47 x 88 n, 31 x 80 n, and 39 x 73 n from anterior

to posterior.

The caudal ramus (Figure I76d) is only a little

wider than long, 32 x 27.5 n. The dorsal seta is 35 n,

the outer lateral seta 57 n, the outermost terminal

seta 64 n, and the innermost terminal seta 125 n.

The two long median terminal setae are 198 n

(outer) and 286 n (inner), both inserted between

dorsal (unornamented) and ventral (with a marginal row of spinules) flaps. All the setae are naked.

The body surface is ornamented with hairs (sensilla) as shown in Figure 176a,6. The posteroventral

margin of the anal segment is smooth.

The egg sacs were incomplete in all ovigerous

females collected. Each egg is about 73 \t in

diameter.

The rostrum (Figure 176*) is broadly rounded

posteroventrally.

The first antenna (Figure 176/) is 296 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 36 (56

n along the anterior margin), 89, 24, 41, 40, 26, and

20 \i respectively. The formula for the armature

is 4, 13, 6, 3, 4 -j- 1 aesthete, 2 -+- 1 aesthete, and

7 -f 1 aesthete. All the setae are naked.

The second antenna (Figure 176g) is 4-segmented. Both the first and second segments bear

an inner seta. The third segment bears three setae.

The fourth segment, 51 M along its outer edge, 36 n

along its inner edge, and 20 n wide at the middle,

bears a single terminal claw 43 n along its axis and

five small hyaline elements. All the setae are naked.

The labrum (Figure 176/i) has two, peculiarly

pointed, posteroventral lobes.

The mandible (Figure 177a) has on the concave

side of the base an indentation distal to which

there is an uninterrupted row of spinules; the convex margin of the base has proximally a roughened

area, more distally a slender, proximally directed

digitiform process, and finally a serrated, hyaline

fringe. The lash is moderately long and bilaterally

barbed. The paragnath (Figure 176h) is a small

hairy lobe. The first maxilla (Figure 1776) has four

elements, one of them finely barbed along one edge.

The second maxilla (Figure 177c) is 2-segmented.

The first segment is unornamented. The second

segment is relatively large, with proximally on its

outer (ventral) margin a small setule, a surficial

anterior seta with short spinules along one edge,

and distally on its inner (dorsal) margin a barbed

seta. The terminal lash has a line of articulation

with the segment. Along the outer edge of the lash

are four or five large teeth followed by small spinules; the inner edge bears only a few minute distal

spinules. The maxilliped (Figure 177d) is 3-segmented. The first segment is unornamented. The

second segment bears two, very unequal, naked

setae. The third segment bears a small naked setule,

a naked spine, and two terminal spiniform processes without articulations, the larger process with

a row of denticles on both anterior and posterior

surfaces.

The ventral area between the maxillipeds and

the first pair of legs (Figure 177e) is only slightly

protuberant. There is a sclerotized line (incomplete medially) between the bases of the maxillipeds.

Legs 1-4 (Figure 177/-J) have the same segmentation and spine and setal formula as in Meringomolgus facetus. The inner seta on the coxa of legs

1-3 is large and feathered, but in leg 4 this seta

is short (11 M-) and naked. The inner margin of the

basis of legs 1-3 has a row of hairs but in leg 4 it

is naked. The exopod of leg 4 is 140 n long, with

the last segment having the formula 111,1,5. The

endopod of leg 4 has hairs along the outer margins

of both segments. The first segment is 33 x 28 n (the

length not including the spinous processes) and

bears an inner distal naked seta, 21 ji. The second

segment is 77 n long (including the terminal

spiniform processes), 28 \i in greatest width proximally, and 16.5 n in least width distally; the outer

terminal spine is 22 n, the inner spine 53 n, both

fringed.

Leg 5 (Figure 177/) has an unornamented free

segment 70 n long, 20 n wide at the slight proximal

inner expansion, and 16.5 n wide distally. The two

naked terminal setae are 33 n and 27 n. The naked

seta on the body near the free segment is 32 n.

Leg 6 is probably represented by the two small

setae near the attachment of each egg sac (Figure

176c).

The color in life in transmitted light is opaque,

the eye red, the egg sacs gray.
SMITHSONIAN CONTRIBUTIONS TO ZOOl^r.v

a

FIGURE 177.—Zamolgus tridens, new species. Female: a, mandible, posterior (E) ; b, first

posterior (F) ; c, second maxilla, anterior (E) ; d, maxilliped, anterior (c); e, area between

maxillipeds and first pair of legs, ventral (B) ; /, leg and intercoxal plate, anterior (D) ; g, leg 2,

anterior (D) ; h, third segment of endopod of leg 3, anterior (D) ; i, leg 4, anterior (D) ; ;, leg 5,

dorsal (E) . Scale: B, 02 mm; D, 0.1 mm; E, F, 0.05 mm.
NUMBER 127 319

MALE.—The body (Figure 178a) is only slightly

more slender than the female. The length (not

including the ramal setae) is 0.85 mm (0.74-0.90

mm) and the greatest width 0.S1 mm (0.29-0.31

mm), based on ten specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.67:1. The ratio of the length of the prosome

to that of the urosome is 1.47:1.

The segment of leg 5 (Figure 1786) is 39 x 97 n.

There is no ventral intersegmental sclerite. The

genital segment is 208 x 200 n. The four postgenital

segments are 27 x 53 n, 27 x 53 n, 21 x 48 n, and 23

x 49 n from anterior to posterior.

The caudal ramus is like that of the female,

but smaller, 24 x 19 n.

The body surface is ornamented with hairs as

in the female.

The rostrum is like that of the female.

The first antenna resembles that of the female,

but additionally it has two long aesthetes on the

second segment and one long aesthete on the fourth

segment, so that the formula is 4, 13 -|- 2 aesthetes,

FIGURE 178.—Zamolgus tridens, new species. Male: a, dorsal (A) ; b, urosome, dorsal (B) ; c,

second antenna, posterior (F) ; d, maxilliped, inner (E) ; e, third segment of endopod of leg 1,

anterior (F) ; /, third segment of endopod of leg 2, anterior (F) ; g, spermatophores, attached to

female, dorsal (B) . Scale: A, B, 02 mm; D, 0.1 mm; E, F, 0.05 mm.
320 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

6, 3 + 1 aesthete, 4 + 1 aesthete, 2 -f 1 aesthete,

and 7 -\- 1 aesthete. All the setae are naked.

The second antenna (Figure 178c) is similar to

that of the female, but there are short stout spines

on the inner margins of the first two segments.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female.

The maxilliped (Figure 178d) is 4-segmented,

assuming that the proximal half of the claw represents a fourth segment. The first segment is unarmed. The second segment bears two naked setae

and two rows of spinules. The small third segment

is unarmed. The claw is 152 n along its axis, with

a line of division about midway, where on the

concave edge there is a row of spinuliform serrations. Proximally the claw bears two unequal

setae, the longer one with fine distal barbules.

The ventral area between the maxillipeds and

the first pair of legs is like that of the female.

Legs 1-4 are segmented and armed as in the

female, except for the last segment of the endopod

of leg 1 (Figure 178e) where the formula is 1,1,4

instead of 1,5 as in the female. Slight sexual dimorphism is seen also in the smaller outer distal

spinous processes on the first and second segments

of the endopods of legs 1 and 2, and in the details

of the armature of the last segment of the endopod

of leg 2 (Figure 178/), where the barbs on the

middle spine are much more prominent than in

the female. Legs 3 and 4 are like those of the

female.

Leg 5 (Figure 1786) has a slender, subrectangular, unornamented free segment, 38 x 10 ji. The two

naked terminal setae are 22 n and 30 n.

Leg 6 (Figure 1786) consists of a posteroventral

flap on the genital segment bearing two naked

setae, 22 n and 29 n.

The spermatophore (Figure 178g), attached to

the female in pairs, is about 192 x 86 n, not including the neck. Its wall is unusually thick and

golden brown in color.

The color in life in transmitted light is like that

of the female.

ETYMOLOGY.—The specific name tridens (Latin,

having three teeth or prongs) alludes to the tripartite extremity of the maxilliped in the female.

REMARKS.—The genus Zamolgus resembles Odontomolgus in having a proximally directed, toothlike

process on the mandible. It differs from that genus,

however, in the armature of the last segment of the

exopod of leg 4 being 111,1,5 and the row of spinules

on the concave side of the base of the mandible

being uninterrupted instead of being arranged on

two lobes.

Zamolgus acanthodes, new species

FIGURES 179-181

TYPE MATERIAL.—9 9 9 > 6 tf tf from single

colony of the alcyonacean Sinularia arborea Verseveldt, in 12 m, west of the harbor at Hellville,

Nosy Be\ northwestern Madagascar, 4 August 1967,

collected by AGH. Holotype $ , allotype, and 9

paratypes (6 9 9 , 3 cTd") deposited in USNM, the

remaining paratypes (dissected) in the collection

of AGH.

OTHER MATERIAL EXAMINED (all from Sinularia

arborea).—3 cf o* from single colony, in 2 m, Pte.

Lokobe, Nosy Be, 3 June 1967; 9 9 9 . 8 rf rf from

single colony, in 23 m, Tany Kely, a small island

south of Nosy Be, 30 June 1967; 1 9 , 2 tf d from

single colony, in 2 m, off Ampombilava, Nosy Bt'.

7 July 1967: and 5 ?? , 1 rf from ten small

colonies, in 13 m, opposite Antsiabe, southern shore

of Nosy Komba, near Nosy hi, 2 September 1967.

All collected by AGH.

FEMALE.—The body (Figure 179a) has a moderately broadened prosome. The length (not including the setae on the caudal rami) is 1.02 mm (0.95-

1.09 mm) and the greatest width 0.50 mm (0.47-0.53

mm), based on nine specimens in lactic acid. The

ratio of the length to the width of the prosome is

1.31:1. The ratio of the length of the prosome to

that of the urosome is 1.93:1. The segment of leg

1 is separated from the head by a dorsal transverse

furrow. The epimera of the segment of leg 3 are

angular as indicated in the figure.

The segment of leg 5 (Figure 1796) is 78 x 180 i*.

There is no ventral intersegmental sclerite between

that segment and the genital segment. The genital

segment is 133 x 159 n in greatest dimensions in

dorsal view, its anterior two-thirds expanded with

rounded lateral margins, its posterior third with

parallel margins. The areas of attachment of the

egg sacs are located dorsolaterally in the middle

of the expanded portion. Each area (Figure 179c)

bears two small, naked setae, 6 n and 11 n, with a

minute spiniform process between them. The three

postgenital segments are 66 x 88 \i, 50 x 76 n, and

44 x 69 n from anterior to posterior.
NUMBER 127 321

FIGURE 179.—Zamolgus acanthodes, new species. Female: a, dorsal (A) ; b, urosome, dorsal (B) ; c,

area of attachment of egg sac, dorsal (c); d, caudal ramus, dorsal (D) ; e, rostrum, ventral (B) ;

/, first antenna, ventral (E) ; g, second antenna, anterior (F) ; h, labrum, with paragnaths indicated by broken lines, ventral (c). Scale: A, 0.2 mm; B, E, F, 0.1 mm; c, 0.05 mm; D, 0.02 mm.
322 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The caudal ramus (Figure 179d) is slightly

wider than long, 24 x 27.5 n. The dorsal seta is 35 n,

the outer lateral seta 72 n, the outermost terminal

seta 83 n, and the innermost terminal seta 140 n.

The two long terminal setae are 275 n (outer) and

350 n (inner), both inserted between dorsal (unornamented) and ventral (with a marginal row

of spinules) flaps. All the setae are naked.

The body surface is ornamented with hairs (sensilla) as shown in Figure 179aA The posteroventral

margin of the anal segment is smooth.

The egg sacs were broken in all ovigerous females seen. The eggs are numerous, each about 46 n

in diameter.

The rostrum (Figure 179c) has a broadly rounded

posteroventral margin. Between this margin and the

anterior edge of the labrum there are pairs of

sclerotized lines.

The first antenna (Figure 179/) is 314 n long.

The lengths of the seven segments (measured along

their posterior nonsetiferous margins) are 27 (66 »*

along the anterior margin), 94, 29, 58, 32, 18, and

17 n respectively. The formula for the armature is

like that of Z. tridens. All the setae are naked.

The second antenna (Figure 179g) has a seta on

the first and second segments and three setae on the

third segment. The fourth segment, 66 n along its

outer edge, 44 H along its inner edge, and 21 n wide,

bears a single terminal claw 51 n along its axis and

six very small, hyaline elements. All the setae are

naked.

The labrum (Figure I79h) has two broad, posteroventral lobes of unusual form.

The mandible (Figure 180a) resembles that of

Z. tridens but the toothlike process on the convex

margin of the base is stouter and more pointed,

and just proximal to its origin there is a row of

spinules (not present in Z. tridens). The paragnath

(Figure 179ft) is a small hairy lobe. The first maxilla

(Figure 1806) has three elements. The second

maxilla (Figure 180c) has on its second segment

a small setule proximally on its outer (ventral)

margin, a surficial anterior seta finely barbed along

one edge, and a long, strongly barbed seta distally

on its inner (dorsal) margin. The terminal lash has

on one side a crest of about nine long falciform

spines followed by a series of short spines; the

opposite side has a row of spinules. The maxilliped

(Figure 180d) has on the first segment a few small

inner spines and on the second segment two very

unequal setae (the longer one barbed) and a

few small spines. The last segment bears a small,

naked setule, a spine with a few minute barbs, and

a terminal barbed spiniform process without an

articulation.

The ventral area between the maxillipeds and the

first pair of legs (Figure \S0e) is not protuberant.

A sclerotized line extends between the bases of the

maxillipeds.

Legs 1-4 (Figure 180/—i) are segmented and

armed as in Z. tridens. On the outer side of the

coxa of leg 1 there is a posterior lobe (Figure 180/).

The inner seta on the coxa of legs 1-3 is large and

feathered, but in leg 4 this seta is short (20 n) and

naked. The inner margin of the basis is haired in

legs 1-3 but naked in leg 4. The three spines on the

last segment of the endopod of leg 2 are 26, 26, and

29 i> from outer to inner. The exopod of leg 4 is

134 n long, with the last segment having the formula 111,1,5. The endopod has hairs along the outer

margins of both segments and a few minute barbs

on the inner margin of the second segment. The

first segment is 35 x 22 ^ (not including the spinous

processes) and bears an inner distal feathered seta

56 n. The second segment is 88 n long (including

the terminal spiniform processes), 17.5 n in greatest

width, and 15 n in least width; the terminal outer

spine is 26 n, the inner spine 55 i*, both fringed.

Leg 5 (Figure 180;*) has a long, slightly recurved

free segment reaching posteriorly beyond the genital

segment. Its greatest dimensions are 187 x 42 p. The

two short terminal naked setae are 28 i* and 33 n.

The segment is ornamented dorsally and outwardly

with moderately long spines. The naked seta on the

body adjacent to the free segment is about 28 n;

near this seta is a group of slender spinules.

Leg 6 is probably represented by the two small

setae near the attachment of each egg sac (Figure

179c).

The color in life in transmitted light is translucid, the prosome with a few small red globules,

the eye red, the egg sacs opaque.

MALE.—The body (Figure 181a) has a somewhat broadened prosome. The length (not including the ramal setae) is 0.76 mm (0.74-0.78 mm)

and the greatest width 0.31 mm (0.30-0.32 mm),

based on six specimens in lactic acid. The ratio

of the length to the width of the prosome is 1.42:1.

The ratio of the length of the prosome to that of

the urosome is 1.56:1.
NUMBER 127

FIGURE 180.—Zamolgus acanthodes, new species. Female: a, mandible, posterior (c); b, first

maxilla, posterior (c); c, second maxilla, posterior (c); d, maxilliped, anterior (c); e,

area between maxillipeds and first pair of legs, ventral (E) ; /, leg 1 and intercoxal plate, anterior (E) ; g, leg 2, anterior (E) ; h, third segment of endopod of leg S, anterior (E) ; i, leg 4,

anterior (E) ; ;, leg 5, dorsal (E) . Scale: c, 0.05 mm; E, 0.1 mm.
324 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The segment of leg 5 (Figure 1816) is 52 x 83 ».

There is no ventral intersegmental sclerite. The

genital segment is 159 x 164 n in dorsal view, its

lateral margins only slightly expanded. The four

postgenital segments are 28 x 54 it, 28 x 53 i»,

21 x 51 n, and 25 x 50 j* from anterior to posterior.

The caudal ramus resembles that of the female

but is smaller, 19 x 21 n.

The body surface has a few hairs as in the female.

The rostrum is similar to that of the female. The

sclerotized lines seen in the female between the

rostrum and the labrum are absent.

The first antenna is like that of the female but

there are three additional aesthetes, so that the

formula is the same as in the male of Z. tridens.

All the setae are naked.

The second antenna (Figure 18\c) is similar in

general form and armature to that of the female

but has added ornamentation. The first segment

has a small spine near the seta. The second segment

has a long inner row of spines. The fourth segment,

55 H along its outer edge, 37 v- along its inner edge,

FIGURE 181 .—Zamolgus acanthodes, new species. Male: a, dorsal (A) ; b, urosome, dorsal (B) ; c,

second antenna, posterior (c); d, maxilliped, outer (E) ; e, endopod of leg 1, anterior (c); /,

third segment of endopod of leg 2, anterior (c); g, leg 5, dorsal (D) ; h, spermatophores, attached to female, dorsal (B) . Scale: A, 02 mm; B, E, 0.1 mm; c, 0.05 mm; D, 0.02 mm.
NUMBER 127 325

and 14 n wide, with its claw 45 n along its axis, has

a row of very small spines on its distal inner surface.

The labrum, mandible, paragnath, first maxilla,

and second maxilla are like those of the female. The

maxilliped (Figure 181d) is more elongated and

slender than in Z. tridens. The second segment

bears a seta unilaterally fringed with spinules, a

naked seta, and a single row of long spines. The

claw is 216 u along its axis including the terminal

lamella and has the usual two unequal proximal

setae.

The ventral area between the maxillipeds and

the first pair of legs resembles that of the female.

Legs 1-4 are segmented and armed as in the

female except for the last segment of the endopod

of leg 1 (Figure 181«), where the formula is 1,1,4,

the two spines being 31 n and 20 u. Sexual dimorphism is present also in the last segment of the

endopod of leg 2 (Figure 181/) where the three

spines from outer to inner are 10, 9, and 11 n, all

shorter than in the female (see Figure 180g). Legs

3 and 4 are like those of the female. The coxa of

leg 1 has a postero-outer lobe as in the female.

Leg 5 (Figure 181g) has a small, elongated free

segment 37 x 10 u. The terminal setae are 23 v- and

18 n. The segment is ornamented with a few small

spines. The seta adjacent to the free segment is

22 u.

Leg 6 consists of a posteroventral flap on the

genital segment bearing two naked setae, 11 n

and 13 u (Figure 1816).

The spermatophore (Figure 18 Ig), attached to

the female in pairs, is ovoid, 117 x 70 n, not including the neck.

ETYMOLOGY. — The specific name acanthodes,

Greek axav*a>8iis (thorny), allules to the spines on

leg 5 in the female.

REMARKS.—This species may be easily distinguished from Z. tridens in the female by the ornamentation of spines on leg 5, the shape of the

genital segment, and the bidentate appearance

(rather than tridentate) of the last segment of the

maxilliped. In the male, this species has a different

ornamentation on the second antenna, only one

row of spines is present on the second segment of

the maxilliped, and the outer of the two spines on

the last segment of the endopod of leg 1 is longer

and stronger than the inner. Both sexes of Z acanthodes may be separated from Z. tridens by details

of the labrum and the mouthparts.

Genus Zygomolgus Humes and Stock, 1972

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented. Caudal ramus with six setae.

Rostrum rounded so far as known. First antenna

7-segmented. Second antenna 4-segmented, the

third segment with a clawlike element and the

fourth segment with four claws.

Mandible with a slender base merging into a

long, slender pectinate lash. First maxilla with twofour elements. Second maxilla of the usual lichomolgid form. Maxilliped in the female 3-segmented.

Legs 1-4 with 3-segmented rami except for leg

4 endopod which is 2-segmented. Armature generally of the typical lichomolgidiform type. Leg 4

exopod with the third segment having 11,1,5. Leg 4

endopod with the formula 0-1; II, the seta being

feathered.

Leg 5 with a small, unornamented free segment

bearing two terminal elements.

Other features as in the species below.

Associated with ascidians or algae (a single female

of Z. tenuifurcatus with a holothurian).

Key to Species of the Genus Zygomolgus

FEMALES

1. Caudal ramus more than twice as long as anal segment Z. tenuifurcatus

Caudal ramus about as long as or scarcely longer than anal segment 2

2. Prosome broad; leg 4 endopod with second segment not much longer than first segment

Z. didemni

Prosome slender; leg 4 endopod with second segment at least li/2 times longer than first

segment 3

3. Leg S exopod with third segment having 11,13 Z. curtiramus

Leg S exopod with third segment having IILL5 Z. poucheti
326 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCY

TYPE-SPECIES.—Zygomolgus tenuifurcatus G. O.

Sars.

ETYMOLOGY.—The generic name is a combination

of twyou (a yoke) and HOXYOC,.

REMARKS.—The male is unknown in all species.

Zygomolgus tenuifurcatus (G. O. San, 1917)

FIGURE 182

I.ichomolgus tenuifurcatus G. O. San, 1917a, pp. 160, 161,

pi. 89 [from Norway].-Gotto, 1954a, p. 133 [from the

holothurian Lapidoplax digitate (Montagu), Strangford

Lough, County Down, Northern Ireland]; 1955, p. 391 [from

the ascidian Diplosoma listerianum, Strangford Lough,

County Down, Northern Ireland]; 1960b, pp. 213. 223, fig.

30; 1966, p. 194.

Zygomolgus curtiramus (Boquet and Stock, 1962)

I.ichomolgus curtiramus Bocquet and Stock, 1962b, pp. 244-

249, figs. 1, 2 [from the alga Lithophyllum incrustans

Saint Efflam, Cdtes-du-Nord. France].

Zygomolgus didemni (Gotto, 1956)

Lichomolgus didemni Gotto, 1956, pp. 600-602, figs. 1-9

[from the ascidian Didemnum maculosum (Milne Edwards),

Northern Ireland]: 1960b, pp. 213, 223. fig. 1; 1966, p.

I'M. • Hocquet and Stock, 1962b, pp. 248. 249.-Glacon.

1971, p. 23 [unnumbered].

Zygomolgus poucheti (Canu, 1891)

Lichomolgus poucheti Canu, 1891a, pp. 478, 479 [from the

surface of colonies of the asddians Morchellium argus M.

Edwards and Fragarium areolatum Giard, Concarneau,

France]; Canu, 1892, pp. 231, 232, pi. 23. fig*. 5-12 [from

Morchellium argus and Fragarium areolatum, Conrarneau,

France].-T. Scott. 1907. p. 371 - G. O. Sara, 1917a. pp.

158-160, pi. 88 [from Norway] .-Cotto, 1960b. p. 227 [on

Morchellium argus and Sidnyum elegans].—Bocquet and

Stock. 1962b, pp. 244. 247-249. fig. 3 [free living. CAin-duNord, France].

FIGURE 182.—Zygomolgus tenuifurcatus (G. O. Sars). Female: a, dorsal; b, second antenna; c;

mandible and first maxilla; d, leg 4. [Redrawn from G. O. Sars, 1917a, pi. 89, figs. 9 (omitting

the first antennae and egg sacs), a* m, p4.] Length of female 1.40 mm.
NUMBER 127 327

Lichomolgidae Insufficiently Described or of

Uncertain Position

Boholia cerianthiphila Kossman, 1877

Boholia cerianthiphila Kossmann, 1877, pp. 22, 23, pi. 4,

fig. 4, pi. 5, fig. 2 [from the ceriantharian Cerianthus,

canal of Lapinig, Bohol, Philippine Islands].

Boholia cerianthiphyla.-De Zulueta, 1912, pp. 11-12.

Kelleria gurneyi Sewell, 1949

Kelleria gurneyi Sewell. 1949, pp. 117-119, fig. 31 A-H [in surface tow-net, Kuiau River, Perak, Federated Malay States].

REMARKS.—The male is unknown.

Kelleria sp.

Kelleria sp., Grice, 1960, p. 220 [in plankton, northwestern

coast of Florida].

Lichomolgus aegyptius Gurney, 1927

Lichomolgus aegyptius Gurney. 1927, pp. 467, 468, fig. 112

[from Suez Canal].

REMARKS.— The female is unknown.

Lichomolgus elegans Thompson and A. Scott, 1903

Lichomolgus elegans Thompson and A. Scott, 1903, p. 280,

pi. 16, figs. 8-13 [from washings of dredged invertebrates,

Ceylon].-A. Scott, 1909, p. 264. [Siboga Station 226, midway between Lucipara and Schildpad Islands].

Lichomolgus (Macrochiron) elegans.—Monod and Dollfus,

1932. p. 139.

Macrochiron elegans.—Stock, 1957, p. 381.

rstellicola elegans.-Stock, 1957, p. 381.

REMARKS.—The male is unknown.

Lichomolgus fuliginosus Brady, 1910

Lichomolgus fuliginosus Brady, 1910, p. 573, fig. 59 [at 385

m, Gauss Station, Antarctica] .-Stock, 1960a, pp. 370, 371,

fig. 4.

REMARKS.—Only one specimen is known—a female according to Brady, but sex uncertain according to Stock (1960a).

Lichomolgus gigas Thompson and A. Scott, 1903

Lichomolgus gigas Thompson and A. Scott, 1903, pp. 280,

281, pi. 16, figs. 21-26 [from washings of dredged invertebrates, Ceylon].—A. Scott, 1909, p. 264 [between Lucipara

and Schildpad Islands, 5° 26.7' S, 127*36.5' E. Siboga Station

226].—Ummerkutty, 1968, p. 315 [in starfish washings, Gulf

of Mannar, southeastern India].

Lichomolgus rotundus Sewell, 1949

Lichomolgus rotundus Sewell, 1949, pp. 97-99, fig. 23 A-* [in

weed-washings, Addu Atoll, Maldive Archipelago; spelling

is rotundus on pp. 19, 97 but rotundatus on pp. 92, 93.

REMARKS.—The male is unknown.

Lichomolgus (Lichomolgus) sp.

Lichomolgus (Lichomolgus) sp., Monod and Dollfus, 1932a,

p. 138, figs. 2c, 3A, 4A, 5D, 6B,C,C,H [from the nudibranch

Aeolidia nebae Risbec, New Caledonia],

"Copopode commensal."—Risbec, 1930, p. 296, fig. 60 [from

Aeolidia nebae Risbec. New Caledonia].

Lichomolgus (Macrochiron) sp.

Lichomolgus (Macrochiron) sp.—Monod and Dollfus, 1932a,

p. 140, figs. 2E, 5f.E, 6A, 7 [from Chrotnodoris mouaci

Risbec, He Mouae, New Caledonia].

"Cope'pode commensal."—Risbec, 1930, p. 281 [from the

nudibranch Chromodoris mouaci Risbec, New Caledonia].

Lichomolgus sp.

Lichomolgus sp.—Marques, 1949, pp. 24, 26, pi. 3, figs, b, c, f

[from Portuguese Guinea].—Monniot, 1961, p. 97, fig. Id

[from the ascidian Microcosmus sabatieri Roule, Banyuls,

Mediterranean coast of France].—Ummerkutty, 1968, pp.

315-317. figs. 22-31 [from weed-washings (sic) of the starfish Pentaceros hedemanni (Lutken), Gulf of Mannar,

southeastern India].

Lichomolgus tenuicornis Brady, 1910

Lichomolgus tenuicornis Brady, 1910, pp. 571, 572, fig. 58

[at 385 m, Gauss Station, Antarctica].

REMARKS.—The male is unknown.

Lichomolgus vagans Gurney, 1927

Lichomolgus vagans Gurney, 1927, pp. 465, 466, fig. HI [from

Suez Canal].

REMARKS.—The female is unknown.

Macrochiron (Macrochiron) longipes Sewell, 1949

Macrochiron (Macrochiron) longipes Sewell, 1949, pp. 104.

105, fig. 26A-I [in weed-washings, Addu Atoll, Maldive

Archipelago].

Lichomolgus longipes.-Stock, 1957, p. 382.
328

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—This is a Lichomolgus according to

Stock (1957, p. 382). The nature of the mandible

is not known. The male is unknown.

Paralichomolgus longicaudatus.—Monod and Dotlfus, 1932a,

p. 139.

Linckiomolgus caeruleus.—Monod and Dollfus, 1932a, p. 142.

Macrocheiron sp.

Macrocheiron sp., Grice, 1960, p. 220 [in plankton, Alligator

Harbor, west coast of Florida] .-Wickstead, 1961, p. 61

[Indo-West-Pacific].—Bruce, Colman, and Jones, 1963, p.

128 [from Isle of Man, England].

Oncaeola specialis Kramer, 1895

Oncaeola specialis Kramer, 1895, p. 220, pi. 18 (figs. 2, 4, 5, 7),

pi. 19 (figs. 1, 3, 7a, 8, 9) [in plankton, New Zealand].

Paralichomolgus orbicular is Monod, 1928

Paralichomolgus orbicularis Monod, 1928, pp. 2-8, fig. 2 [from

the gills of the nudibranch Platydoris cruenta (Quoy and

Gaimard), Noumea, New Caledonia].—Risbec, 1928, p. 35,

fig. 1 ter [from Platydoris cruenta (Quoy and Gaimard),

New Caledonia].

Lichomolgus (Stellicola) orbicularis.—Monod and Dollfus,

1932a, p . 139; p. 143, figs. 2A,B, 5A,B, 6F,F.

Pseudanthesshu minimus Tanaka, 1960

Pseudanthessius minimus Tanaka, 1960, pp. 75-77, pi. 33,

figs. 1-18 [in plankton, Indian Ocean, 08° 15' S, 76s 15' £].

REMARKS.—This species is perhaps near Parameter ochir on.

Stellicola alabatensis Kossmann, 1877

Stellicola alabatensis Kossmann, 1877, pp. 14, 15, pi. 1, figs.

2, 3 [from Philippine Islands].

"Cop£pode commensal."—Risbec, 1930, p. 290 [from Trevelyana kouaouae Risbec, New Caledonia].

Lichomolgus (Macrochiron) alabatensis.—Monod and Dollfus,

1932a, p. 140, figs. 2D, 4B, 6D, 9A-B [from the nudibranch

Gymnodoris (=Trevelyana) kouaouae (Risbec), Noumea,

New Caledonia].

Lichomolgus alabatensis.—Stock, 1957, p. 381.

Macrochiron alabatensis.—Stock, 1957, p. 381.

Stellicola semperi Kossmann, 1877

Stellicola semperi Kossmann, 1877, pp. 13, 14, pi. 3, fig.

1 [from the asteroid Ophidiaster miliaris, Isabela, Philippine Islands].

Lichomolgus (Stellicola) semperi.-Monod and Dollfus, 1932a,

p. 139.

Species Undesignated

"Copepodes," Risbec, 1928, p. 34 [on gills of the nudibranchs

Platydoris immonda Risbec and Trevelyana (=Gymnodoris)

ceylonica (Kelaart)] New Caledonia], pp. 34, 169 [on gills

of the nudibranch Noumea flava (Eliot), New Caledonia].

"Lichomolgide (a decrire)," Pruvot-Fol. 1954, p. 25 [from

Armina maculatd].

Undetermined Lichomolgidae.—Semper, 1862, pp. 105, 106

[from naked mollusks, Philippine Islands].—Boas, 1886, p.

34 [from Cavolina (=Hyalaea) tridentata (Fortkil), locality unknown; "en Ergasilus-lignende Snyltekreb*"].—

Pelseneer, 1903, p. 7 [from mantle cavity of the bivalve

Nacella mytilina (Helbling), Tierra del Fuego].

Copepods Placed by Various Authors in the Old

Family Lichomolgidae but Belonging Elsewhere

Anthessius Delia Valle, 1880 a,b. Myicolidae. Listed

in the Lichomolgidae by Monod and Dollfus,

1932a, pp. 148-151.

Boeckia Brady, 1872 a.b, pp 430-431. Preoccupied.

(=Anthessius.)

Cholidya polypi Farran, 1914. Harpacticoida. Listed

in the Lichomolgidae by Monod and Dollfus,

1932a, p. 153. (See discussion in Bresciani, 1970,

p. 15.)

Hermannella arenicola (Brady, 1872).—Farran,

1913, p. 6.—Norman and T. Scott, 1906, p. 199.

(=Anthessius.)

Herrmannella concinna A. Scott, 1909, pp. 262-

263, pi. 68, figs. 1-10. (=Anthessius.)

Lichomolgus aberdonensis T. Scott and A. Scott,

1892b, pp. 149-151, pi. 6 figs. 1-12. (=Hcmicyclops.)

Lichomolgus adhaerens Williams, 1907, pp. 75, 76,

pi. 2. (=Hemicyclops.)

Lichomolgus arenicolus (Brady, 1880), vol. 3, pp.

46, 47, pi. 87, figs. 1-7 (=Anthessius.)

Lichomolgus brexneaudis Leigh-Sharpe, 1934, pp.

11-13, fig. 7. (=Anthessius. See Stock, Humes

and Gooding, 1963b, p. 2; Stock, 1964c, pp. 121,

123.)

Lichomolgus littoralis T. Scott, 1892, pp. 260, 261,

pi. 10, figs. 1-9. {=Hemicyclops.)

Lichomolgus major Williams, 1907, pp. 77, 78, pi.

3. (=Myocheres.)
NUMBER 127 329

Lichomolgus (Sabelliphilus) spinosus Raffaele and

Monticelli, 1885, pp. 303-306, figs. 1-12. ( =

Midicola=Pseudomyicola.)

Myicola metisiensis Wright, 1885, and M. major

(Williams, 1907). Myicolidae. Listed in the

Lichomolgidae by C. B. Wilson, 1932, pp. 346-

348.

Pachysoma punctatum Claus, 1863, pi. 25, figs. 6-11.

(Transferred to the genus Pachos Stebbing, 1910,

p. 556).) Oncaeidae. Listed in the Lichomolgidae by Ganapati and Shanthakumari, 1962, p. 10,

and by A. Scott, 1909, p. 261.

Panaietis incamerata Stebbing, 1900, pp. 666, 667,

pi. 7IF.. Myicolidae. {Panaietis placed in Clausiidae by Yamaguti, 1936).

Parapanaietis Hoshina and Sugiura, 1953. Myicolidae. Placed in the Lichomolgidae by Gooding,

1963, p. 13.

Pseudomolgus leptostylis G. O. Sara, 1918, pp. 182.

183, pi. 103. (=Anthessius.)

Pseudomolgus navanacis C. B. Wilson, 1935, pp.

780, 781. pi. 26, figs. 13-24. (=Anthessius; see

Stock. Humes, and Gooding, 1965b, p. 2.) Pseudomolgus placed in the Lichomolgidae by San,

1916, p. 3; 1918, p. 181.

Pseudomolgus groenlandicus Ha men, 1923, pp. 19,

20, pi. 2, fig. 7a-g. ( = Anthessius.)

Pseudomolgus dilatatus G. O. Sars, 1918. p. 184, pi.

104 (=Anthessius.)

Pseudomolgus sp., in C. B. Wilson, 1923, p. 4.

(Probably = Anthessius.)

Rhinomolgus anomalus G. O. Sars, 1918, pp. 185-

187, pis. 105. 106. Myicolidae. Listed originally

in the Lichomolgidae by Sars, 1918, p. 184.

Synaptiphilus luteus Canu and Cu£not, 1892. Synaptiphilidae. (See Bocquet and Stock, 1957b, p.

695.)

Family UROCOPIIDAE Humes and Stock, 1972

Genus Vrocopia G. O. Sars, 1917

DIAGNOSIS.—Body cyclopiform, elongated. Urosome in the female 5-segmented. Caudal ramus

elongated, lamellar, slender proximally, pointed

distally, with four setae. Rostrum absent. First

antenna 6-segmented. Second antenna 4-segmented, with the last segment having a short,

straight claw (regarded by Sars as a fifth segment).

Labrum incised medially. Mandible lamellar

and falcate with the distal part of the base having

on its convex side a denticulated crest; lash short.

First maxilla with four elements. Second maxilla

of the general lichomolgid type. Maxilliped in

the female 3-segmented, the third segment pointed.

Legs 1-4 with 3-segmented rami. Armature of

the lichomolgidiform pattern. Leg 4 exopod with

the third segment having 11,1,5. Leg 4 endopod

with the formula 01;0—1;II. Leg 5 rudimentary

with two setae, without a free segment.

Other features as in the species below.

TYPE-SPECIES.—Urocopia singularis G. O. Sars,

1917.

REMARKS.—The male is unknown.

Urocopia singularis G. O. Sars, 1917

FIGURE 183

Urocopia singularis G. O. Sara. 1917b, pp. 5-11. figs. 1-15

[in plankton, 600-700 m. depth. 59*35' N, 20-40' E. south of

Iceland]-Lysholm and Nordgaard. 1921, p. 29.

FIGURE 183.—Urocopia singularis G. O. Sars. Female: a, dorsal;

b, caudal ramus; c, mandible and first maxilla; d, tip of

mandible. (Redrawn after G. O. Sars. 1917b. figs. 1, 7, 7a.

15.) Length of female 1.90 mm.
330 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

REMARKS.—We have been unable to locate type

or other specimens of Urocopia singularis. There

are none in the Zoologisk Museum, Bergen (letter

from Dr. Johanne Kjennerud, 2 February 1970),

and none in the Zoologisk Museum, Oslo (letter

from Dr. Marit Christiansen, 20 January 1970).

Family PSEUDANTHESSIIDAE Humes and

Stock, 1972

Lichomolgidae, Lichomolginae (part).—Gurney, 1927, p. 463.

Lichomolginae (part).-Sewell, 1949, p. 91.

First antenna usually 7-segmented, but 3 - or

possibly 6-segmented in Kombia. Legs 1-4 with a

reduction occurring in a posterior-to-anterior series. In Heteranthessius and Pseudanthessius with

3-segmented rami except leg 4 endopod which is

1-segmented or reduced to a small knob (in one

species of Heteranthessius). In Meomicola legs 1

and 2 with 3-segmented rami and leg 3 exopod 3 -

segmented, leg 4 exopod 1-segmented, legs 3 and

4 endopods absent. In Temnomolgus legs 1 and 2

with 3-segmented rami, leg 3 reduced to a small

sclerotization with two setae, leg 4 absent. In Kombia legs 1 and 2 with 3-segmented exopods and

2-segmented endopods, leg 3 exopod 3-segmented

but endopod absent, leg 4 absent. Leg 5 without

a free segment and represented by two or three

elements.

Key to Genera of the Family Pseudanthessiidae

1. Leg 4 absent 2

Leg 4 present, though in some cases much reduced 3

2. Legs 1 and 2 with 3-segmented exopods and 2-segmentcd endopods; second antenna with one

terminal claw KomMa

Legs 1 and 2 with 3-segmented rami; second antenna with two terminal claws Temnomolgus

3. Leg 3 with 3-segmented exopod, endopod absent; prosome truncated anteriorly .. Meomicola

Leg 3 with both rami 3-segmented; prosome not truncated anteriorly 4

4. Leg 4 with endopod represented only by a small segment or knob without spines or setae;

body of female transformed, with swollen prosome HeteranUietsku

Leg 4 with endopod 1-segmented with two elements; body of female cydopiform, not

transformed Pueudantheisnit

Genus Heteranthessius T. Scott, 1904

DIAGNOSIS—Body of the female transformed, the

prosome swollen. Body of the male cyclopiform,

elongated. Urosome in the female 5-segmented, in

the male 6-segmented. Caudal ramus with five or

six setae. First antenna 7-segmented. Second antenna 4-segmented, with one terminal claw.

Mandible consisting of a broad basal area and

a slender, attenuated recurved blade with denticulated edges. First maxilla with two elements. Second maxilla with the terminal armature simplified,

consisting of one spine and one claw (modified

lash). Maxilliped in the female 3-segmented with

a rounded rather than pointed tip, in the male

4-segmented (assuming that the proximal part of

the claw represents a fourth segment).

Legs 1-4 with 3-segmented rami except leg 4

endopod which consists of a single small segment

or is represented only by a small mucronate knob

with two terminal setae. Setae on all four legs

short. Leg 4 exopod with third segment having

111,1,5. Leg 5 without a free segment and represented only by two setae.

Other features as in the species below.

Found free or one species associated with an

ascidian.

TYPE-SPECIES.—Heteranthessius dubius (T. Scott,

1903).

REMARKS.—Keys for the three species cannot be

satisfactorily prepared, as only the male is known

for H. dubius and only the female for H. scotti

and H. furcatus.

Heteranthessius dubius (T. Scott, 1903)

Paranthessius dubius T. Scott, 1903a, pp. ISO, 131, pi. 6,

figs. 16-24 [from Clyde, Scotland]. (Paranthessius gen. nov.,

preoccupied by Paranthessius Claus, 1889. Replaced by

Heteranthessius T. Scott, 1904, p. 259).

REMARKS.—The female is unknown.
NUMBER 127

Heteranthessius fur cat us Stock, 1971

Heteranthessius furcatus Stock, 1971, pp. 5S5-S4O, figs. 1, 2

[from the ascidian Microcosmus sabatieri Runic, in 30-40

m, between Canet and St. Cyprien, southwestern (Mediterranean) eoast of France].

REMARKS.—The male is unknown.

Heteranthessius scotti Bocquet, Stock, and Blnard,

1959

Ficimx 184

Heteranthessius scotti Bocquet, Stock, and Benard, 1959, pp.

111-117, figs. 1 3 [in washings of the calcareous alga Lithophyllum incrustans Phil., Bretagne, France],

REMARKS.—The male is unknown.

Genus Kombia Humes, 1962

DIAGNOSIS.—Body transformed, elongated, the

prosome swollen, especially in the female. Urosoine with weakly denned segmentation, but apSS1

parently in the female 5-segmented and in the

male 6-segmented. Caudal ramus with five very

short setae. Rostrum weak, without a distinct posteroventral border. First antenna small, with reduced segmentation, 3- (possibly 6-) segmented.

Second antenna 4-segmented, with a single terminal claw.

Labrum a broad, transverse plate. Mandible with

the basal region beyond the indentation having on

its convex side a scalelike area with spinules and

on its concave side a row of spinules; lash forming

a short, serrated blade. Paragnath a small lobe.

First maxilla with three elements. Second maxilla

of the usual lichomolgid type. Maxilliped in the

female 3-segmented, with a pointed tip; in the

male possibly 4-segmented (if the proximal part

of the claw is assumed to represent a fourth segment).

Legs 1 and 2 with 3-segmented exopods and 2-

segmented endopods. Leg 3 with a 3-segmented

exopod but lacking an endopod. Leg 4 absent.

Reduced armature on these legs, similar in both

FICUKE HA.—Heteranthessius scotti Bocquet, Stock, and Benard. Female: a, dorsal; t>, ventral; c,

mandible and first maxilla; d, second maxilla; e, maxilliped; /, leg 4 and detail of the mucronate

knob representing the trace of the reduced endopod. (From Bocquet, Stock, and Benard, 1959,

figs. 1, 2d-f, Se.) Length of female 4.1-4.3 mm. Male unknown.
352

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 185.—Kombia angulata Humes, from Porites somaliensis in Mauritius. Female: a, first

antenna (A) ; b, second antenna (B) ; c, maxilliped (c) ; d, leg 1 (D) ; e, leg 2 (D) ; /, leg 3 (D) .

Scale: A, C, D, 0.05 mm; B, 0.1 mm.
NUMBER 127 333

sexes, without sexual dimorphism on leg 1 endopod. Leg 5 represented only by two setae, without a free segment. Leg 6 in the male represented

by two small posterolateral setae on the genital

segment.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES.—Kombia angulata Humes.

Kombia angulata Humes, 1962

FICURES 185. 186

Kombia angulata Humes, 1962, pp. 49-56, figs. I-S8 [from

the hard coral Psammocora sp., region of Nosy Be\ Madagascar]. -Bouligand, 1966, p. 27O.-Humes and Ho, 1968a,

pp. S71, 372 [from the hard corals Porites (Synaraea) sp.,

Porites sp. cf. nigrescent Dana, and Porites, young colony,

region of Nosy Bi, Madagascar].

NEW RECORDS.—3 ? ? , 3 cf cf from Porites sp.,

in 2 m, Ambariotelo, a small island almost between

Nosy Komba and Nosy B6, Madagascar, 22 August

1967, collected by AGH.

NEW HOST.—Porites somaliensis Gravier, in Mauritius: 6 9 9» 1 cT» depth about 4 m, offshore reef,

57°21' E, 20°22' S, 4 February 1964; 12 ? ? , 5

o"o*. depth 1-2 m, lagoon of Flic en Flacq, 13

February 1964; 1 ?, depth 1 m, same locality, 3

February 1964. All collected by JHS.

REMARKS.—The Mauritian material from Porites

somaliensis disagrees in some minor points with

that described in the original description (host

Psammocora in Madagascar). Humes and Ho

(1968a) recorded K. angulata from Porites but did

not mention any deviating characters in relation

to this different host.

FIGURE 186.—Kombia angulata Humes. Female: a, dorsal; b, ventral; c, lateral; d, first antenna;

e, mandible, /, /eg 1; g, leg S; h, leg 5. Male: i, dorsal; ;, lateral. (From Humes, 1962, figs. 1-3,

7, 10, 16, 20-23.) Length of female 1.40 mm, of male 0.97 mm.
334 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

The present specimens deviate in the following

ways: (1) the first antenna of the female is clearly

5-segmented (versus 3-segmented), through subdivision of the slender terminal portion (Figure

185a); (2) the second antenna in both sexes has a

slightly more slender claw (Figure 1856); (3) the

maxilliped of the female has more slender terminal processes on the third segment (Figure 185c)

and bears only one seta on the second segment;

(4) leg 1 of the female has longer exopod spines,

and on the endopod the spines are about as long

as the seta (Figure 185a*); (5) the exopod spines

in leg 2 are slightly longer (Figure 185c); and (6)

the exopod spines in leg 3 are much longer (Figure

185/).

On the other hand, complex structures such as

the first antenna and the maxilliped of the male

are very similar to those figured by Humes (1962).

In the light of the variability (especially in the

chaetotaxis of the legs) recorded by Humes (1962),

it seems wise to consider the present material and

the original samples as one variable species.

Genus Meomicola Stock, Humes, and Gooding,

1963

DIAGNOSIS.—Body modified, the prosome truncated anteriorly. Urosome in the female 3-segmented, in the male 4-segmented. Caudal ramus

with six short setae. Rostrum probably the median

part of a broad frontal plate with short lateral

horns. First antenna 7-segmented, in both sexes

with the formula 3, 11, 3, 4, 4, 3, 8, including aesthetes. Second antenna 4-segmented, the armature

being 1, 1, 2 + a large hooklike claw, and four

short, bent, dawlike elements -f- three setae.

Labrum incised medially. Mandible consisting

of a small base and a curved, attenuated, bipectinate blade. First maxilla with two elements.

Second maxilla of the usual lichomolgid type. Maxilliped of the female 3-segmented with a pointed

tip, in the male 4-segmented (assuming that the

proximal part of the claw represents a fourth segment).

Legs 1 and 2 with 3-segmented rami. Leg 3 with

3-segmented exopod, the endopod absent. Leg 4

with a protopod and 1 exopod segment. These legs

with reduced armature which is alike in both sexes.

Leg 5 represented by three setae, without a free

segment. Leg 6 in the male represented by the ventral posterolateral flap on the genital segment having two setae.

Other features as in the species below.

Associated with echinoid echinoderms.

TYPE-SPECIES.—Meomicola amplectans Stock,

Humes, and Gooding.

Meomicola amplectans Stock, Humes, and

Gooding, 1963

FIGURE 187

Meomicola amplectans Stock, Humes, and Gooding, 196Sa,

pp. 57-71, figs. 20-24 [from the echinoid Meoma ventricosa

(Lamarck), Curasao, Jamaica].

NEW RECORD.—One large-form 9» 5 small-form

9 9»

 ant* 2 small-form <$&, free in 6 m, Station

no. 26-68, Long Island, Bahamas, 22°52'10" N,

74°63'45" W, collected by A. Fosshagen.

Genus Pseitdanthessius Clans. 1889

DIAGNOSIS.—Body cyclopiform. Urosome in the

female 5-segmented, in the male 6-segmented except in P. deficient (female 4-, male 5-segmented)

and in P. dubius (female 4-segmented, male unknown). Caudal ramus with six setae. Rostrum

not well developed. First antenna 7-segmented, in

the female with the armature 4, 13, 6, 3, 4 + 1

aesthete, 2 + 1 aesthete, and 7 —f- 1 aesthete; in

the male with the same armature or in some species with an additional aesthete on the second segment. [In P. luculentus, P. notabilis, and P. liber

(sensu Sewell, non Brady and Robertson) with one

or two very long elements on the first segment.]

Second antenna 4-segmented, with the formula

1,1,3 and variable terminal armature (1-4 claws or

clawlike spines and a few setae; or, in P. tortuosus,

two lamelliform elements, one seta, and three

spines; or, in P. mucronatus, all elements setiform).

Labrum deeply incised medially. Mandible with

the basal region having its convex edge smooth or

with a weak scalelike area or a process and on its

concave edge a row of spinules; lash long or short.

Paragnath a small lobe. First maxilla with three

or four elements. Second maxilla of the usual

lichomolgid type. Maxilliped in the female 3-segmented, with a pointed tip; in the male 4-segmented (assuming that the proximal part of the

claw represents a fourth segment).
NUMBER 127 335

Legs 1-4 with 3-segmented rami, except for leg 4

endopod which is 1-segmented with two distal elements. Leg 1 endopod of the male with the same

armature as in the female, or with the same formula but with slight differences in the form of the

processes or elements, or in P. deficiens 1,1,4 or in

both P. tortuosus and P. aestheticus 1,1,1,4. (In

nearly one-third of the species males are unknown.)

Leg 5 represented by three elements (a subterminal

seta and a terminal spine and seta); no free segment. Leg 6 represented by the two setae near

the genital openings.

Other features as in the species below.

Associated with various invertebrates, chiefly turbellarians, nemerteans, polychaetes, crinoids, asteroids, ophiuroids, and echinoids.

TYPE-SPECIES.—Pseudanthessius gracilis Claus.

REMARKS.—Although the genus Pseudanthessius

is inconveniently large (containing 34 species), attempts to discover natural divisions within it are

FIGURE 187.—Meomicola amplectans Stock, Humes, and Gooding. Female, small form: a, dorsal;

b, lateral; c, second antenna; d, mandible; e, leg 2; /, leg S; g, leg 4; A, legs 5 and 6 in situ. Male,

small form: t, dorsal; /, maxilliped. (From Stock, Humes, and Gooding, 1963a, figs. 20a,b.e,

21b,g-i, 22b,f, 23a.) Length of small-form female 0.82 mm, of small-form male 0.57 mm.
336 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

plagued by incomplete descriptions (for example,

of the mandible) and by the fact that males are

unknown in about one-third of the species. The

genus appears to be much more variable than other

genera considered here. Nevertheless, certain features, such as the form of the mandible and the

terminal armature of the second antenna, suggest

that it may be possible to arrange the species in

several groups. However, since information is lacking about many of the species, not all members

of the genus can at present be assigned to such

groups. The existing descriptions of species of

Pseudanthessius demonstrate by their incompleteness the need for detailed and careful descriptions

in order to provide the basis for revisionary work.

We have noted the following similarities in the

species, but without any intention of arranging

them in definite taxonomic groups. (Undoubtedly,

when all the species are better known other groupings will become apparent.)

(1) Mandible with a long lash and without a tooth or

scale on the convex side; with one or two very long elements

on the first segment of the first antenna; second antenna

with two claws: P. notabilis, P. luculentus, and probably P.

liber sensu Sewell.

(2) Mandible with a long lash and without a tooth or

scale on the convex side; elements on the first segment of the

first antenna not unusually long; second antenna with two

claws: P. procurrens and P. pusillus.

(3) Mandible with a long lash and without a tooth or

scale on the convex side; elements on the first segment of the

first antenna not unusually long; second antenna with three

claws: P. assimilis, P. liber, and P. weberi. (Perhaps P.

pectinifer belongs here also, but it differs in having a small

spinelike element on the convex side of the mandible.)

(4) Mandible with a broad blade and short lash, with a

toothlike process or scale on the convex side; second antenna

with two claws: P. angularis, P. ferox, P. madrasemis, P.

major, P. minor, and P. rostellatus.

(5) Mandible with a short lash, without a tooth or scale

on the convex side; second antenna with several slender

setiform claws: P. gracilis and P. sauvagei.

(6) Mandible with a relatively broad blade, but with

almost no lash and without a tooth or scale: P. aestheticus,

P. obscurus, and P. tortuosus.

(7) Urosome of ft-malc 4-srgmented instead of 5-iegmented: /'. deficiem and P. dubius.

A key to the males has not been prepared because that sex is unknown in so many of the species. In the following key to the females, the species P. anormalus, P. brevicauda, and P. minutus

are omitted because of insufficient information; in

addition, the female of P. faouzii is unknown.

Partial Key to Species of the Genus Pseudanthessius

FEMALES

1. Urosome 4-segmented 2

Urosome 5-segmented 3

2. Each postgenital segment about two times longer than wide; claw on second antenna

slender and gently recurved P. dubius

Each postgenital segment about as long as wide; claw on second antenna stout and strongly

recurved P. deficient

3. At least one element on basal segment of first antenna almost as long as rest of appendage... 4

Elements on basal segment of first antenna less than half as long as rest of appendage 6

4. Two elements on basal segment of first antenna much better developed than the others . 5

A single element on basal segment of first antenna better developed than the others

P. luculentus

6. Terminal spine on exopods of legs 2-4 strongly curved; distal segment of second antenna

about six times as long as wide P. liber sensu Sewell

Terminal spine on exopods of legs 2-4 almost straight; distal segment of second antenna

about three times as long as wide P. notabilis

6. Caudal ramus about as long as wide or slightly longer than wide (but distinctly less than

1-5:1) 7

Caudal ramus distinctly more than 1.5:1 13

7. Two claws on second antenna with denticles 8

Two claws on second antenna smooth 9

8. Fourth segment of second antenna very long and slender, more than eight times longer

than wide p . foliatus

Fourth segment of second antenna shorter, about 3.5:1 P. pectinifer
NUMBER 127 337

9. Rostrum snoutlike; pair of conspicuous spines on dorsal surface of genital segment

P. rostellatus

Rostrum weakly developed, not snoutlike; genital segment without such a pair of spines ...10

10. Genital segment abruptly indented posteriorly with two acute posterolateral angles

P. angularit

Genital segment with broadly rounded lateral margins without acute angles 11

11. Prosome slender (ratio of length to width 1.96:1); caudal ramus slightly longer than wide

(about 1.3:1) p. minor

Prosome broader (ratio of length to width 1.37-1.45:1); caudal ramus about as long as

wide 12

12. Cephalosome broadly subtriangular; spine on terminal segment of leg 1 endopod straight

P. madrasensu

Cephalosome broadly rounded; spine on terminal segment of leg 1 endopod curved

P. major

13. Caudal ramus close to two times longer than wide 14

Caudal ramus more than two times longer than wide 15

14. Body length 1-1.3 mm; claws on second antenna slender, setiform P. sauvagei

Body length 052 mm; claws on second antenna recurved and unguifonn P. pusillus

15. Caudal ramus not more than four times longer than wide 16

Caudal ramus more than four times longer than wide 23

16. Lateral margins of genital segment produced medially into conspicuous pointed expansions;

two elements on third segment of second antenna modified as slender jointed claws

P. spinijer

Lateral margins of genital segment produced at most into small dorsal processes (P. tenuis),

not conspicuously expanded; not more than one element on third segment of second

antenna modified as a claw 17

17. Outer margin of leg 4 endopod smoothly convex 18

Outer margin of leg 4 endopod sinuate, or broken by a swelling or indentation which may

become a conspicuous knob or notch 20

18. Length of leg 4 endopod more than four times the width; second antenna terminally with

recurved claws 19

Length of leg 4 endopod less than three times the width; second antenna terminally with

modified armature not including recurved claws P. tortuotus

19. Caudal ramus a little longer than anal segment; latter about as long as preceding segment

P. liber

Caudal ramus one-fourth longer than anal segment; latter nearly twice as long as preceding segment P. asdmOis

20. Leg 4 endopod with outer margin bulging or sinuate, without a pronounced notch or

indentation 21

Leg 4 endopod with outer margin regular, with a pronounced notch or indentation 22

21. Leg 4 endopod no longer than basal segment of exopod, with proximal bulge on outer

edge but no notch P. mueronatus

Leg 4 endopod much longer than basal segment of exopod, with outer margin distally

sinuate P- jerox

22. Leg 4 endopod with marked notch on outer edge at proximal third; segments of first

antenna short and compact P- obscunu

Leg 4 endopod with notch at center of outer edge; not all segments of first antenna short

and compact ?• tenuis

23. Caudal ramus not more than 7.5 times as long as wide 24

Caudal ramus at least ten times as long as wide 50

24. Greatest width of leg 4 endopod over one-half length P. lofcu

Greatest width of leg 4 endopod less than one-half length 25

25. Leg 4 endopod on the outer edge with a notch or at least a break in the sclerotization 26

Leg 4 endopod with outer edge lacking a notch or break in the sclerotization 29

26. Genital segment elongated, not much expanded laterally 27

Genital segment expanded in its anterior part but abruptly constricted in its posterior

part 28

27. Genital segment with a pair of anterior lateral lobes; caudal ramus about 4.5:1

P. fitOCUl f €f 15

Genital segment without such lobes: caudal ramus about 5:1 P. weberi
338 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

28. Leg 4 endopod less than four times as long as wide; all elements on caudal ramus setiform

P. gracilis

Leg 4 endopod more than five times as long as wide; two elements on caudal ramus flattened

and spinelike P. gracUoides

29. Caudal ramus 4-5 times as long as wide; leg 4 endopod with little proximal swelling

P. titmertophilui

Caudal ramus 6-7 times as long as wide; proximal swelling on leg 4 endopod well marked

P. anthcticus

30. Last two segments of urosome subequal in length P- concinmu

Anal segment at least twice as long as preanal P« thoreUi

Pseudanthessins gracilis Claus, 1889

Pseudanthessius gracilis Claus, 1889, pp. 18, 19, pi. 4, figs.

1-7.-T. Scott and A. Scott, 1893a, pp. 241-242, pi. 12,

figs. 15-20 [from the polychaete Filograna, Moray Firth,

Scotland] .-Graeffe, 1900, p. 41 (=Lichomolgus thorelli

Brady) [with algae, Trieste, Italy].—Thompson and A.

Scott, 1903, p. 276, pi. 14, figs. 19-23 [from washings of

invertebrates, Ceylon].—T. Scott, 1894b, pp. 233, 258, 260

[from Firth of Forth, Scotland]; 1906, p. 353 [from Musselburg, Firth of Forth, Scotland].—Norman and T. Scott,

1906, p. 198 [from £ngland].-G. O. Sars, 1917a, pp. 167,

168, pi. 93 [from Norway].-Lang, 1949, pp. 8, 9. figs. 10.

11 [on the red alga Rhodymenia palmata, Smedjan,

Sweden].—Marine Biological Association, 1957, p. 176-

Bresciani and Lutzen, 1962, p. 377.—Bocquet, Stock, and

Kleeton, 1963, p. 38 [from the polychaete Pomatoceros

triqueter (L.), Roscoff, France, and from the bivalve

Spisula subtruncata (Da Costa), Brest, France].

Pseudanthessius aestheticus Stock, Humes, and

Gooding, 1963

Pseudanthessius aestheticus Stock, Humes, and Gooding

1963a, pp. 23-34, figs. 6-8 [from the polychaete Hermodice

carunculata (Pallas) and probably Eurythoe complanata

(Pallas), Jamaica].

Pseudanthessius angularis Humes and Ho, 1970

Pseudanthessius angularis Humes and Ho, 1970b, pp. 12-15,

figs. 52-66 [from the conoids Stephanometra indica (Smith)

and Dichrometra spJ afra A. H. Clark, region of Nosy Be,

Madagascar].

Pseudanthessius anormalus Ummerkutty, 1966

Pseudanthessius anormalus Ummerkutty, 1966, pp. 108-110,

figs. 1-10 [from sponge washings, Gulf of Mannar, southeastern India].

Pseudanthessius assimilis G. O. Sars, 1917

Pseudanthessius assimilis G. O. Sars, 1917a, pp. 170, 171,

pi. 95 [from the ecbinoid Echinus elegans, Norway].

Pseudanthessius brevicauda Ummerkutty, 1966

Pseudanthessius brevicauda Ummerkutty, 1966, pp. 110-112,

figs. 11-17 [from washings of littoral weeds. Gulf of Mannar, southeastern India].

Pseudanthessius concinnus Thompson and A. Scott,

1903

Pseudanthessius concinnus Thompson and A. Scott, 1903, p.

277, pi. 14, figs. 24-30 [from washings of invertebrates, Ceylon].

REMARKS.—The male is unknown.

Pseudanthessius deficiens Stock, Humes, and

Gooding, 1963

Pseudanthessius deficiens Stock, Humes and Gooding, 1963a,

pp. 45-57, figs. 13-19 [from the ophiuroid Ophioderma

cinereum Miiller and Troschel, Curasao, St. Martin. Puerto

Rico; a collection from the holothurian Ludwigothuria

mexicana (Ludwig) in Curasao probably represents contamination].

Pseudanthessius dubius G. O. Sars, 1918

Pseudanthessius dubius G. O. Sars, 1918, pp. 214, 215, pi.

117 [from Norway].—Bresciani and Lutzen, 1962, p. 402

[from the asteroid Asterias rubens, Swedish west coast}.

REMARKS.—The male is unknown.

Pseudanthessius faouzii Steuer, 1940

Pseudanthessius faouzii Steuer, 1940, pp. 21-24, figs. 9-14

[from Amphioxus-sznd, Alexandria, Egypt].

REMARKS.—The female is unknown.

Pseudanthessius ferox Humes and Ho, 1967

Pseudanthessius ferox Humes and Ho, 1967c, pp. 382-386,

figs. 30-59 [from the polychaete Sabella fusca (Grube),

region of Nosy Be, Madagascar].
NUMBER 127

Pseudanthessius foliatus Stock, 1967

Pseudanthessius foliatus Stock, 1967a. pp. 235-238, figs, lad,

2a,b, 3ae [from the cchinoid Echinothrix calamaris (Pallas), Gulf of Aqaba, Israel}.

Pseudanthessius graciloides Sewell, 1949

Pseudanthessius graciloides Sewell, 1949, pp. 123-125, fig.

34A-F [in weed-washings, Addu Atoll, Maldive archipelago).

REMARKS.—The male is unknown.

Pseudanthessius lotus Illg, 1950

Pirudanthessius latus Illg, 1950, pp. 130-133, fig. 1 [from

the i in In ll.ii i.i11 Cryptophallus magnus Freeman, San Juan

Island, Washington, and from unidentified turbcllarians,

California and Washington].

Pseudanthessius ubscuius.—Wilson, 1935, p. 780, pi. 27, fig.

36 [from turbellaiians, California].

Pseudanthessius liber (Brady, 1880)

l.ichomolgus liber Brady and Robertson, 1876, p. 197 [nomen nudum].-Brady, 1880, pp. 44-46, pi. 86, figs. 1-13

[from England]-Thompson. 1893, p. 207, pi. 25, fig.

2a,b [from Liverpool Bay, England]; 1903, p. 34 [in 270

fathoms at 52°4.5' N, 12°27' W].-Herdman, 1896, p.

52.-Stock, Humes, and Gooding, 1963a, p. 10.

Pseudanthessius liber.-T. Scott and A. Scott, 1893a, p. 240.—

T. Scott, 1894b, pp. 233. 258 [from Firth of Forth, Scotland]; 1897a, p. 154 [from Loch Gair, off Inveraray, Upper

Loch Fyne, Scotland]: 1902, p. 470 [from North Craig,

Firth of Forth, Scotland]; 1906, p. 353 [from North Craig

and Inchkeith, Firth of Forth, Scotland].—d'Arcy Thompson, 1901, p. 47.-Thompson and A. Scott, 1903, p. 277

[from washings of invertebrates, Ceylon].—Pearson, 1905,

p. 163 [from Lough Swilly, Ireland].-G. O. Sars, 1916,

p. 3; 1917a, pp. 169, 170, pi. 94 [from the echinoids

Echinus sphaera and Strongylocentrotus droebakiensis,

Norway].—Stephensen, 1929, pp. 3. 12 [from the Faroes].—

Bocquet, 1952, p 502 [from the echinoids Paracentrotus

lividus, Psammechinus miliaris, and Echinus esculentus,

region of Roscoff, France].—Gooding, 1957, p. 212 [from

the echinoid Echinus esculentus, Plymouth, England].—

Marine Biological Association, 1957, p. 176].—Bocquet and

Stock, 1962a, p. 90 [from the asteroid Marthasterias glacialis, region of Roscoff, northern France].—Bresciani and

Liitzen, 1962, pp. 377, 378 [from Echinus esculentus,

Flatholmen, mouth of Gullmar Fiord, Swedish west

coast].—Bocquet, Stock, and Louise, 1963, pp. 149-152, fig.

1 [from the echinoids Echinus esculentus Linnaeus, Psammechinus miliaris (Gmelin), and Paracentrotus lividus

(Lamarck), region of Roscoff, France].—Bruce, Colman,

and Jones, 1963, p. 128.-Gotto. 1966, p. 194.—Barel and

Kramers, 1970, table 5 [from the echinoids Echinus escu339

lentus Linnaeus and Psammechinus miliaris (Gmelin),

Roscoff, France, and Paracentrotus lividus (Lamarck), Morgat. Channel coast of France].

Pseudanthessius liber sensu Sewell, 1949

Pseudanthessius liber sensu Sewell, 1949, non Brady and

Robertson, 1876.-Sewell, 1949, p. 121, fig. 32A-C [from a

short-spined, regular echinoid, Nankauri Harbor, Nicobar

Islands].

Pseudanthessius luculentus Humes and Cressey,

1961

Pseudanthessius luculentus Humes and Cressey, 1961b, pp.

75-81, figs. 23-56 [from the echinoid Stomopneustes variolaris (Lamarck), Nosy Be, Madagascar].—Ummerkutty,

1968, pp. 314, 315.

Pseudanthessius agilis Ummerkutty, 1962. pp. 43-47, fig. 7

(1-11), fig. 8 (1-3) [from sponge washings, Gulf of Mannar, southeastern India; misspelled Pseudoanthessius].—

Stock, Humes, and Gooding. 1963a, p. 10.

Pseudanthessius madrasensis Reddiah, 1968

FIGURE 188

Pseudanthessius madrasensis Reddiah, 1968b, pp. 320-323,

figs. 1, 2 [from crinoids, Madras, India].—Humes and Ho,

1970b, pp. 1-5, figs. 1-28 [from the crinoid Tropiometra

carinata (Lamarck), region of Nosy Be, Madagascar].

Pseudanthessius major Stock, 1967

Pseudanthessius major Stock, 1967a, pp. 243-248, figs. 7, 8

[from the crinoids Heterometra savignyi (J. Muller) and

iMtnprometra klunzingeri (Hartlaub), Gulf of Aqaba, Israeli-Humes and Ho, 1970b, pp. 5-11, figs. 29-44 [from

the crinoids Cenometra emendatrix (Bell), Heterometra

africana (A. H. Clark), Stephanometra indica (Smith),

Lamprometra klunzingeri (Hartlaub), Liparometraf sp.,

and Dichrometra sp.? afra A. H. Clark, region of Nosy

IW. Madagascar].

Pseudanthessius minor Stock, 1967

Pseudanthessius minor Stock, 1967a, pp. 238-242, figs. 4-6

[from the crinoid Lamprometra klunzingeri (Hartlaub),

Gulf of Aqaba, Israel].-Humes and Ho, 1970b, pp. 11, 12,

figs. 45-51 [from the crinoids Heterometra africana (A.

H. Clark), Lamprometra klunzingeri (Hartlaub), Liparometra? sp., and Dichrometra sp.? afra A. H. Clark, region

of Nosy Be, Madagascar.

Pseudanthessius minutus Reddiah, 1968

Pseudanthessius minutus Reddiah, 1968b, pp. 323-328, figs.

3, 4 [from tunicates, Madras, India].
340

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

FIGURE 188.—Pseudanthessius madrascnsis Rcddiah. Female: a, dorsal; b, urosome; c, second

antenna; d, mandible and first maxilla; e, leg 4 and intcrcoxal plate; /. leg 5. Male: g, dorsal.

(From Humes and Ho, 1970b, figs. 1, 2, 7, 10, 17-19.) Length of female O.f>8 mm, of male

0.48 mm.

Pseudanthessius mucronatus Gurney, 1927

Pseudanthessius mucronatus Gurney, 1927, pp. 469, 470, fig.

115 [from Suez Canal].

REMARKS.—The male is unknown.

Pseudanthessius obscurus A. Scott, 1909

Pseudanthessius obscurus A. Scott, 1909, pp. 270, 271, pi. 69,

figs. 8-14 [from plankton off Laiwui, coast of Obi Major,

0°24' S, 127°36' E].-Sewell, 1949, pp. 121-123, fig. 33A-I

[in weed-washings, Nankauri Harbor, Nicobar Islands].

Pseudanthtssius nemertophilus Gallien, 1936

Pseudanthessius nemertophilus Gallien, 1936, pp. 444—459,

figs. 1-3 [from the nemertean Lineus longissitnus Sowerby,

Wimereux, France].

Pseudanthessius notabilis Humes and Cressey, 1961

Pseudanthessius notabilis Humes and Cressey, 1961b, pp.

67-75, figs. 1-31 [from the echinoid Echinometra mat had

(Blainville), Nosy Be, Madagascar].—Stock, 1967a, pp. 232-

235, fig. 2c,d [from the echinoid Echinometra mathaei,

Gulf of Aqaba, Israel].

Pseudanthessius pectinijer Stock, Humes, and

Good ing, 1963

Pseudanthessius pectinifer Stock, Humes, and Gooding,

1963a, pp. 15-23, figs. 3-5 [from the echinoid Clypeaster

rosaceus Linnaeus, Bahamas, Puerto Rico, Jamaica].—Ummerkutty, 1966, pp. 107, 108.

Pseudanthessius procurrens Humes, 1966

Pseudanthessius procurrens Humes, 1966b, pp. 2-7, figs. 1-29

[from the echinoid Phyllacanthus imperialis (Lamarck),

region of Nosy Be, Madagascar].
NUMBER 127 341

Pseudanthessius pusillus Humes, 1969

Pseudantltessius pusillus Humes, 1969c, pp. 268-277, figs.

1-30 [from the echinoid Echinodiscus auritus Leskc, region

of Nosy Be, Madagascar].

Pseudanthessius rostellatus Humes and Ho, 1970

Pseudanthessius rostellatus Humes and Ho, 1970b, pp. 15-20,

figs. 67-90 [from the crinoid Comaster sp.? nov. aff. distinctus (P. H. Carpenter), Nosy Be, Madagascar].

Pseudanthessius sauvagei Canu, 1891

Pseudanthessius sauvagei Canu, 1891a, p. 481 [from the

echinoid Echinocardium cordatum Pennant, Pointe aux

Oies. Wimereux, and Boulogne, France]; 1892, pp. 243,

244, pi. 25, figs. 1-17 [from the echinoid Echinocardium

cordatum, Channel coast of France].—T. Scott, 1894b, pp.

23S, 258, 260 [from Firth of Forth, Scotland]; 1905, p. 202

[from the echinoid Echinus esculentus, Scotland]; 1906, pp.

S.r>3, 354 [from Echinus esculentus, St. Monans, Firth of

Forth, Scotland].—T. Scott and A. Scott. 1894. p. 146 [from

off St. Monans, Firth of Forth, Scotland].—Hcrdman, 1895,

p. 45 [record from I. C. Thompson: from the echinoid

Spatangus purpureus, Port Erin, F.ngland].—Thompson,

1895, p. 102 [from Spatangus purpureus, Port Erin, EnglandJ.-G. O. Sars. 1917a. pp. 171, 172, pi. 96 [from Norway].—Bocquet, 1952, p. 502 [from Echinocardium

cordatum, region of Roscoff, northern France].—Stock,

1960c, p. 248 [from Spatangus purpureus O. F. Miiller,

Cap Bear, near Banyuls, Mediterranean coast of France].—

den Hartog, 1961, p. 29 [from Echinocardium cordatum,

The Netherlands].-Bresciani and Liitzen, 1962, p. 378

[from Spatangus and Echinocardium flavescens, Bonden,

Swedish west coast].—Barel and Kramers, 1970, p. 166,

table 5 [from Echinocardium cordatum, Morgat, France,

and Salcombe (Plymouth), England].

Pseudanthessius spinifer Lindberg, 1946

Pseudanthessius spinifer Lindberg, 1946, pp. 81-84, figs, a-1

[from Bombay, India].

Pseudanthessius tenuis Nicholls, 1944

Pseudanthessius tenuis Nicholls, 1944, pp. 55, 56, fig. 25

[from South Australia].

NEW RECORD.—1 9 in plankton net towed

obliquely from bottom to mid-depth (total depth

10 m). Surface temperature 16° C, bottom temperature 15.6° C. Bottom salinity approximately

37 percent. One-half mile from south of Yarra

River, Port Phillip Bay, Victoria, Australia. Collected 5 May 1969 by G. H. Arnott of Monash University, who has permitted us to include his record

here.

REMARKS.—The male is unknown.

Pseudanthessius thorelli (Brady, 1880)

I.ichomolgus thorellii —Brady and Robertson, 1876, p. 197

[nomen nudum].-Brady, 1880, pp. 47^*9, pi. 88, figs. 1-19

[from Englandj.-Thompson, 1893, p. 207, pi. 25, fig. 2c

[from Liverpool Bay, England].—Stock, Humes, and Gooding, 1963a, pp. 10, 32.

Pseudanthessius thorellii—T. Scott and A. Scott, 1893a, p.

240.-T. Scott, 1894b, pp. 233, 258, 259 [from Firth of

Forth, Scotland]; 1900, p. 399 [from Clyde area, Scotland].—Norman and T. Scott, 1906, p. 198 [from the bivalve Pecten opercularis, Plymouth and Salcombe,

England].

Pseudanthessius gracilis (syn. Lichomolgus thorelli).—Graeffe,

1900, p. 41 [with algae, Trieste, Italy].

Pseudanthessius thorelli.-T. Scott, 1906, p. 353 [St. Monans

and Inchkeith, Firth of Forth, Scotland].—Monod and

Dollfus, 1932a, p. 153 [from Chlamys (Aequipecten) opercularis (Linnaeus)].—Leigh-Sharpe, 1935, p. 47 [from the

mantle cavity of Chlamys (Aequipecten) opercularis (Linnaeus), England].—Marine Biological Association, 1957, p.

176.—Bresciani and Liitzen, 1962, p. 378 [from the asteroid

Asterias rubens, near Flatholman, Gullmar Fiord, Swedish

west coast].

Pseudanthessius tortuosus Stock, Humes, and

Gooding, 1963

Pseudanthessius tortuosus Stock, Humes, and Gooding, 1963a,

pp. 34-45, figs. 9-12 [from the polychaete Hermodice

carunculata (Pallas) , Puerto Rico, Jamaica, and Barbados].

Pseudanthessius weberi A. Scott, 1909

Pseudanthessius weberi A. Scott, 1909, pp. 267-268, pi. 67,

figs. 18-24 [from washings of dredged invertebrates, midway between Lucipara and Schildpad Islands, 1395 m,

5° 26.7' S, 127° 36.5' E].

Genus Temnomolgus Humes and Ho, 1966

DIAGNOSIS.—Body transformed, with a broad

swollen prosome in the female; somewhat less transformed in the male. Urosome 5-segmented in the

female, 6-segmented in the male, but in both sexes

the last two segments clearly separated only dorsally. Caudal ramus with six setae. Rostrum

linguiform. First antenna 7-segmented, the formula in the female being 3, 13, 4, 5, 3 -|- 1 aesthete,

2 —j— 1 aesthete, and 7 —J— 1 aesthete; in the male
342 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCV

3, 12 + 1 aesthete, 4, 4 -j- 1 aesthete, 3 -f- 1 aesthete, 2 + 1 aesthete, and 7 -f 1 aesthete. Second

antenna 4-segmented, with the formula 1, 1, 3, and

II -(- 4 small setae, there being two terminal claws.

Labrum with a median notch between the two

lobes. Mandible with the base divided into two

areas by an indentation, the proximal area unornamented, the distal area having on its convex side

a scale bearing a row of spinules followed by a

striated fringe and on its concave side a similar

spinulose scale and a row of hairlike setules; lash

moderately long. Paragnath an elongated hairy

lobe. First maxilla with two elements. Second

maxilla of the lichomolgid type. Maxilliped in the

female 3-segmented, terminating in pointed processes, in the male 4-segmented (considering the

proximal part of the claw as a fourth segment).

Legs 1 and 2 with 3-segmented rami. Leg 2 exopod with the formula I—0;0—1;II,I,4, the outer

spine on the second segment being absent. Leg 3

reduced to a small sclerotization bearing two setae.

Leg 4 absent. Male with similar segmentation and

armature of the legs. Leg 5 in both sexes rudimentary, without a free segment, and consisting of

only three setae. Leg 6 in the female represented

by two small spines near the area of attachment

of each egg sac, in the male by a posteroventral

Hap on the genital segment bearing two setae.

Other features as in the species below.

Associated with zoanthidean coelenterates.

TYPE-SPECIES.—Temnomolgus eurynotus Humes

and Ho.

Temnomolgus eurynotus Humes and Ho, 1966

189

Temnomolgus eurynotus Humes and Ho. 1966a, pp. 7-10,

figs. 29-55 [from the gastrovascular cavity of the zoanthklcan Pnlythoa tuberculosa (Fspcr). region of Nosy Be.

northwestern Madagascar].

9

FIGURE 189.—Temnomolgus eurynotus Humes and Ho. Female: a, dorsal; b, ventral; c, lateral;

d, urosome; e, mandible; /, leg 1 and intercoxal plate; g, leg S. Male: h, dorsal. (From Humes

and Ho, 1966a, figs. 29-32, 41, 47, 49, 50.) Length of female 1.18 mm, of male 1.21 mm.
NUMBER 127 343

Pseudanthessiidae Insufficiently Described or of

Uncertain Position

Pseudanthessius sp., Herbst, 1955

Pseudatithessius sp., Herbst, 1955, p. 218 [from S3o Scbastiao, s."n> Paulo, Brasil].

Family RHYNCHOMOLGIDAE Humes and Ho,

1972

Genus Rhynchotnolgus Humes and Ho, 1967

DIAGNOSIS.—Body transformed, in the female

with expanded cephalosome, narrow metasome and

urosome; in the male the body elongated and almost vermiform. Urosome 5-segmented in the

female, with the postgenital segments sometimes

telescoped. Urosome in the preadult male 5-segmented, in the adult male 6-segmented. Caudal

r.nuns with six short setae in the female; in the

male reduced, lobate, without distinct articulation

with the anal segment, and bearing three spinules.

Rostrum a conspicuous tumid snoutlike lobe. First

antenna in the female short and 5-segmented, with

the formula 3, 10, 4 -|- 1 aesthete, 2 -f- 1 aesthete,

and 7 -f- 1 aesthete; in the male very short, 1-segmented, with 13 short terminal spines. Second antenna in the female 4-segmented and much longer

than the first antenna, with the formula 1, 1, 3, I

-(- 5 spinules, there being a single terminal claw.

Second antenna in the male probably 3-segmented,

terminally with eight short spines. In both sexes a

conspicuous lobe external to the base of the second

antenna and behind the first antenna.

Mouthparts of the female: Labrum with the two

lobes separated by a median indentation. Mandible with the base divided into two areas by an indentation. The proximal area unornamented, the

distal area bearing on its convex side a large, digitiform, posteriorly directed process and a row of

small dentations and on its concave side a row of

spinules; lash relatively short and stout. ParagFICURE 190.—Rhynchomolgus corallophilus Humes and Ho. Female: a, dorsal; b, lateral; c, first

antenna; d, mandible; e, second maxilla; /, maxilliped; g, leg 1; h, leg 3; i, leg 4; ;, leg 5. Male:

*, dorsal; /, second maxilla. (From Humes and Ho, 1967b, figs. 80, 81, 86, 90, 92, 93, 95, 97-99,

112, 115.) Length of female 1.07 mm, of mature male 2.04 mm, of small male 1.17 mm.
344 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

nath a small hairy lobe. First maxilla with two

elements. Second maxilla of the lichomolgid type.

Maxilliped 3-segmented, terminating in a pointed

process.

Mouthparts of the male: Labrum a triangular

bulbous area without posteroventral lobes or median cleft. Postlabral region with a fold overlapping the end of the labrum. Mandibles, paragnaths, and first maxillae absent. Second maxilla in

the preadult male a well-elongated, recurved hook;

in the adult male a small spinous process. Maxilliped in the preadult male 1-segmented with a

clawlike tip; in the adult male absent.

Legs 1-4 much modified. Legs of the female:

Legs 1, 2 with 2-segmented rami. Leg 3 reduced to

2 setae and leg 4 to 1 seta, both legs lacking intercoxal plates. Leg 5 without a free segment and

reduced to three setae. Leg 6 represented by the

two setae near the area of attachment of each egg

sac. Legs of the preadult and adult males: Legs

1, 2 with poorly defined protopods and lacking intercoxal plates, with a setiform unsegmented exopod and weakly sclerotized unsegmented endopod

bearing one hyaline element. Leg 3 a single seta.

Legs 4, 5 absent. Leg 6 represented by a posteroventral flap on the genital segment bearing two

setae and a spinous process.

Other features as in the species below.

Associated with madreporarian corals.

TYPE-SPECIES. — Rhynchomolgus corallophilus

Humes and Ho.

Rhynchomolgus corallophilus Humes and Ho,

1967

FIGURE 190

Rhynchomolgus corallophilus Humes and Ho, 1967b, pp.

15-20, figs. 80-115 [from the gastrovascular cavity of Psammocora contigua (Esper), region of Nosy Be, northwestern

Madagascar].

Previously Known Copepods Here Recorded

from New Hosts

SABEIXIPHILIDAE

Herrmannella columbiae (Thompson, 1897), from the bivalve Schizothaerus (=Tresus) capax Could, Washington.

Herrmannella panopeae (Illg, 1949), from the bivalves

Clinocardium nuttalli (Conrad) and Schizothaerus capax Gould, Washington.

Lichomolgidium sardum Kossmann, 1877, from the ascidian

Halocynthia papillosa (Linnaeus), Banyuls, France.

Lichomolgidium cynthiae (Brian, 1924), from the ascidian

Pyura microcosmus (Savigny), Roscoff, France.

Modiolicola insignis Aurivillius, 1882, from the bivalve

Modiolus barbatus (Linnaeus), Brest, France.

Modiolicola gracilis C. B. Wilson, 1935, from the bivalve

Mytilus californianus Conrad, California.

Scambicornus idoneus (Humes and Cressey, 1961), from

the holothurian Holothuria edulis (Lesson), Madagascar.

Scambicornus tuberatus (Humes and Cressey, 1961), from

the holothurian Bohadschia vitiensis (Semper), Madagascar.

Thamnomolgus robustus Humes, 1969, from the gorgonian

Acanthogorgia aspera Pourtales, Madagascar.

LlCHOMOLClDAF.

Acanthomolgus cuneipes (Humes and Ho, 1968), from the

alcyonacean Dendronrphlhya mucronata (Ptittcr), Madagascar.

Acanthomolgus exilipes (Humes and Ho, 1968), from the

alcyonaceans Dendronephythya speciosa Kiikenthal and

Stereonephthya cordylophora Verscvcldt, Madagascar.

Acanthomolgus fissiseliger (Humes and Ho, 1968), from

the alcyonacean I.emnalia humesi Verseveldt, Madagascar.

Acanthomolgus gentilis (Humes and Ho, 1968), from the

alcyonaceans Urnbellulifera striata (Thompson and Henderson) , Dendronephthya lokobeensis Vcrseveldt, D.

speciosa Kiikenthal, and Slrreonephthya cordylophora

Versevcldt, Madagascar.

Acanthomolgus varirostratus (Humes and Ho, 1968), from

the alcyonaceans Dendronephthya cirsium Kiikenthal,

D. lokobeensis Verseveldt, D. speciosa Kiikenthal, and

Stereonephthya cordylophora Vcrseveldt, Madagascar.

Acanthomolgus verseveldti (Humes and Ho, 1968), from

the alcyonaceans Heteroxenia fuscescens (Hemprich and

Ehrenberg) and Xenia lepida Verseveldt, Madagascar.

Anchimolgus digitatus (Humes and Ho, 1968), from the

madreporarian Favia sp., Madagascar.

Anisomolgus protentus (Humes and Frost, 1964), from the

alcyonaceans Sarcophyton glaucum (Quoy and Gaimard),

and S. globosum Tixier-Durivault, Madagascar.

Ascidioxynus jamaicensis (C. B. Wilson, 1921), from the

ascidian Ascidia atra (Savigny), Curasao.

Doridicola foxi (Gurney, 1927), from the alcyonaceans

Cladiella latissima (Tixier-Durivault) and C. sphaerophora (Hemprich and Ehrenberg), Madagascar.

Gelastomolgus spondyli Humes, 1968, from the madreporarian Plerogyra sp., Madagascar.

Lichomolgus asaphidis Humes, 1959, from the bivalve Solen conneus Lamarck, Madagascar.

Macrochiron sargassi G. O. Sars, 1916, from the ascidians

Didemnum candidum Savigny and Tridemnum lorbiculare (Van Name), Curacao, and the pennatulacean

Renilla reniformis (Pall.), St. Martin, West Indies.

Metaxymolgus aculeatus (Humes and Ho, 1968), from the

alcyonaceans Nephthea amentacea Studer, N. bumasta

Verseveldt, N. filamentosa Verseveldt, N. galbuloides

Verseveldt, N. lanternaria Verseveldt, Stereonephthya

nosybearia Verseveldt, and S. scaphis Verseveldt, Madagascar.
NUMBER 127 345

Metaxymolgus commodus (Humes, 1964), from the nudibranch Hexabranchus sp. (cf. H. marginatus Quoy and

Caimard), Madagascar.

Metaxymolgus httaericus (Humes and Ho, 1968), from

the alcyonacean Cladiella laciniosa (Tixier-Durivault),

Madagascar.

Metaxymolgus patulus (Humes, 1959), from the alcyonacean Sinularia mayi Liittschwager and the crinoid Cenometra emendatrix (Bell), Madagascar.

Metaxymolgus securiger (Humes, 1964), from the nudibranch Asteronotus cespitosus (van Hasselt), Madagascar.

Metaxymolgus spinulifer (Humes and Frost, 1964), from

the alcyonaccans I.emnalia longiramus Versveldt, L. digitata (May), /-. cervicornis (May), L. crassicaulis Verseveldt, /.. tenuis Verseveldt, /.. madagascarensis Verseveldt, Paralemnalia clavata Verseveldt, and Sinularia

polydactyla (Hcmprich and Ehrenberg), Madagascar.

Metaxymolgus venusttis (Humes, 1959), from the crinoid

Cenometra emendatrix (Bell), Madagascar.

Monomolgus unihastatus Humes and Frost, 1964, from the

alcyonacran Parerythmpodium fulvum Forskal, Madagascar.

Octopicola superbus antillensis Stock, Humes, and Gooding, 1963. from the cephalopod Octopus briareus Robson, Florida.

Odontomolgus campulus (Humes and Ho, 1968), from the

madrcporarian Goniopora sp., Madagascar.

Odontomolgus rhadinus (Humes and Ho, 1967), from the

III.KIK |.in.il i.ui Pavona sp.. Madagascar.

Paradoridirola glabripes (Humes and Ho, 1968), from the

alcyonaceans Xetiia macrospiculata Cohar and X. viridis

Schenk, Madagascar.

Paramolgus clai'atus (Humes and Ho, 1968), from the

alcyonaceans Lemnalia longiramus Verseveldt, L. cervicornis (May), and /.. crassicaulis Verseveldt, Madagascar.

Paramolgus spathophorus (Humes and Ho, 1968), from

the alcyonacean Sarcophyton acutangulum (Marenzeller), Madagascar.

Paredromolgus decorus (Humes and Frost, 1964), from the

alcyonaceans Cladiella latissima (Tixier-Durivault), and

C. sphaerophora (Hemprich and Ehrenberg), Madagascar.

Schedomolgus lobophorus (Humes and Ho, 1968), from

the madreporarian Acropora florida (Dana), Eniwetok

Atoll.

Stellicola pollex Humes and Ho, 1967, from the asteroids

Linckia diplax (Miiller and Troschel), L. multiflora

(Lamarck), and L. guildingi Gray, Hawaii.

PSEUDANTHESSHDAE

Kombia angulata Humes, 1962, from the madreporarian

Porites somaliensis Gravier, Mauritius.

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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1960a. Observations on the Orientation and Feeding of the

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1961a. A Polychaet-Infesting Lichomolgid Copepod New

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1961b. A New Lichomolgid Copepod, and the Occurrence

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Crustaceana, 3(2):85-92. figures 1,2.

1962. Egg-Number and Ecology in Commensal and Parasitic Copepods. Annals and Magazine of Natural

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1966. Copepods Associated with Marine Invertebrates

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1887. Sur quelques crustaces parasites des Phallusies.

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1900. Uebersicht der Fauna des Golfes von Triest nebst

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1968. The Biology of Estuarine Animals. 401 pages, 105

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1960. Calanoid and Cyclopoid Copepods Collected from

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1927. Zoological Results of the Cambridge Expedition to

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1944. A New Copepod (Scambicomus hamatus) Parasitic on a Japanese Holothurian. Videnskabelige

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Herbst, H. V.

1955. Cyclopoida Gnathostoma (Crustacea Copepoda)

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Herdman. W. A.

1888. The Foundation and First Season's Work of the

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1894. Seventh Annual Report of the Liverpool Marine

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Ho, J.-S.

1970. Revision of the Genera of the Chondracanthidae,

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Hoffman, H.

1926. Opisthobranchia. In: G. Grimpe and E. Wagler,

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Hoshina, T., and Y. Sugiura

1953. On Two New Species of Parasitic Copepods of

Mollttsks. Journal of the Tokyo University of

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Humes, A. G.

1957. Octopicola superba n. g., n. sp., copepode cyclopoide parasite d'un Octopus de la Mediterranee.

Vie et Milieu, 8(1): 1-8, plates 1, 2, figures 1-22.

1959. Copcpodes parasites dc mollusques a Madagascar.

Me'moires de I'Institut Scientifique de Madagascar,

1958, seiic r, 2:285-542, figures 1-268.

1961. Errata—Copcpodes parasites de mollusques a Madagascar. Me'moires de I'Institut Scientifique de

Madagascar, serie f, 4:2 pages.

1962. Kombia angulata, n. gen., n. sp. (Copepoda, Cyclopoida) Parasitic in a Coral in Madagascar.

Crustaceana, 4(l):47-56, figures 1-38.

1963. Octopicola stocki n. »p. (Copepoda, Cyclopoida)

Associated with an Octopus in Madagascar. Crustaceana, 5(4):27l-280. figures 1-30.

1964. New species of Lichomolgus (Copepoda, Cyclopoida) from Sea Anemone* and Nudibranchs in

Madagascar Cahiers ORSTOM Odanographie

1963, 6(se>ic Nosy Be II) :59-130. figures 1-197.

1966a. New Species of Macrochiron (Copepoda, Cyclopoida) Associated with Hydroids in Madagascar.

lieaufortia, 14 (165):5-28, figures 1-88.

1966b. Pseudanlhessius procurrens n. sp., a Cyclopoid

Copepod Associated with a Cidarid Echinoid in

Madagascar. Breviora, 246:1-14, figures 1-29.

1967a. A New Species of Scambicornus (Copepoda, Cyclopoida, Lichomolgidae) Associated with a Holothurian in Madagascar, with Notes on Several

Previously Described Species. Beaufortia, 14(173):

135-155, figures 1-73.

1967b. Cyclopoid Copepods of the Genus Paranthessius

Associated with Marine Pelecypods in Chile. Proceedings of the United States National Museum,

124 (3628): 1-18, figures 1-43.

1968a. Lecanurius kossmannianus, a New Cyclopoid Copepod Parasitic in Holothurians in Madagascar. Proceedings of the Biological Society of Washington,

81:179-190, figures 1-32.

1968b. Two New Copepods (Cyclopoida, Lichomolgidae)

from Marine Pelecypods in Madagascar. Crustaceana, supplement 1, Studies on Copepoda, pages

65-81, figures 1-64.

1969a. Aspidomolgus stoichactinus n. gen., n. sp. (Copepoda, Cyclopoida) Associated with an Actiniarian

in the West Indies. Crustaceana, 16(3) :225-242,

figures 1-69.

1969b. Cyclopoid Copepods Associated with Antipatharian

Coelenterates in Madagascar. Zoologische Mededelingen, 44(1): 1-30, figures 1-98.

1969c. Pseudanthessius pusillus n. sp., a Cyclopoid Copepod Associated with a Clypeastroid Echinoid in

Madagascar. Zoologischer Anzeiger 183 (3/4) :268-

277, figures 1-30.

1969d. Stellicola dent ifer n. sp. (Copepoda, Cyclopoida)

Associated with a Starfish in Jamaica. Breviora,

315:1-11, figures 1-30.

1969e. A Cyclopoid Copepod, Sewellochiron fidens n. gen.,

n. sp.. Associated with a Medusa in Puerto Rico.

lieaufortia, 16(219) :171-183, figures 1-27.

I969f. Copepods of the Genus Scambicornus (Cyclopoida,

Lichomolgidae) Associated with Holothurians in

the West Indies. Studies on the Fauna of Curacao

and other Caribbean Islands, 29(109):79-95, figures

1-83.

1970a. Cyclopoid Copepods of the Genus Paranthessius

Associated with Marine Pelecypods in the West

Indies. Bulletin of Marine Science, 20 (3) :605-«25,

figures 1-51.

1970b. Stellicola acanthasteris n. sp. (Copepoda, Cyclopoida) Associated with the Starfish Acanthaster

planci (L.) at Eniwetok Atoll. Publications of

the Seto Marine Biological laboratory, 17(5):329-

338, figures 1-30.

1970c. A Census of Copepods Associated with Marine

Invertebrates in a Tropical Locality. Journal of

Parasitology, 56(4) (2) (1): 160-161.

197Od. Paramachochiron japonicum n. sp., a Cyclopoid

Copepod Associated with a Medusa in Japan. Publications of the Seto Marine Biological Laboratory,

18(4):223-232, figures 1-31.

1972. Cyclopoid Copepods Associated with Tridacnidae

(Mollusca, Bivalvia) at Eniwetok Atoll. Proceedings of the Biological Society of Washington,

84 (42) :345-358, figures 1-32.

Humes, A. G., and R. F. Cressey

1958a. Copepod Parasites of Mollusks in West Africa.

Bulletin de I'Institut Francois de VAfrique Noire,

series A, 20(3) :921-942, figures 1-156.

1958b. Four New Species of Lichomolgid Copepods Parasitic on West African Starfishes. Bulletin de I'Institut Francois de VAfrique Noire, series A, 20 (2):

330-341, figures 1-94.

1961a. Copepodes cyclopoides du genre Preherrmannclla

parasites d'holothuries et d'un oursin a Madagascar. Me'moires de I'Institut Scientifique de Madagascar, 1959, series F, 3:25-65, figures 1-157.

1961b. Deux nouvelles especes de Pseudanthessius (Copepoda, Cyclopoida) parasites des oursins a Madagascar. Memoires de I'Institut Scientifique de

Madagascar, 1959, series F, 3:67-82, figures 1-56.

1961c. Lichomolgus oreastriphilus (Kossmann) copepode

cyclopoide parasite des etoiles de mer a Madagascar, Mimoires de I'Institut Scientifique de

Madagascar, 1959, series F, 3:83-92, figures 1-31.
352 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Humes, A. G., and A. De Maria

1969. The Cyclopoid Copepod Genus Macrochiron from

Hydroids in Madagascar. Beaufortia, 16(216): 137-

155, figures 1-52.

Humes, A. G., and B. W. Frost

1964. New Lichomolgid Copepods (Cyclopoida) Associated with Alcyonarians and Madreporarians in

Madagascar. Cahiers ORSTOM Ocianographie,

1963, 6(s*rie Nosy Be II): 131-212, figures 1-236.

Humes, A. G., and J.-S. Ho

1966a. New Lichomolgid Copepods (Cyclopoida) from

Zoanthid Coelenterates in Madagascar. Cahiers

ORSTOM Ocianographie, 4 (2) :S-47, figures 1-143.

1966b. Cyclopoid Copepods Associated with the Starfish

Choriaster granulatus (Liitken) in Madagascar.

Cahiers ORSTOM Oceanographie 4(3):93-108,

figures 1-59.

1967a. New Cyclopoid Copepods Associated with the Alcyonarian Coral Tubipora musica (Linnaeus) in

Madagascar. Proceedings of the United States

National Museum, 121 (3573): 1-24, figures 1-69.

1967b. New Cyclopoid Copepods Associated with the

Coral Psammacora contigua (Espcr) in Madagascar. Proceedings of the United States National

Museum, 122(3586) :l-32, figures 1-115.

1967c. New Cyclopoid Copepods Associated with Polychaete Annelids in Madagascar. Bulletin of the

Museum of Comparative Zoology, Harvard University, 135 (7) :377-413, figures 1-142.

1967d. Two New Species of Lichomolgus (Copepoda, Cyclopoida) from an Actiniarian in Madagascar.

Cahiers ORSTOM Oceanographie, 5(1):S-21, figures 1-53.

1967e. New Species of Stellicola (Copepoda, Cyclopoida)

Associated with Starfishes in Madagascar, with a

Redescription of S. caeruleus (Stebbing, 1900).

Bulletin of the British Museum (Natural History),

Zoology, 15 (5) :201-255, figures 1-141.

1968a. Lichomolgid Copepods (Cyclopoida) Associated

with Corals in Madagascar. Bulletin of the

Museum of Comparative Zoology, Harvard University, 136(10):353-413, figures 1-259.

1968b. Cyclopoid Copepods of the Genus Lichomolgus

Associated with Octocorals of the Family Nephtheidae in Madagascar. Proceedings of the United

States National Museum, 125(3661): 1-41, figures

1-113.

1968c. Cyclopoid Copepods of the Genus Lichomolgus

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Society of Washington, 81:635-692, figures 1-187.

1968d. Cyclopoid Copepods of the Genus Lichomolgus

Associated with Octocorals of the Families Xeniidae,

Nidaliidae, and Telestidae in Madagascar. Proceedings of the Biological Society of Washington,

81:693-750, figures 1-148.

1969a. Cyclopoid Copepods of the Genus Kelleria (Lichomolgidae) from Intertidal Burrows in Madagascar.

Bulletin of the British Museum (Natural History),

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1969b. Cyclopoid Copepods Parasitic in Holothurians in

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894, figures 1-82.

1970a. The Genus Diogenella (Copepoda, Cyclopoida)

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1970b. Cyclopoid Copepods of the Genus Pseudanthessius

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1971. The Genus Diogenidium (Copepoda, Cyclopoida)

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Humes, A. G., and J. H. Stock

1972. Preliminary Notes on a Revision of the Lichomolgidae, Cyclopoid Copepods Mainly Associated

with Marine Invertebrates. Bulletin Zoologisch

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133.

Illg, P. L.

1949. A Review of the Copepod Genus Paranthessius

Claus. Proceedings of the United States National

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1950. A New Copepod, Pseudanthessius latm (Cyclopoida: Lichomolgidae), Commensal with a Marine

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Ulg, P. L., and A. G. Humes

1971. Henicoxiphium redactum, a New Cyclopoid Copepod Associated with an Ascidian in Florida and

North Carolina. Proceedings of the Biological Society of Washington, 83 (49):569-577, figures 1-26.

Jepsen, U.

1965. Die Struktur der Wattenbiozdnosen im Vormundsgebiet der Elbe. Archiv fur Hydrobiologie, Supplement, 29:252-370, figures 1-28, plates 1-6.

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1933. Halbparasitische Copepoden aus der Kieler Bucht.

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K6, Y.

1961. The Biology of the Commensal Copepods in the

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1969a. On the Reproduction and Metamorphosis of a

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1969b. On the Occurrence of Four Commensal Copepods

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K6, Y.; Y. Murakami; and K. Daiku

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1 histogram.

1952. Some New Species of Copepods from Madras Coast.

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336, figures 1-5.

1953. Pelagic Copepoda of the Madras Coast. Journal of

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1949. On Some Swedish Marine Semi-Parasitic and Parasitic Copepods. Arkiv fur Zoologi, 42A (22): 1-10.

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1966. Lc Coral ligenc des Alberes. Monographic biocenotique. Annales de I'lnstitut Ocianographique,

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1933. A Second List of Parasitic Copepoda of Plymouth

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1935. A List of British Invertebrates with Their Characteristic Parasitic and Commensal Copepoda.

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1946. Un nouveau copepode poecilostome de I'Inde de

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1888. Notizie biologiche riguardanti specialmente il periodo di maturita sessuale degli animali del golfo di

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1970. La liste des especes de copepodes trouvees dans les

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1967. A Field and Experimental Study of the Symbiotic

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1949. Nova contribuicSo para o conhedmento dos copepodes da Guine Portuguesa. Anais da Junta de Investigacocs Coloniais, 4 (4): 17-28, plates 1-3.

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1874. Die wirbellosen Thiere der Ostsee. IX. Copepoda

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1961. Les parasites de Microcosmus Heller et les modalites de leur repartition. Vie et Milieu, 12 (1) :97-

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1928. Sur quelques copepodes parasites de nudibranches.

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1932a. Les copepodes parasites de mollusques. Annales

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S54 SMITHSONIAN CONTRIBUTIONS TO ZOOLOCV

1932b. Les copepodcs parasites de mollusques. (Premier

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1934. Des copdpodes parasites de mollusques. (Deuxiemc

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1956. De Wester- en de Oosterscheide, tegenstelling tussen riviermond en binnenzee. Zeepaard, 16(1):

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1944. Littoral Copepods from South Australia (II) Calanoida, Cyclopoida, Notodelphyoida, Monstrilloida

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1903. Notes on the Natural History of East Finmark.

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1905. Crustacea Copepoda New to Science from Devon

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1922. Norwegian Opisthobranchiate Mollusca in the Collections of the Zoological Museum of Kristiana.

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Pearson, J.

1905. A List of the Marine Copepoda of Ireland. Part 1.

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1903. Mollusques (Amphineures, Gastropodes et Lamellibranches). Resuttats du Voyage du S.Y. Belgica,

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1929. Cop^podes parasites de mollusques. Annales de la

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1909. Beitrage zur Kenntnis parasitischer Copepoden.

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Porumb, F., and I. Andriescu

1964. Asupra presenjei a doua copepode in caviatea

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1954. Mollusques opisthobranches. Faune de France,

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Raffaele, F., and F. S. Monticelli

1885. Descrizionc di un nouva Lichomotgus parassita del

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Rancurel, P.

1954. Un nouveau copdpode parasite de taret: Teredophilus renicola. Bulletin de I'Institut Francois

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Reddiah, K.

1960. A New Lichomolgid Copepod, Paranthessius colwant n. sp., from the Laevicardium crassum

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1968a. Three New Species of Paramacrochiron (Lichomolgidae) Associated with Medusae. Crustaceana,

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1868b. Two New Pseudanthessius Species (CopepodaLichomolgidae) from the Madras Harbour. Journal of the Marine Biological Association of India,

1966, 8(2):320-328, figure* 1-4.

1968c. Scambicomus brachysetosus n. sp. (Copepoda: Lichomolgidae) from a Holothurian in the Gulf of

Mannar. Journal of the Marine Biological Association of India, 1967, 9(1) -.126-131, figures 1, 2.

1969. Pseudomacrochiron stocki n. g., n. sp., a Cyclopoid

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Reddiah, K., and D. I. Williamson

1959. On Modiolicola inermis Canu and Modiolicola

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Ricketts, E. F., and ]. Calvin

1952. Between Pacific Tides. 3rd ed.; xii+502 pages. Sunford University Press, Stanford.

Ridley, H. N.

1879. On a New Copepod of the Genus Doridicola. Annals and Magazine of Natural History, series 5,

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Rioja, E.

1935. Sobre un copepodo parasito de Sabella pavonina

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Risbec, J.

1928. Contribution a l'ltude des nudibranches n£ocalddoniens. (These.) Faune des Colonies Francoises, 2:1-328, figures 1-98, plates A-D, 1-8, 3 maps.
NUMBER 127 S55

1930. Nouvelle contribution a l'eiudc des nudibranches

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Rosoll, A.

1889. Uber zwei neue an Echinodermen lebende parasitische Copepoden: Ascomyzon comatulae und

Astericola clausii. Sitiungsberichte der Mathematisch-naturwissenschaftlichen Class der Kaiserlichen

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1951. Inventaire dc la faune marine dc RoscofT. Annelides. Polychctes. Travaux de la Station Biologique de Roscoff, new series. I (supplement 3) :8-57.

Sars. G. O.

1916. Liste systematique des Cyclopides, Harpacticoides

et Monstrilloidcs recueillis pendant let campagnes

de s.A.s. lc Prince Albert de Monaco, avec descriptiom ct figures des especes nouvellcs. Bulletin de

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1917a. An Account of the Crustacea of Norway with

Short Descriptions and Figures of All the Species.

Volume VI, Copepoda, Cyclopoida, Parts XI, XII,

Clausidiidae, Lichomolgidae (part), pages 141-172,

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1917b. Vrocopia singularis G. O. Sara, a Peculiar Semiparasitic Copepod from Great Deeps of the North

Atlantic Ocean. Bergens Museums Aarbok 1916-17,

1 (Naturvidcnskabelig raekke, number 4): I—11. figures 1-15.

1918. An Account of the Crustacea of Norway with Short

Descriptions and Figures of All the Species. Volume

VI, Copepoda, Cyclopoida, Parts XIII, XIV, Lichomolgidae (concluded), Oncaeidae, Corycaeidae,

Ergasilidae, Clausiidae, Eunicicolidae, Supplement,

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1921. An Account of the Crustacea of Norway with Short

Descriptions and Figures of All the Species. Volume VII, Copepoda Supplement, Parts IX, X,

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Sars, M.

1862. Beskrivelse med Afbildninger af fire nye parasitiskc

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Scott, A.

1896a. Description of New and Rare Copepoda. Proceedings and Transactions of the Liverpool Biological

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1896b. Description and Notes of Some New and Rare

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Bay and Neighbouring Seas, Fifth Report, 1900:

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1902. On Some Red Sea and Indian Ocean Copepoda.

Proceedings and Transactions of the Liverpool

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1909. The Copepoda of the Siboga Expedition. Part I.

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Scott, T.

1891. Additions to the Fauna of the Firth of Forth. Part

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1892. Additions to the Fauna of the Firth of Forth.

Part IV. Tenth Annual Report of the Fishery

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plates 7-13.

1893. Additions to the Fauna of the Firth of Forth. Part

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1894a. Report on the Entomostraca from the Gulf of

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1894b. Additions to the Fauna of the Firth of Forth.

Part VI. Twelfth Annual Report of the Fishery

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plates 5-10.

1897a. The Marine Fishes and Invertebrates of Loch Fyne.

Fifteenth Annual Report of the Fishery Board for

Scotland, 1896, part 3. pages 129-158. 168-174

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1897b. Zoological Notes. Sabelliphilus sarsi, Claparede, in

the Clyde. Annals of Scottish Natural History,

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1897c. Zoological Notes. Sabelliphilus sarsi, Claparede,

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1898. Some Additions to the Invertebrate Fauna of Loch

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1900. Notes on Some Gatherings of Crustacea Collected

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1901. Notes on Gatherings of Crustacea, Collected for

the Most Part by the Fishery Steamer "Garland"

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and Examined During the Year 1900. Nineteenth

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1900, part 3, pages 235-281, plates 17, 18.

1902. Notes on Gatherings of Crustacea Collected by the

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Report of the Fishery Board for Scotland, 1901,

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356 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

1903a. On Some New and Rare Crustacea Collected at

Various Times in Connection with the Investigations of the Fishery Board for Scotland. TwentyFirst Annual Report of the Fishery Board for Scotland, 1902, part 3, pages 109-135, plates 2-6.

1903b. Notes on Some Copepoda from the Arctic Seas

Collected in 1890 by the Rev. Canon A.M. Norman, F.R.S. In: A. M. Norman, Notes on the

Natural History of East Finmark. Annals and

Magazine of Natural History, series 7, 11 (61) :4-32,

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1904. Notes on Some Rare and Interesting Marine Crustacea. Twenty-Second Annual Report of the Fishery Board for Scotland, 1903, part 3, pages 242-261,

plates 13-15.

1905. An Account of Some Copepoda That Live as Parasites on, or Messmates with, Other Invertebrata.

Transactions of the Edinburgh Field Naturalists'

and Microscopical Society, 5 (3): 197-207.

1906. A Catalogue of Land, Fresh-Water, and Marine

Crustacea Found in the Basin of the River Forth

and Its Estuary. Proceedings of the Royal Physical

Society, 16(7):267-386.

1907. Observations on Some Copepoda That Live as

Messmates or Commensals with Ascidians. Transactions of the Edinburgh Field Naturalists' and

Microscopical Society, 5 (5): 357-372.

1912. The Entomostraca of the Scottish National Antarctic Expedition, 1902-1904. Transactions of the

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1-14.

Scott, T., and A. Scott

1892a. On Some New or Rare Crustacea from the Firth

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1892b. On New and Rare Crustacea from the East Coast

of Scotland. Annals of Scottish Natural History,

1892 (3):149-156, plates 6, 7.

1893a. On Some New or Rare Crustacea from Scotland.

Annals and Magazine of Natural History, series 6,

12 (70) :237-246, plates 11-13.

1893b. On Some New or Rare Scottish Entomostraca.

Annals and Magazine of Natural History, series 6,

11 (63) :210-215, plates 7, 8.

1894. On Some New and Rare Crustacea from Scotland.

Annals and Magazine of Natural History, series 6,

13 (74) .137-149, plates 8, 9.

1901. On Some Entomostraca Collected in the Arctic Seas

in 1898 by William S. Bruce, F.R.S.G.S. Annals

and Magazine of Natural History, series 7, 3 (47):

337-356, plates 3-6.

Semper, C.

1862. Reisebericht des Herrn Dr. Semper. Fortsetzung.

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Sewell, R. B. S.

1949. The Littoral and Semi-Parasitic Cyclopoida, the

Monstrilloida and Notodelphyoida. John Murray

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199, figures 1-41, 1 map.

Sharpe, R. W.

1910. Notes on the Marine Copepoda and Cladocera of

Woods Hole and Adjacent Regions, Including a

Synopsis of the Genera of the Harpacticoida. Proceedings of the United States National Museum,

38 (1758):405-436, figures 1-20.

Shen, C.J., and F.-S. Lee

1966. On the Estuarine Copepods of Chaikiang River,

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Shih, C.T.; A. J. G. Figueira; and E. H. Grainger

1971. A Synopsis of Canadian Marine /ooplankton. Fisheries Research Board of Canada, Bulletin, 176:1-

264, 1 map.

Sowinsky. W.

1884. K faune rakoobraznikh Cernago Morya. I. O njekotorykh parazitnykh formakh iz gruppy Copepoda.

Memoirs of the Kiev Society of Natural History,

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1904. Introduction a 1'etude de la faunc du bassin matin

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province zoo-geographique indrpcndante. Memoirs

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Stcbbing. T. R. R.

1900. On Crustacea Brought by Dr. Willey from the

South Seas. An: A. WHIey, Zoological Results Based

on Material from New Britain, New Guinea, Loyalty Islands and Elsewhere, Collected During the

Years 1895, 1896 and 1897, part 5, pagei 605-690.

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1910. General Catalogue of South African Crustacea (Part

V of S. A. Crustacea, for the Marine Investigations

in South Africa). Annals of the South African

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Stephensen, K.

1929. Marine Parasitic, Semiparasitic and Non-Pelagic

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Steuer, A.

1940. Uber einige Copepoda Cyclopoida der mediterranen Amphioxussande. Note dcll'Instituto ltaloCermanico di Biologia Marina di Rovigno d'Istria,

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Stock, J. H.

1952. On a Semi-Parasitic Copepod from a Dutch Nudibranch. Basteria, 16(4) :58.

1956. Lichomolgus longicauda (Claus, 1860), Copepod

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1957. Some Notes on the Genus Macrochiron Brady.

1872 (Copepoda, Cyclopoida). Annals and Magazine of Natural History, series 12, 10:378-382, figures 1-10.

1959a. Copepoda Associated with Neapolitan Mollusca.

Pubblicazioni delta Stazione Zoologica di Napoli,

31 (1) :43-58, figures 1-7.

1959b. Copepoda Associated with Neapolitan Invertebrates. Pubblicazioni della Stazione Zoologica di

Napoli, 31 (1) :59-75, figures 1-8.
NUMBER 127 357

1960a. A Note on Some of G. S. Brady's Types of Antarctic Copepoda. Crustaceana, 1 (4) :366-S71, figures

1-4.

1960b. Presence de Lichomolgus canui Sars, copepode cyclopoide confondu avec L. albens Thor. et L. marginatus Thor., sur les cotes neerlandaises. Crustaceana, 1 (1): 72-74.

1960c. Sur quelques copepodes associes aux invertebres des cdtes du Roussillon. Crustaceana, 1 (3):

218-257. figures 1-20.

1962. Lichomolgus pterophilus n. sp., a Cydopoid Copepod Associated with the East Indian Sea-Pen

Pteroeides. Beaufortia, 9(105): 155-163. figures

1-20.

1964a. On Copepoda Associated with Dutch Molluscs.

Basteria, 29(1-4) :65-71.

1964b. On Scatnbicornus Heegaard, 1944, a Senior Synonym of Preherrmannclla Seymour Sewell, 1949

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183-192, figures 1-3.

1964c. Sur deux especes d'Anthessius (Copepoda) des

Indcs Orientates. Zoologische Mededelingen, 39:

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1967a. Copepoda Associated with Invertebrates from the

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1967b. Copepoda Associated with Invertebrates from the

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1968. Copepoda Endoparasitic of Tropical Holothurians.

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1969. The Names of Certain Cyclopoid Copepods Associated with Invertebrates in the Black Sea. Crustaceana, 17 (2)-.220-222.

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1963a. Copepoda Associated with West Indian Invertebrates. IV. The Genera Octopicola, Pseudanthessius and Meomicola (Cyclopoida, Lichomolgidae).

Studies on the Fauna of Curacao and Other Caribbean Islands, 18(77) 1-74, figures 1-24.

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1970. Some Associates of Tresus nuttallii (Conrad, 1837)

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NUMBER 127 359

Wright, R. R.

1885. On a Parasitic Copepod of the Clara. American

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Yamaguti, S.

1936. Parasitic Copepods from Mollusks of Japan, I.

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Yeatman, H. C.

1962. The Problem of Dispersal of Marine Littoral Copepods in the Atlantic Ocean, Including Some Redescriptions of Species. Crustaceana, 4 (4) :253-272,

figures 1-76.
Index to Generic and Specific Names of the Lichomolgoidea

aberdonensis, Lichomolgus, 328

Acaenomolgus, 85

protulae, 86

serpulae, 87

acanthasteris, Synstellicola, 302

acanthodes, Zamolgus, 320

Acanthotnolgus, 87

astrictus, 89

bilobipes, 96

cuneipes, 100

exilipes, 89

fissise tiger, 100

gentilis, 100

hales, 101

hians, 105

longispinifer, 106

plantei, 106

telestophilus, 110

varirostratus, 110

verseveldti, 111

actiniae, Doridicola, 168

actinophorus, Odontomolgus, 254

aculeatus, Metaxymolgus, 222

adduensis, Scambicornus, 71

adelphus, Paradoridicola, 265

adhaerens, Lichomolgus, 328

aegyptius, Lichomolgus, 327

aestheticus, Pseudanthessius, 338

affinis, Synstellicola, 302

agilis, Doridicola, 167

alabatensis, Stellicola, 328

albens, Lichomolgus, 191

Alimeda, 180

Amarda, 111

compta, 116

cultrata, 112

amplectans, Mcomicola, 334

amygdalae, Philoconcha, 279

Anchimolgus, 119

digitatus, 120

prolixipes, 121

andamanensis, Kelleria, 186

Andrianellus, 121

exsertidens, 121

anemoniae, Paranthessius, 66

angoti, Telestadcola, 304

angularis, Pseudanthessius, 338

angulata, Kombia, 332

Anisomolgus, 126

incisus, 127

insolens, 127

protentus, 126

spinipes, 127

anomalus, Paramolgus, 274

Rhinomolgus, 329

anormalus, Pseudanthessius, 338

antheae, Doridicola (=Paranthessius?)

81, 168

Lichomolgus, 81

antheliae, Metaxymolgus, 222

Anthessius, 328

arcanus, Lichomolgus, 191

arcuatipes, Schedomolgus. 290

arenicola, Herrmannella, 328

arenicolus, Lichomolgus, 328

armoricanus, Scambicornus, 71

asaphidis, Lichomolgus, 191

Ascetomolgus, 127

plicatus, 128

Ascidioxynus, 133

bermudcnsis, 138

floridanus, 134

jamaicensis, 142

Aspidomolgus, 146

stoichactinus. 147

assimilis, Pseudanthessius, 338

Astericola, 148

astropectinis, 149

clausi, 148

dentifer, 149

frequens, 149

lautus, 149

luidiae, 149

asterinae, Lichomolgus, 148

Stellicola, 149

astrictus, Acanthomolgus, 89

astropectinis, Astericola, 149

astrophyticus, Doridicola, 168

audens, Doridicola, 174

australiensis, Kelleria, 186

barneae, Herrmannella, 18

bermudensis, Ascidioxynus, 138

bifidus, Modiolicola, 50

bilobipes, Acanthomolgus, 96

bispirae, Sabelliphilus, 82

Bispiraphilus, see Sabelliphilus

Boeckia, 328

Boholia, 327

botulosus, Metaxymolgus, 227

brachysetosus, Scambicornus, 71

brevicauda, Pseudanthessius, 338

Scambicornus, 71

brevicaudis, Lichomolgus, 328

brevifurcatus, Metaxymolgus, 227

brevipes, Doridicola, 174

brevisetosus, Indomolgus, 184

buddhensis, Doridicola, 174

bulbipes, Doridicola, 174

bullata, Herrmannella. 19

caeruleus, Stellicola. 295

Calypsarion, 8

carinatum. 9

leprum, 9

sentosum, 9

Calypsina, 9

changeuxi, 10

camortensis, Kelleria. 186

campanulipes. Scambicornus. 72

campulus, Odontomolgus, 254

canui, Lichomolgus, 192

caribaea, Herrmannella, 24

Caribulus, 10

sculpt us, II

carinatum, Calypurion, 9

cerianthiphila. Boholia, 327

chamanim. Lichomolgui, 192

changeuxi, Calypsina. 10

chcliferum. Macrochiron, 196

chlamydis, Metaxymolgus, 227

Cholidya, 328

cinctus, Metaxymolgus, 232

daudus, Metaxymolgus, 236

clausi, Astericola, 148

clavatus, Paramolgus, 274

clavelinae, Debruma. 161

colmani, Paranthessius, 29

Colobomolgus, 150

cristatus, 150

dentipes, 150

laboutei, 151

columbiae, Herrmannella, 24

commodus, Metaxymolgus, 240

compositus, Spaniomolgus, 293

compta, Amarda, 116

concinna, Herrmannella, 328

concinnus, Lichomolgus, 167

Pseudanthessius, 338

congoensis, Metaxymolgus, 240

conjunctus, Plesiomolgus, 280

constrictus, Paramolgus, 275

Contomolgus, 154

lokobeensis, 155

corallophilus, Rhynchomolgus, 344

crassus, Spaniomolgus, 293

cristatus, Colobomolgus, 150

Nasomolgus, 250

cuanensis, Lichomolgides, 189

360
NUMBER 127 361

cultrata, Amarda, 112

cuneipcs, Acanihomolgus, 100

curticaudatiis. Stcllicola, 295

curtiramus, Zygomolgus, 326

cuspis, Metaxymolgus, 240

cynthiae, Lichomolgidium, 44

Debruma, 161

davelinae, 161

decorus, Parcdromolgus, 276

dcficicns, Pscudanthcssius. 338

deforme, Diogcnidium. 14

deichmannac, Diogcnclla, 13

dcntifer, Astericola. 149

dentipcs, Colobomolgus. 150

dcvotus, Mcringomolgus, 211

diazonac. Lichomolgus, 192

didrmni, Zygomolgus. 326

digilatus, Anchimolgus, 120

dilatatut, Pscudomolgus. 329

Diogenella. 12

tl«- n 1IIII.IIIII.II- . 1 3

mi|I.Ii. IS

M'l K .lllll.l . 1 3

spinicauda, 13

Diogcnidium, 13

dcformc, 14

n.iMitiiin. 14

spinulosum, 14

tectum, 14

dissident, Hcrrmannclla, 29

diversus, Indomolgus, 184

Doridicola, 165

actiniae, 168

agilis, 167

antheac, 81

astrophyticus, 168

audens, 174

brevipes, 174

hiidd lunsis, 174

bulbipes, 174

fishelsoni, 174

foxi, 174

gemmatus, 174

magnificus, 174

ptilosarci, 174

rigidus, 180

trispinosus, 180

dubia, Herrmannella, 81

dubius, Heteranthessius, 330

Pseudanthessius, 338

duplus, Serpuliphilus, 77

echinicolum, Macrochiron, 196

elegans, Lichomoigus, 327

elegantulus, Lichomolgus, 192

elongacus, Sabelliphilus, 68

ennorense, Paramacrochiron, 271

F.olidicola. see Doridicola

Epimolgus, 180; see also Metaxymolgus

orientalis, 180

trochi, 180

Ergasilus, see Doridicola

euiynotus, Temnomolgus, 342

exilipes, Acanthomolgus, 89

exsertidens, Andrianellus, 121

facet us, Mcringomolgus. 205

faouzii, Pseudanthessius, 338

femineus, Stellicola, 296

ferox, Pseudanthessius. 338

fidi-ns, Scwel loch iron, 292

finmarchicus, Scambicornus, 72

firmus, Nasomolgus, 250

fishelsoni. Doridicola, 174

fissisctiger, Acanthomolgus. 100

floridanus, Ascidioxynus, 134

foliatus, Pseudanthessius. 339

fni In uI.i. I.ichomolgus, 192

foxi. Doridicola. 174

frequens, Astcricola, 149

ini iioluiii. Macrochiron, 195

I ' M iiiliirn.i l i< w Inion . 28 2

fucicolus, Lichomolgus, 168

fuliginosus, Lichomolgus, 327

furcalus, Heteranthessius, 331

furcillatus, Lichomoigus, 193

Gclastomolgus, 181

spondyli, 182

geminus, Spaniomolgus, 293

gemmatus, Doridicola, 174

gentilis, Acanthomolgus, 100

gigas, Lichomolgus, 327

glabripes, Paradoridicola, 265

gracilipes, Metaxymolgus, 241

gracilis, Modiolicola, 50

Pseudanthessius, 338

Synstellicola, 303

graciloides, Pseudanthessius, 339

gradata, Kelleria, 186

groenlandicus, Pseudomolgus, 329

gurneyi. Kelleria, 327

hales, Acanthomolgus, 101

hamatus, Meringomolgus, 216

Scambicornus, 71

haploceras, Herrmannella, 29

Haplomolgus, 182

montiporae, 183

Henicoxiphium, 14

redactum, 15

Herrmannella, 15, 328; see also Lichomolgidium, Modiolicola, Scambicornus

barneae, 18

bullata, 19

caribaea, 24

columbiae, 24

dissidens, 29

haploceras, 29

inflatipes, 30

mesodesmatis, 30

panopeae, 30

parva, 33

pecteni, 33

rostrata, 18

saxidomi, 33

tivelae, 37

valida, 39

hetaericus, Metaxymolgus, 241

Heteranthessius, 330

dubius, 330

furcatus, 331

scotti, 331

liians, Acanthomolgus, 105

hirsutipes, Metaxymolgus, 241

holothuriae, Stellicola, 296

hydnophorae, Panjakus, 256

idoneus, Scambicornus, 72

icversi, Lichomolgus, 193

illgi. Stellicola, 296

impar, Diogenella, 13

inaequalis, Metaxymolgus, 241

incamerata, Panaietis, 329

incisus, Anisomolgus, 127

indicus, Lichomolgus, 193

Indomolgus, 183

brevisctosus, 184

di versus, 184

mutatus, 185

panikkari, 185

inermis, Modiolicola, 55

Paranthessius, 56

inflatipes, Herrmannella, 30

inflatiseta, Metaxymolgus, 241

inflatus, Lichomolgus, 193

insectus, Paramolgus, 275

insignis, Lichomolgus, 48

Modiolicola, 48

insolens, Anisomolgus, 127

intcstinalis, Lecanurius, 41

Ischnurella, see Heirmannella

isseli, Lichomolgella, 188

jamaicensis, Ascidioxynus, 142

japonicum, Paramacrochiron, 271

Kelleria, 185, 327; see also Pseudomacrochiron

andamanensis, 186

australiensis, 18fi

camortensis, 186

gradata, 186

pectinata, 186

propinqua, 187
362

SMITHSONIAN CONTRIBUTIONS TO ZOOLOf.Y

purpurocincta, 187

regalis, 186

rubimaculata, 187

Kombia, 331

angulata, 333

kossmanni, Synstellicola, 303

kossmannianus, Lecanurius, 41

laboutci, Colobomolgus, 151

lanceolatus, Prionomolgus, 281

lankensis, Synstellicola, 303

latus, Pseudanthessius, 339

lautus, Astericola, 149

Lecanurius, 39

intestinalis, 41

kossmannianus, 41

leprum, Calypsarion, 9

leptodermatus, Lichomolgus, 193

leptostylis, Pseudomolgus, 329

leptus, Nasomolgus, 251

leuckarti, Sabelliphilus. 82

liber, Pseudanthessius, 339

Lichomolgella, 187

isseli, 188

pusilla, 188

Lichomolgides, 188

cuancnsis, 189

Lichomolgidiutn, 41

cynthiae, 44

sardum, 42

Lichomolgus, 189, 327, 328, 329; see also

Acaenomolgus, Acanthomolgus, Anchimolgus, Anisomolgus, Astericola,

Colobomolgus, Doridicola, Epimolgus, Herrmannella, Macrochiron,

Metaxymolgus, Modiolicola, Odontomolgus, Paradoridicola, Paralichomolgus, Paramacrochiron, Paramolgiis, Paredromolgus, Pennatulicola, Plesiomolgus, Pseudanthessius,

Pseudomacrochiron, Sabelliphilus,

Scambicornus, Schedomolgus, Spaniomolgus, Stellicola, Synstellicola,

Zygomolgus

albens, 191

arcanus, 191

asaphidis, 191

canui, 192

chamarum, 192

diazonae, 192

elegantulus, 192

forficula, 192

furcillatus, 193

ieversi, 193

indicus, 193

inflatus, 193

leptodermatus, 193

longicauda, 193

marginatus, 193

minor, 194

spondyli, 194

tridacnae, 194

Lichothuria, 44

mandibularis, 47

Linckiomolgus, see Paralichomolgus;

Stellicola

littoralis, Lichomolgus, 328

lobatum, Macrochiron, 200

lobophorus, Schedomolgus, 290

lobulatus, Scambicornus, 72

lokobeensis, Contomolgus, 15")

longicauda, Lichomolgus, 193

longicaudatus, Stellicola, 301

longipes, Macrochiron, 327

longiseta. Syntellicola, 303

longispinifer, Acanthomolgus, 106

luculentus, Pseudanthessius, 339

luidiae, Astericola, 149

luteus, Synaptiphilus, 329

lytocarpi, Macrochiron, 200

Macrochiron, 194, 327; see also Anisomolgus, Doridicola, Epimolgus.

Lichomolgus, Metaxymolgus, Paramacrochiron, Pseudomacrochiron,

s..mihi(i)iiiiis. Stellicola

cheliferum, 196

echinicolum, 196

fucicolum, 195

lobatum, 200

lytocarpi, 200

mutatum, 200

rostra turn. 200

sargassi, 201

valgum, 201

vervoorti, 201

madrasensis, Pseudanthessius, 340

magnifkus, Doridicola, 174

major, Lichomolgus, 328

Pseudanthessius, 339

malayense, Pseudomacrochiron, 283

mandibularis, Lichothuria, 47

marginatus, Lichomolgus, 193

maxima, Herrmannella, 56

maximum, Paramacrochiron, 271

maximus, Lichomolgus, 56

Modiolicola, 56

Meomicola, 334

amplectans, 334

meretricis, Paraphiloconcha, 275

Meringomolgus, 204

devotus, 211

facetus, 205

hamatus, 216

mesodesmatis, Herrmannella, 30

Metaxymolgus, 220

aculeatus, 222

antheliae, 222

botulosus, 227

brevifurcatus, 227

chlamydis. 227

cinctus, 232

claudus. 236

commodus, 240

congoensis, 240

cuspis, 240

gracilipes, 241

hetaericus, 241

hirsutipes. 241

inacqualis, 241

inflatiseta, 241

myorae, 246

patulus, 247

securiger, 222

M-nsilis, 247

simplex, 247

singularipes, 247

spinulifcr, 247

venustui, 248

mctisicnsii, Myicola, 329

minimus . I' M iicl.mili . SMUS . 32 8

minor, I.ichnmolguv I'M

I ' M ll'l.llltll . sslll v 33 9

minutui, Psrudanlhnsius, 339

modestllS . Si .imliKiuiiiis , 7 2

Modiolicola, 47 see also Aiddioxynui;

Herrmannella

IMIHIIIS . 5 0

gracilis, 50

mi i mis, 55

insignis, 48

maximus. 56

trabalis. 61

Monomolgus, 248

psammocorae, 250

unihastatus, 249

montiporae, Haplomolgus, 183

mucronatus, Pseudanthessius, 340

mutatum, Macrochiron, 200

mutatus, Indomolgus, 185

Myicola, 329

myxicolae, Myxomolgus. 62

Myxomolgus, 61

myxicolae. 62

proximus, 62

myorae, Metaxymolgus. 246

Nasomolgus, 250

cristatus, 250

firmus, 250

leptus, 251

parvulus, 251

rudis, 251

nasutum, Diogenidium, 14

navanacis, Pseudomolgus, 329

nemertophilus, Pseudanthessius, 340

nicobaricus, Scambicornus, 72
NUMBER 127

notabilis, Pscudanthcssius, 340

obscurus, Pseudanthcssius, 340

Octopicola, 251

stocki, 252

superbus antillensis, 252

superbus superbus, 252

Odontomolgus, 252

actinophorus, 254

campulus, 254

rhadinus, 255

Oncaeola. 328

mini ul.ii is. Paralichomolgus, 328

oreastriphilus. Steilicola, 301

organicus, Plesiomolgus, 280

orirntalis, Epimolgus, 180

oniatum. Pseudomacrochiron, 283

Pachos, 329

Pachysoma, 329

p.ic itu urn. P.u.tm.K MM hit.in. 272

Paclabius. 81

tumidus. 81

Panaietis. 329

panikkari, Indomolgus. 185

Panjakus. 255

hydnophorae. 256

pl.Kv i'\ i:ir. 261

panopcae. Herrmannella, 30

paphiac, Philoconcha. 280

Paradoridicola. 264

adelphus. 265

glabripes, 265

robustus, 265

sinulariae, 266

squamiger, 265

triquetrus, 271

Paralichomolgus, S28; see also Stellicola

Paramacrochiron, 271

ennorense, 271

japonicutn, 271

maximum, 271

pacific urn. 272

rhizostomae, 272

sewelli, 272

Paramolgus, 272

anomalus, 274

clavatus, 274

constrictus, 275

insectus, 275

politus, 274

simulans, 275

spathophorus, 275

Paranthessius, 66; see also Herrmannella, Lichomolgidium, Modiolicola

anemoniae, 66

Parapanaictis, 329

Paraphiloconcha. 275

meretrids, 275

Paredromolgus, 275

decorus, 276

parva, Herrmannella, 33

parvulus, Nasomolgus, 251

parvum, Pseudomacrochiron, 282

parvus, Paranthessius, 33

patulus, Metaxymolgus, 247

pecteni, Herrmannella, 33

Lichomolgus, 33

pectinata, Kelleria, 186

pectinifer, Pseudanthessius, 340

pectinis, Paranthessius, 33

Pseudolichomolgus, 33

Pennatulicola, 276

pteroidis, 278

pterophilus, 278

serratipes, 278

Pestalichomolgus, see Herrmannella

peiiti, Scambicornus, 72

Philoconcha. 278

amygdalae, 279

paphiae. 280

pichoni, Synstellicola, 303

plantei, Acanthomolgus, 106

platygyrae, Panjakus, 261

Plesiomolgus, 280

conjunctus, 280

organicus, 280

pleurobranchi, Steilicola, SOI

plicatus, Ascetomolgus, 128

poculiferus, Scambicornus, 72

politus, Paramolgus, 274

pollex, Stellicola, 301

polypi, Cholidya, 328

pouched, Zygomolgus, 326

prehensilis, Scambicornus, 72

Preherrmannella, see Scambicornus

Prionomolgus, 280

lanceolatus, 281

procurrens, Pseudanthessius, 340

prolixipes, Anchimolgus, 121

propinqua, Kelleria, 187

propinquus, Scambicornus, 72

protentus, Anisomolgus, 126

proteus, Rakotoa, 284

protulae, Acaenomolgus, 86

proximus, Myxomolgus, 62

pruvoti, Strongylopleura, 181

psammocorae, Monomolgus, 250

Pseudanthessius, 328, 334; see also Kelleria, Lichomolgus, Macrochiron,

Pseudomacrochiron

aestheticus, 338

angularis, 338

anormalus, 338

assimilis, 338

brevicauda, 338

concinnus, 338

deficit ns, 338

365

dubius, 338

faouzii, 338

ferox, 338

foliatus, 339

gracilis, 338

graciloides, 339

latus, 339

liber, 339

luculentus, 339

madrasensis, 339

major, 339

minor, 339

minutus, 339

mucronatus, 340

nemertophilus, 340

notabilis, 340

obscurus, 340

pectinifer, 340

procurrens, 340

pusillus, 341

rostellatus, 341

sauvagei, 341

spinifer, 341

tenuis, 341

thorelli, 341

tortuosus, 341

weberi, 341

Pseudolichomolgus, see Herrmannella

Pseudomacrochiron, 281

fucicolum, 282

malayense, 283

ornatum, 283

parvum, 282

stocki, 283

Pseudomolgus, 329

pteroidis, Pennatulicola, 278

pterophilus, Pennatulicola, 278

ptilosarci, Doridicola, 174

punctatum, Pachysoma, 329

purpurocincta, Kelleria, 187

pusilla, Lichomolgella, 188

pusillus, Pseudanthessius, 341

Rakotoa, 283

proteus, 284

Ravahina, 289

tumida, 289

redactum, Henicoxiphium, 15

regalis, Kelleria, 186

rhadinus, Odontomolgus, 255

Rhinomolgus, 329

rhizostomae, Paramacrochiron, 272

Rhynchomolgus, 329

corallophilus, 344

rigidus, Doridicola, 180

robustus, Paradoridicola, 265

Scambicornus, 72

Thamnomolgus, 81

rostellatus, Pseudanthessius, 341
364 SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

rostrata, Herrmannella, 18, 33

rostratum, Macrochiron, 200

rotundus, Lichomolgus, 327

rubimaculata, Kelleria, 187

rudis, Nasomolgus, 251

sabellae, Lichomolgus, 68

Sabelliphilus, 66

bispirae, 82

elongatus, 68

leuckarti, 82

sarsi, 68

sardum, Lichomolgidium, 42

sargassi, Macrochiron, 201

sarsi, Lichomolgus, 68

Sabelliphilus, 68

sauvagei, Pseudanthessius, 341

saxidomi, Herrmannella, 33

Scambicornus, 68

adduensis, 71

armoricanus, 71

brachysetosus, 71

brevicauda, 71

campanulipes, 72

finmarchicus, 72

idoneus, 72

hamatus, 71

lobulatus, 72

modestus, 72

nicobaricus, 72

petiti, 72

poculiferus, 72

prehensilis, 72

propinquus, 72

robustus, 72

serendibicus, 73

subgrandis, 73

subtilis, 73

tenuicaudis, 73

tuberatus, 73

Schedomolgus, 290

arcuatipes, 290

lobophorus, 290

scotti, Heteranthessius, 331

sculptus, Caribulus, 11

securiger, Metaxymolgus, 222

semperi, Stellicola, 328

sensilis, Metaxymolgus, 247

sentosum, Calypsarion, 9

Sepicola, see Lichomolgus

serendibicus, Scambicornus, 73

serpulae, Acaenomolgus, 87

Serpuliphilus, 73

duplus, 77

tenax, 74

serratipes, Pennatulicola, 278

scticauda, Diogenella, 13

sewelli, Paramacrochiron, 272

Sewellochiron, 291

fidens, 292

simplex, Metaxymolgus, 247

simulans, Paramolgus, 275

singularipes, Metaxymolgus, 247

singularis, Urocopia, 329

sinulariac, Paradoridicola, 266

Spaniomolgus, 292

compositus, 293

crassus, 293

geminus, 293

spathophorus. Paramolgus, 275

specialis, Oncaeola, 328

spinicauda, Diogenella, 13

spinifer, Pseudanthessius, 341

spinipes, Anisomolgus, 127

spinosus, Lichomolgus, 329

spinulifer, Metaxymolgus, 247

spinulosum, Diogenidium, 14

spondyli, Gelastomolgus, 182

Lichomolgus, 194

squamiger. Paradoridicola. 265

Stellicola, 293, 328; see also Astcricola.

Lichomolgus, Pennatulicola

caeruleus, 295

curticaudatus, 295

femineus, 296

holothuriae, 296

illgi, 296

longicaudatus, 301

oreastriphilus, 301

pleurobranchi, 301

pollex, 301

thorelli, 295

stocki, Octopicola, 252

Pseudomacrochiron, 283

stoichactinus, Aspidomolgus, 147

Strongylopleura, 181

subgrandis, Scambicornus, 73

subtilis, Scambicornus, 73

superbus antillensis, Octopicola, 252

superbus superbus, Octopicola, 252

Synapticola, 80

teres, 80

Synaptiphilus, 329

Synstellicola, 301

acanthasteris, 302

affinis, 302

gracilis, 303

kossmanni, 303

lankensis, 303

longiseta, 303

pichoni, 303

tectum, Diogenidium, 14

Telestacicola, 304

angoti, 304

telestophilus, Acanthomolgus, 110

Temnomolgus, 341

eurynotus, 342

u-nax, Bispiraphilus, 68

Serpuliphilus, 74

tenuicaudis, Scambicornus, 73

tenuicornis, Lichomolgus, 327

tenuifurcatus, Zygomolgus, 326

tennis, Pseudanthessius, 341

teres, Synapticola, 80

Thamnomolgus, 81

robustus, 81

thorelli, Pseudanthessius. 341

Stellicola, 295

tivelae. Herrmannella, 37

torttioMis, I\< ml.unli c ssiti v 34 1

trabalii. Modiolicola. 61

tridacnae. Lirhomolgus. I'll

tridens, /-amnlgus, 315

triqtietrus. Paradoridirola, 271

trispinoMi*. Doridicola. ISO

trochi. F.piinolgu<i, 180

tuberatus, Scambicornus, 73

tumida, Ravahina, 289

tumidus, Paclabiu*. 81

uniliast.itus. Monomolgin, 249

Urocopia, 329

singularis, 329

vagans, Lichomolgus, 327

valgum, Macrochiron, 201

valida, Herrmannella, 39

varirostratus, Acanthomolgus, 110

varius, Xenomolgus, 309

venustus, Metaxymolgus, 248

verseveldti, Acanthomolgus, 111

vervoorti, Macrochiron, 201

weberi, Pseudanthessius, 341

Xenomolgus, 309

varius, 309

Zamolgus, 314

acanthodes, 320

tridens, 315

Zygomolgus, 325

curtiramus, 326

didemni, 326

poucheti, 326

tenuifurcatus, 326
Index to Host Genera

(Arranged in major taxonomic groups, with associated copepod genera)

CN ID ARIA

Hydrozoa

Aglaophenia (with Macrochirori), 201

Gymnangium (with Macrochiron), 201

Lytocarpui (with Macrochiron), 200, 201

Scyphozoa

Cassiopea (with Sewellochiron), 292

Dactylomelra (with Pseudomacrochiron), 283

l.ychnorhiza (with Paramacrochiron), 272

Rhizostoma (with Paramacrochiron), 272

Thysanostoma (with Paramacrochiron), 271

Anthozoa

Octocorallia

Stolonifera

Tubipora (with Plesiomolgus). 280

Tcle»tacea

Coelogorgia (with Paramo/guj), 274

(with Telestaeieola). 504

Telesto (with Acanthomolgus), 110

Alcyonacea

i4/cyon/um (with Doridicola) , 174

Anthelia (with Metaxymolgus), 222

(with Paradoridicola), 271

Cespitularia (with Zamo/giu), 515

Cladiella (with Doridicola), 174

(with Metaxymolgus) . 241

(with Paredromolgus), 276

Dendronephthya (with Acanthomolgus), 89, 100,

101. HO. HI

(with Contomolgus), 155

Heteroxenia (with Acanthomolgus), 111

Lemnalia (with Acanthomolgus), 100

(with Metaxymolgus), 247, 248

(with Paramolgus), 274

Litophyton (with Metaxymolgus) , 222

Lobophytum (with ^nuomo/giw), 127

Nephthea (with Metaxymolgus), 222

Paralemnalia (with Metaxymolgus), 247, 248

Parerythropodium (with Doridicola), 174

(with Metaxymolgus), 247

(with Monomolgus), 249

Sarcophyton (with /4n«5omo/gu5), 126, 127

(with Paramolgus), 275

Sinularia (with Colobomolgus), 150, 151

(with Meringomolgus), 205, 212, 216

(with Metaxymolgus), 247

(with Paradoridicola), 265, 266

(with Zamolgus), 320

Siphonogorgia (with Acanthomolgus), 105, 106

Stereonephthya (with Acanthomolgus), 89, 100, 101,

111

(with Metaxymolgus), 222

Studeriotes (with Ascetomolgus), 128

(with Contomolgus), 155

Umbellulifera (with Acanthomolgus), 101, 106

Xenia (with Acanthomolgus), HI

(with Paradoridicola), 265

Gorgonacea

Acanthogorgia (with Acanthomolgus), 89

(with Thamnomolgus), 81

Antillogorgia (with Acanthomolgus), 96

Eunicella (with Herrmannella), 35

(with Afr/axymo/guf), 227

Muricella (with Telestaeieola), 304

Paramuricea (with Metaxymolgus), 227

Psammogorgia (with Metaxymolgus), 232

Solenocaulon (with Acanthomolgus), 101

Pennatulacea

Leioptilum (with Doridicola), 174

Pennatula (with Doridicola), 180

Pteroeides (with Pennatulicola), 278

Pteroides (with Pennatulicola), 278

Ptilosarcus (with Doridicola), 174

Renilla (with Macrochirori), 201

Hexacorallia

Actiniana

Actinia (with Doridicola), 168

Anemonia (with Doridicola), 81, 168

(with Paranthessius), 66

Anthea (with Doridicola), 81, 168

Phytocoeteopsis (with Indomolgus), 185

Radianthus (with Metaxymolgus), 240

Rhodactis (with Paramolgus), 274, 275

Stoichactis (with Aspidomolgus), 147

(with Doridicola), 174

(with Metaxymolgus) , 241, 246

Madreporaria

Acropora (with Schedomolgus), 290

(with Spaniomolgus) , 293

Alveopora (with Odontomolgus) , 254

Fawa (with Amarda), 112, 116

(with Anchimotgus), 120

(with Andrianellus), 121

(with Rakotoa), 284

Goniopora (with Anchimolgus), 120

(with Odontomolgus) , 255

Hydnophora (with Panjakus), 256

Montipora (with Haplomolgus), 183

365
366

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Pachyseris (with Prionomolgus), 281

Pavona (with Odontomolgus), 254, 255

Platygyra (with Andrianellus), 121

(with Panjakus), 261

Plerogyra (with Gelastomolgus), 182

Porites (with Anchimolgus), 121

(with Kombia), 333

(with Afonomo/guj), 249

(with Ravahina), 289

(with Xmomo/gitt), 309

Psammocora (with Kombia), 333

(with Monomolgus), 250

(with Odontomolgus), 255

(with Rhynchomolgus), 344

Seriatopora (with Spaniomolgus), 293

Stylophora (with Spaniomolgus), 293

Zoanthidea

Palythoa (with Indomolgus), 184, 185

(with Metaxymolgus), 241

(with Temnomolgus), 342

Antipatharia

Antipathes (with Paramolgus), 275

(with Thamnomolgus), 81

Ceriantharia

Cerianthus (with Boholia), 327

PLATYHELMINTHES

Turbellaria

Cryptophallus (with Pseudanthessius), 339

RHYNCHOCOELA or NEMERTINA

linens (with Pseudanthessius), 340

ANNELIDA

Polychaeta

Bispira (with Sabelliphilus), 82

Eurythoe (with Pseudanthessius), 338

Filograna (with Pseudanthessius), 338

Hermodice (with Pseudanthessius), 338, 341

Myxicola (with Myxomolgus), 62

Polymnia (with Scamfeicornui) , 71

Potynoe (with Doridicola), 168

Pomatoceros (with Pseudanthessius), 338

Pomatostegus (with Serpuliphilus), 77

Protula (with Acaenomolgus), 86

Sabella (with Metaxymolgus), 241

(with Pseudanthessius), 338

(with Sabelliphilus), 68, 82

Sabellasiarte (with Nasomolgus), 250, 251

Serpula (with Acaenomolgus), 87

Spirobranchus (with Serpuliphilus), 74

Spirographis (with Sabelliphilus), 68

MOLLUSCA

Gastropoda*

Prosobranchia

Gibbula (with Epimolgus), 180

Monodonta (with Epimolgus), 180

Trochus (with Epimolgus), 180

Opisthobranchia

Acclesia (with S-trongylopleura), 181

Aeolidia (with Doridicola), 167, 168

(with Lichomolgus), 327

Antiopa (with Doridicola), 167, 168

Antiopella (with Doridicola), 168

Aplysia (with Doridicola), 168

(with Epimolgus), 180

(with Strongylopleura), 181

Archidoris (with Doridicola), 167, 168

/4rminfl (with Doridicola). 167, 168

(with undetermined Lichomolgidae), 328

Asteronotus (with Metaxymolgus), 222

Cavolina (with undetermined Lichomolgidae), 328

Chromodoris (with Doridicola), 167, 168

(with Lichomolgus), 327

(with Metaxymolgus), 241

Dendrodoris (with Doridicola), 168

Doridopsis (with Doridicola), 167

(with Metaxymolgus), 240

Dom (with Doridicola), 167, 168

(with Metaxymolgus), 222

Dorodigitata (with Doridicola), 167

O«f« (with Doridicola), 167, 168

£O/IJ (with Doridicola), 167, 168

Euphurus (with Doridicola), 168

Facelina (with Doridicola), 167. 168

Gasieropteron (with Doridicola), 168

Glossodoris (with Doridieola), 168

(with Afrtaxymo/gui), 241

C.ymnodoris (with Stellicola), S28

(with undetermined Lichomolgidae), 328

Hexabranchus (with Metaxymolgus), 240, 241

Hyalaea (with undetermined Lichomolgidae), 328

/du/ia (with Doridicola), 167, 168

Janolus (with Doridicola), 167, 168

Jorunna (with Doridicola), 168

Limacia (with Doridicola), 168

Noumea (with undetermined Lichomolgidae), 328

Onchidium (with Metaxymolgus), 241

Pa/io (with Doridicola), 168

Phyllidia (with Af«-/axymo/guj), 247, 248

Platydoris (with Doridicola), 174

(with Paralichomolgus), 328

(with undetermined Lichomolgidae), 328

Pleurobranchus (with Stellicola), 301

Pleurophyllidia (with Doridicola), 167

Polycera (with Doridicola), 167, 168

Proctonotus (with Doridicola), 167, 168

Sphaerostoma (with Doridicola), 167, 168

Staurodoris (with Doridicola), 167

Thordisa (with Doridicola), 168

Trevelyana (with Metaxymolgus), 241, 247

(with Stellicola), 328

• Instead of adopting the classification proposed by Taylor

and Sohl (1962), for convenience we have retained an older

classification of the Gastropoda. Taylor and Sohl include

tectibranchs, nudibranchs, and pulmonates (among others)

in the subclass Euthyneura. They place the family Onchidiidae—containing the genus Onchidium and formerly considered to belong to the Pulmonata and more recently to

the Opisthobranchia—in the euthyneuran order Soleolifera.
NUMBER 127 367

(with undetermined Lichomolgidae), 328

Triopa (with Doridicola) , 168

Tritonia (with Doridicola) , 167, 168

Zephyrina (with Doridicola) , 168

Bivalvia

Amygdala (with Philoconcha), 279

Anomia (with Herrmannella) , 33

/Irra (with Lichomolgus), 191

(with Modiolicola), 61

Asaphis (with Lichomolgus) , 191

Barnea (with Herrmannella) , 18

Cardium (with Herrmannella) , 18, 29

(with Lichomolgus), 193

Chama (with Herrmannella) , 24, 29

(with Lichomolgus). 192

Chlamys (with Herrmannella) , 18, 19, 33

(with Aff/axymo/gMj), 227

(with Modiolicola), 55, 56

(with Pseudan(hessius), 341

Clinocardium (with Herrmannella). 30

Emii (with Herrmannella). 18

I.aevicardium (with Herrmannella), 29

(with Lichomolgus), 193

I.utraria (with Herrmannella), 18

Macoma (with Herrmannella), 18

Mactra (with Herrmannella), 18

(with Modiolicola). 50

(with Paranthessius), 81

Meretrix (with Herrmannella), 18

(with Paraphiloconcha) . 275

Mesodesma (with Herrmannella), 30

Modiola (with Modiolicola), 48

Moliolus (with Modiolicola), 48

Mytilus (with Herrmannella), 30

(with Modiolicola), 48, 50

Nacella (with undetermined Lichomolgidae), 328

Ostrea (with Lichomolgus), 191

(with Herrmannella), 19

Panope (with Herrmannella), 30

Paphia (with Herrmannella), 18

(with Lichomolgus), 193

(with Modiolicola), 50

(with Philoconcha), 280

Pecten (with Herrmannella), 18, 33

(with Modiolicola), 55, 56, 57

(with Pseudanthessius), 341

Pholas (with Herrmannella), 18, 19

Pinctada (with Lichomolgus), 193

Pi/ar (with Lichomolgus), 191

Protothaca (with Herrmannella), 24

Pseudochama (with Herrmannella), 24, 29

P/<rria (with Lichomolgus), 192

Saxidomus (with Herrmannella), 33

(with Lichomolgus), 193

Schizothaerus (with Herrmannella), 24, 30

Sofen (with Herrmannella), 18

(with Lichomolgus), 192

Spisula (with Herrmannella), 18

(with Pseudanthessius), 338

Spondylus (with Gelastomolgus), 182

(with Lichomolgus) , 194

Standella (with Lichomolgus), 191

Tapes (with Herrmannella), 18

(with Lichomolgus), 193

(with Modiolicola), 50

(with Paranthessius), 81

Tivela (with Herrmannella), 37

Tresus (with Herrmannella), 24, 30

Tridacna (with Lichomolgus) , 194

(with Paclabius), 81

Venerupis (with Herrmannella) , 18

(with Philoconcha), 279

Cephalopoda

Octopus (with Octopicola), 252

Ommastrephes (with Doridicola) , 168

Srpia (with Lichomolgus), 193

Tritaxeopus (with Octopicola), 252

ARTHROPODA

Crustacea

/fxiuj (with Kelleria), 187

ECHINODERMATA

Crinoidea

Cenometra (with Metaxymolgus), 247, 248

(with Vseudantliessius) , 339

Comaster (with Pseudanthessius), 341

Dichrometra (with Pseudanthessius), 338, 339

Heterometra (with Kelleria), 186

(with Pseudanthessius), 339

Lamprometra (with Pseudanthessius), 339

Liparometra (with Pseudanthessius), 339

Oligometra (with Doridicola), 174

Stephanomelra (with Pseudanthessius), 338, 339

Tropiometra (with Pseudanthessius), 339

Asteroidea

Acanthaster (with Synstellicola), 302

Asteracanthion (with Astericola), 148

Aiterias (with Pseudanthessius), 338, 341

Asterina (with Astericola), 148, 149

Asteropsis (with Stellicola), 301

Astropecten (with Astericola), 148, 149

Choriaster (with Stellicola), 301

(with Synstellicola) , 303

Culcita (with Stellicola). 301

Leiaster (with Stellicola), 2%

Linckia (with Stellicola), 295, 296, 301

Luidia (with Astericola) 149

(with Synstellicola), 302

Maculaster (with Synstellicola), 302

Marthasterias (with Astericola), 148, 149

(with Pseudanthessius), 339

Mithrodia (with Synstellicola), 303

Ophidiaster (with Stellicola), 295, 328

Pentaceraster (with Stellicola), 301

Pentaceros (with Lichomolgus), 327

Poraster (with Stellicola), 301

Protoreas-ter (with Stellicola), 301

(with Synstellicola), 303

Retaster (with Synstellicola), 302
368

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY

Ophiuroidea

Astrophyton (with Doridicola) , 168

Euryale (with Metaxymolgus), 236

Ophioderma (with Pseudanthessius), 338

Echinoidea

Clypeaster (with Pseudanthessius), 340

Diadema (with Scambicornus), 72

Echinocardium (with Pseudanthessius), 341

Echinodiscus (with Pseudanthessius), 341

Echinometra (with Macrochiron), 196

(with Pseudanthessius), 340

Echinothrix (with Pseudanthessius), 339

Echinus (with Pseudanthessius), 338, 339, 341

Lytechinus (with Macrochiron) , 196

Meoma (with Meomicola), 334

Paracentrotus (with Pseudanthessius), 339

Phyllacanthus (with Pseudanthessius), 340

Psammechinus (with Lichomolgus), 193

(with Pseudanthessius), 193, 339

Spatangus (with Pseudanthessius), 341

Stomopneustes (with Pseudanthessius), 339

Strongylocentrotus (with Pseudanthessius), 339

Tripneustes (with Macrochiron), 196

Holothuroidea

Actinopyga (with Calypsarion), 9

(with Caribulus), 12

(with Diogenella), 13

(with Diogenidium), 14

(with Lecanurius), 41

(with Scambicornus), 72

Argiodia, see Microthele

Bohadschia (with Calypsarion), 9

(with Scambicornus), 72, 73

Brandtothuria (with Diogenella), 13

(with Diogenidium), 14

(with Scambicornus), 72

Halodeima (with Diogenella), 13

(with Lichothuria), 47

(with Scambicornus), 72, 73

Holothuria (with Calypsina), 10

(with Diogenella), 13

(with Diogenidium), 14

(with Lichothuria), 47

(with Scambicornus), 71, 72, 73

Isostichopus (with Caribulus), 11

(with Diogenidium), 14

Lapidoplax (with Zygomolgus), 326

Ludwigothuria (with Caribulus), 11, 12

(with Diogenella), 13

(with Diogenidium), 14

(with Lichothuria), 47

(with Pseudanthessius), 338

(with Scambicornus), 72

Microthele (with Lichothuria), 47

(with Scambicornus), 72

# U. S. GOVERNMENT PRINTING OFFICE: I»73 516-215/2

Muelleria (with Diogenidium), 14

(with Lecanurius), 41

Opheodesoma (with Scambicornus) , 73

Semperothuria (with Diogenella), 13

Stichopus (with Calypsarion), 9

(with Scambicornus) , 72

Synapta (with Lecanurius), 41

(with Scambicornus) , 72

(with Synapticola), 80

Thelenota (with Scambicornus), 73

Thyonidium (with Scambicornus), 71

Urodernas (with Scambicornus), 72

CHORDATA

Ascidiacea

Ascidia (with ^jf/dioxynui), 142

(with Herrmannella) , 18

(with Lichomolgus), 191. 192, 193

Ascidiella (with Lichomolgus), 191. 192

Ciona (with Lichomolgus), 191, 192. 193

Clavelina (with Debruma), 161

(with Lichomolgus), 192

Coretla (with Lichomolgus), 191. 193

Cynthia (with Lichomolgidium). 42. 44

(with Lichomolgus). 191

Dendrodoa (with Lichomolgus), 192

Diazona (with Lichomolgus), 192

Didemnum (with Zygomolgus), 326

(with Macrochiron), 201

Diplosoma (with Zygomolgus), 326

Ecteinascidia (with .-Jw IV//«\VHIM) , 138

Fragarium (with Zygomolgus), 326

Halocynthia (with Lichomolgidium), 42. 43

Microcosmus (with Heteranthessius), 531

(with Lichomolgus), 327

Molgula (with Lichomolgus), 191. 192

Morchellium (with Zygomolgus), 326

Perophora (with Lichomolgus), 191

Phallusia (with Lichomolgus), 191, 192, 193

Pyura (with Lichomolgidium), 42, 43, 44

Sidnyum (with Zygomolgus), 326

.S7)>e/<j (with Henicoxiphium), 15

(with Lichomolgidium), 44

(with Lichomolgus), 192, 193

Styelopsis (with Lichomolgus), 192

Trididemnum (with Lichomolgides), 189

(with Macrochiron), 201

ALGAE

Fwcus (with Macrochiron), 195

Lithophyllum (with Heteranthessius), 331

(with Zygomolgus), 326

Rhodymenia (with Pseudanthessius), 338

Sargassum (with Macrochiron), 195, 201

TRACHEOPHYTA

Posidonia (with Lichomolgella), 188

Zoitera (with Metaxymolgus), 241


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