ISBN 2-85653-106-7
MÉMOIRES
DU
MUSÉUM NATIONAL
D’HISTOIRE NATURELLE
NOUVELLE SÉRIE
Série A, Zoologie
TOME 117
Paul L. ILLG and Patricia L. DUDLEY
THE FAMILY ASCIDICOLIDAE AND ITS SUBFAMILIES
(COPEPODA, CYCLOPOIDA), WITH DESCRIPTIONS
OF NEW SPECIES
PARIS
EDITIONS DU MUSÉUM
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Source : MNHN, Paris
Source ; MNHN, Paris
MÉMOIRES DU MUSÉUM NATIOXAL D’HISTOIRE \ATURELLE
Série A, Zoologie, Tome 117
THE FAMILY ASCIDICOLIDAE AND ITS SUBFAMILIES
(COPEPODA, CYCLOPOIDA), WITH DESCRIPTIONS
OF NEW SPECIES
by
Paul L lllg * and Patricia L Dudley **
SUMMARY
I. — Introduction. 5
Acknowledgments. 8
Matériel and methods. 8
II. — Family Ascidicolidae Thorell, 1859 . 9
Key to the subfamilies. 15
Subfamily Ascidicolinae Thorell, 1859. 16
Key to the généra. 17
Ascidicola Thorell, 1859. 17
Ascidicola ro»ea Thorell, 1859. 17
Styelicola Lûtzen, 1968. 24
Key to species. 25
Styelicola bakusia Lützen, 1968. 25
Styelicola lighti new species. 25
Subfamily Buprorinae Thorell, 1859. 30
Buprorus Thorell, 1859. 31
Key to females of Buprorus . 32
Buprorus Icveni Thorell, 1859. 32
Buprorus rwrdgaardi Sars, 1921. 32
Buprorus caudalm new species. 32
Subfamily Enterocolinae Délia Valle, 1883 . 35
Key to généra. 36
* Laboratoire d'Ëcologie générale et Laboratoire de Biologie des Invertébrée marine, Muséum national d'Hie-
toire naturelle, 57, rue Cuvier, 75231 Paris Cedex 05, France - Zoology Department, University of Washington, Seattle,
Washington 98195, U.S.A.
*• Biology Department, Barnard College, Columbia University, New-York, N.Y. 10027, U.S.A.
Source : MNHN, Paris
2
SUMMARY
Enterocoiinae sp. incertae sedis . 37
Lequerrea Chatton & Harant, 1924. 37
Lequérrea perezi Chatton & Harant, 1924. 37
Lequerrea canui new species. 39
Enlerocolides Chatton & Harant, 1922. 42
Enlerocolides ecaudalus Chatton & Harant, 1922. 43
Enterocola van Beneden, 1860 . 45
Key to species. ^7
non Enterocola sp. 48
Species of Enterocola . 49
Enterocola bilamellata Sars, 1921. 49
Enterocola hessi Chatton & Harant, 1924. 49
Enterocola mammifera Chatton & Harant, 1922. 49
Enterocola megalooa Gotto, 1962. 49
Enterocola setifera Hansen, 1923. 49
Enterocola sp. Claus, 1875. 50
Enterocola sp. A Chatton & Brément, 1909 . 50
Enterocola sp. B Chatton & Brément, 1909 . 50
Enterocola sp. Scheilenberg, 1922 . 50
Enterocola pterophora Chatton & Brément, 1909. 50
Enterocola fulgens van Beneden, 1860 . 63
Enterocola claoelinae Chatton & Harant, 1924. 68
Enterocola precaria new species. 72
Enterocola ianthina new species. 74
Enterocola sydnii Chatton & Harant, 1924. 76
Enterocola petiti Guille, 1964. 80
Enterocola brementi new species. 82
Enterocola fertilis new species. 85
Enterocola laiiceps new species. 91
Indéterminable species. 94
Subfamily Enteropsinae Aurivillius, 1885. 94
Key to females of généra of Enteropsinae. 97
Mychopkilus Hesse, 1865. 97
Key to species. 98
Mychophilus roseus Hesse, 1865 . 98
Mychopkilus fallax Stock, 1967. 103
Enleropsis Aurivillius, 1885. 103
Enleropsis fifth male copepodid. 105
Key to the species. 106
Enteropsis chattoni Monniot, 1961. 106
EnteropsU superbus new species. 108
Enteropsis minor new species. 111
Enteropsis roscoffensis Chatton & Brément, 1909. 114
Eiüeropsis sphinx Aurivillius, 1887. 120
Enteropsis capitulatus new species. 123
Enleropsis georgianus Schellenberg, 1922. 126
Enteropsis abbotti new species. 128
ErUeropste orychopkorua Schellenberg, 1922. 131
Indéterminable species. 131
Excluded species. 132
Subfamily Haplostominae Chatton & Harant, 1924. 132
Key to the généra. 133
Haplosloma (Canu, 1886}. 133
Key to the species. 133
Haplostomides Chatton & Harant, 1924. 134
Key to the species. 134
Haplosaccm Chatton & Harant, 1924. 135
Source : MIJHN, Paris
SUMMARV
3
Key to the speoies.
Haploslomella Chatton & Harant, 1924.
Key to the species.
Subfamiiy Botryllophilinae nomen conservandum .
Key to généra.
Botryllopkilus Hesse, 1864.
Species of Botryllopkilus .
Botryllopkilus ruber Hesse, 1864.
Botryllopkilus macropus Garni, 1891.
Botryllopkilus banyulsnsis Brément, 1909. . .
Botryllopkilus bredpes Brément, 1909.
Botryllopkilus bergemis Schellenberg, 1921.,
Botryllopkilus noruegicus Schellenberg, 1921.
Botryllopkilus hrevipes Sars, 1921.
Botryllopkilus aspinosus Schellenberg, 1922.
Botryllopkilus inasquipes Hansen, 1923....
Botryllopkilus randalli Stock, 1970.
Indéterminable species.
Sckizoproctus Aurivillius, 1885.
Key to the species.
Sckizoproctus inflatus Aurivillius, 1885....
Sckizoproctus oestilus (Sars, 1921).
Indéterminable genus, Botryllophilinae.
Enterognathinae, New Subfamiiy.
Key to généra.
Enterognathus Giesbrecht, 1900.
Enterognatkus comaiulae Giesbrecht, 1900..
Enterognatkus laleripes Stock, 1966.
Zanclopus Caïman, 1908.
Zanclopus cepkalodisci Caïman, 1908.
Zanclopus anlarcticus Gravier, 1912.
III.— Development.
Development of Ascidicola rossa .
Development of ErUerocola fertilis .
Development of ErUerocola laticeps .
Third Copepodid of Enlerocola laticeps .
Developmental stages of Enteropsis capüulatus.
Bibliography.
135
135
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141
143
145
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145
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Source : MNHN, Paris
IIÊSUMÉ-ABSTRACT
ABSTRACT
A review of the family Ascidicolidae Thorell is presented, with reviews of the included
subfamilies Ascidicolinae, Buprorinae, Enterocolinae, Enteropsinae and the new subfamily Entero*
gnathinae. A synoptic présentation is ofTered of the subfamilies Botryllophilinae and Haplos
tominae. New species are described in the généra Styelicola Lûtzen, Buprorus Thorell, EiUerocola
van Beneden, Lequerrea Chatton & Harant and Enteropsis Aurivillius. Anatomical details of deve-
lopmental stages in selected species in Ascidicola, Enterocola and ErUeropsU are figured and descri*
bed in support of our morphological interprétations.
RÉSUMÉ
Une révision de la famille des Ascidicolidae est présentée ici avec une revue des sous-familles qui en font
partie : Ascidicolinae, Buprorinae, Enterocolinae, Enteropsinae et la nouvelle sous-famille des Ente-
rognathinae. Les familles des Botryllophilinae et Haplostominae sont présentées de façon synoptique. De
nouvelles espèces sont décrites dans les genres Styelicola Lützen, Buprorus Thorell, Enterocola Van Beneden
et EnieropsU Aurivillius. Des détails anatomiques de différents stades de développement chez certaines espèces
de Ascidicola, Enterocola et Enteropsis sont décrits et figurés et viennent renforcer nos interprétations morpho¬
logiques.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAHILIES
I. — INTRODUCTION
For many years we hâve pursued a joint study o( copepods living in association with ascidians.
The interest goes back to an early phase of study of copepods from the West Coast of North America
by the senior author. Early in the research it was found necessary to restudy some of the forras des-
cribed in the classic literature and a great quantity of material became availakle to us through a period
of time spent by the junior author in laboratories on the Mediterranean, particularly at the Labora¬
toire Arago, Banyuls-sur-Mer. In addition, the unusual opportunities afforded to us at the Friday
Harbor Laboratories hâve made it possible for us to develop an acquaintance with hosts and their
associâtes in the living State to a degree which has not ordinarily been the lot of previous students of
the group.
It is noteworthy that a great variety of organisnis live internally with ascidians in varying
degrees of relationship. Of the copepods there are members of several distinct lineages that furnish
examples. A few members of the Harpacticoida live in definite association with certain ascidians.
Most of the associated copepods are assigned to the broad group formerly referred to the Cyclopoida.
Of these a few species, and these very probably not in a definite and obiigate association, are referrable
to the cyclopoid siphonostomes. A few more species, now distributed in a few généra, are members
of the poecilostomes. There are particularly numerous and long-standing référencés to members
of the genus Lichomolgus. The greatest number of species, and these in a sériés of généra showiiig
a very wide range of adaptational divergences, are assigned to the Cyclopoida Gnathostoma. These
include the many species of the family Notodelphyidae, in the sense of Illc, 1958 ; ail but one of the
few species so far known in the family Archinotodelphyidae ; and the species of the assemblage to be
discussed beiow, the Ascidicolidae of the présent treatment. Ail the copepods occurring in associa¬
tion with ascidians, and indeed any organism so existing, may properly be termed ascidicoles. The
term ascidicolids has historically reccived a very wide application, as we demonstrate below, and in the
treatment of several authors has included the Notodelphyidae as well as the diverse assemblage to
which we now propose to restrict the concept. These gnathostome ascidicoles, including the three
families mentioned above emphatically demonstrate a very important biological considération. In
every représentative for which we hâve been able to find information the development is based on a
lecithotrophic pattern, .\lthough there are free-living developmental stages they are non-feeding
and so there is throughout a trophic dependence, in every known species, on the symbiotic existence
with an ascidian host. There is a graduation in the pattern expressed in this trophic dependence
although exact information on feeding mechanisms and the précisé formulation of the dietary compo-
nents are very difficult to ascertain and rarely recorded. In the many species, probably the majority,
living at the level of a rather simple commensalism, the problem is perhaps most difficult. In the cases
of some of the actual parasites, which seem to be of rare occurrence, there are available some detailed
findijigs (Dudley, 1968, for the remarkable notodelphyid species Scolecodes hurUsinani, for example).
The great number of ascidicoles belonging to généra referrable to the family Notodelphyidae
probably influenced Sars (1901-1903 ; 1921) when he proposed an ordinal classification for the copepods,
to erect a suborder Notodelphyoida, including essentially ail the known gnathestome ascidicolc généra
of that time. Wilson (1932) adopted Sars’ taxon and expanded it by including a sériés of strongly
anatomically modified symbionts of various invertebrates. Most of these généra hâve since been allo-
cated to families substantially removed from the ascidicole complex, falling for the most part among
the poecilostomes. Lang (1948) arrived at the important conclusion that the order Notodelphyoida
had become obsolète. We concur and do not recognize the order as valid in the overall classification
Source : MNHN, Paris
6
PAUL L. ILLG AND PATRICIA L. DUDLEY
of tlie copepods. Lanc’s further suggestion thaï a considérable sériés of ascidicole forms, the “ Eiite-
rocolidae and its relatives ” are directly dérivative from the poecilostomes as well, furnished in part the mo¬
tivation leading us to study this group extensively, and particularly ihrough the compilation of représen¬
tative life historiés. The information we hâve developed leads us to concludo that the gnathostome
ascidicole sériés is for the most part a naturel assemblage deriving from free-living ancestors among
the Cyclopoida Gnathostoma. We thus revise Lang’s suggested outline (1948, p. 27) which proposed
a classification for the Cyclopoida Gnathostoma and Cyclopoida Poecilostoma. W'e alter the ranks
of sonie of his categories and remove his Enterocolidiformes from the poecilostomes to restore it to the
gnathostomes. W’e arrive then at the families Archinotodelphyidae, Notodelphyidae and Ascidicolidae,
the latter subdivided into subfamilies Ascidicolinae, Uuprorinae, Enterocolinae, Enteropsinae, Botryllo-
philinae and Haplostominae, with a new subfamily, Enterognathinae, for related forms which arc not
ascidicole but show many indications of relationship.
W'e hâve given much considération to the matter of Buprorus and to Enlerognatkus and Zanclopus.
Few specimens hâve ever been studied of these rare organisms and we hâve very little material. It
seems clearly indicated that the only logical position for them in the overall system of the copepods
must be in close relation to our group. It has seemed inconsistent to maintain a monotpyic family
for Bupwriis, althoiigh there is a long-standing tradition for such treatment. In the case of Buprorus
ail of the information so far available concerns the anatomy of the adult female, in which the appendages
are considerably reduced. Study of the male and of the developmental stages has not yet been possible
and reinains essentiel for final resolution of the position of this form. Our conclusions dérivé very
substantially from our familiarity with the remarkable séries of modifications within the family Noto-
delphyidae. W'ith these Buprorus shares the incubatory cavity in the adult female. However the
most modiiied members among the notodelphyids hâve very reduced and altered appendages, these
modifications falling into a very extensive sériés as now known, and there is no situation in which the
pattern of modification conforms to the condition in Buprorus. The appendages here hâve struck us
as basically ascidicolid, as we will attempt to demonstrate in discussion below. This leads us to deal
with the question of the incubatory cavity in Buprorus, an extremely important diagnostic feature
of ail notodelphyids, by postulating a convergence. Since we are convinced that the notodelphyid
and ascidicolid stocks are genetically fairly close in the first place, and since much more remotely
counected copepods also hâve incubatory cavities, we adopt the convergence explanation as our work-
iiig hypothesis.
A feature in certain of the lineages that make up the Ascidicolidae in our concept is great taxono¬
mie rcfractoriness in certain généra. The genus Bolryllophilus has been widely recorded as ofPering
ppculiarly great difficulty to the systematist and our own direct expérience amply confirms this idea.
A very similar case is involved when an attempt is made to deal with the taxonomy of the European
species of Enlerocola. The particularly accompllshed w'orkers Canu and Chatton and his colleagues
hâve already recorded or intimated their problems with the best-known species E. fulgens (and its
forma E. belencourli). Every collection, or even, every new specimen, has shown a tendency to con¬
fuse the concept based on pre-existing studies.
.\8 we will discuss below in connection with the Botryllophilinae we are able to prosent only
a sketch of a classification for this subfamily and must leave unresolved matters of synonymy and indeed
actiial identity of some of the long recorded species.
One of us, with a collaborator (Ooishi & Illg, 1977) hâve dealt with the basic taxonomy of
the subfamily Haplostominae, so only a résume of our findings is included in the présent paper.
The name Ascidicolidae stems from the work of Thobell (1859) ; in his treatment, so remarkable
for its day in its anatomical compétence, the author restricted the taxon to the genus Ascidicola, as the
subfamily Ascidicolidae in the family Notodelphyidae. He proposed the monotypic family Buproridae
in the same work. The tradition of pioneering anatomical discrimination of the ascidicoles was con-
tinued for the Scandinavian and Arctic faunas in the work of Aurivillius and culminated in the mono-
graph of G. O. Sars, (1921). In this approach the descriptionist attitude led to the proposai of many
higher level taxa, many of which can be reduced in the general treatment of the présent study. Subs-
tantial leadership in the modem Ireiid of more synthetic treatment of the ascidicoles has corne from
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
7
the work o( Lang {1948, 1949). It is remarkable that throughout the taxonomie history of the ascidi-
colids there has persisted a great emphasis on the ecological characteristics of association with ascidians.
In the periods of synthetic treatment of the group, this criterion has continued to emerge. Even in
the strongly comparative morphological and evolutionary outlook of E. Canu (1886, 1892, etc.), there
is great persistcnce of this viewpoint. His family définition was one of the widest in scope of any
and was directly influenced by this considération. In his monograph (1892, p. 107), he States, in dis-
cussing Ascidicolidae : « Dans cette famille, j’ai réuni toutes les espèces commensales et parasites des
Tuniciers, qui formaient autrefois les diverses familles des Notodelphidés Tborell, des Buproridés
Thorell, des Kosmechtridés Della Valle équivalente aux Schizoproctidés Aurivillius, des Ente*
rocolidés Della Valle, des Enteropsidés Aurivillius... Les Ascidicolidés, qui étaient à l’origine
de parfaits gnathostomes nageurs, abrités seulement dans la liranchie de leur hôte {Notodelphys), se
sont peu à peu adaptés à l’existence cavicole en modifiant leur mode de locomotion dans différents
sens (Doropygus, Bonnierilla, Guenenolophorus, d’une part ; Ascidicola, d’autre part). Changeant leur
genre de vie et devenant de vrais parasites, ils se sont transformés en siphonostomes, par les divers
stades ; Agnathancr, Enterocola, Enteropsis, Aplosloma. »
It is only fair to emphasize that it was actually by comparative anatomical characters that
C.vNU diagnosed the varions categories within his group, and further, he was a most successful innovator
in the cxtremely important approach of study of comparative life historiés as a taxonomie supplément.
In reaching a concept of the family he was much influenced by the strong evolutionary implications
and (p. 186) States, in connection with his definitive diagnosis of the family : «... Mais c’est bien pour les
Ascidicolidae que semble écrite la phrase de Lamarck choisie comme épigraphe de ce travail (« Il n’y a
que ceux qui se sont longtemps et fortement occupés de la détermination des espèces, et qui ont consulté
de riches collections, qui peuvent savoir jusqu’à quel point les espèces, parmi les corps vivants, se fondent
les unes dans les autres, et qui ont pu se convaincre que dans les parties où nous voyons des espèces
isolées, cela n’est ainsi que parce qu’il nous en manque d’autres qui en sont plus voisines et que nous
n’avons pas encore recueillies. » Lamarck, Philosophie zoologique, Première partie, chapitre IlL) ;
il n’est point, dans cette famille, d’espèces isolées : toutes s’enchaînent et se relient ; mes prédécesseurs
ont pensé différemment parce qu’ils ignoraient encore les espèces établissant les relations les plus impor¬
tantes, les types de passage sans lesquels ne peut être comprise la biologie de ces êtres.
Malgré la diversité des types qui fa composent, je crois inutile de subdiviser actuellement cette
faible série de Copépodes, où les délimitations génériques suffisent à tous les besoins de la systématique. »
\Ve acknowledge as Canu’s great contribution to the study of these forms his insistence on the
evolutionary significance of the inany degrees of adaptation to symbiotic existence and on the equal
importance of study of development as an approach in analysis of the morphological complexities
resulting.
The excellent start produced by Canu in analysis of the strangely modified ascidicolous forms
was subsequently continued with great brilliancc by Professer Édouard Chatton and two successive
associâtes, M. Ernest Bbément and, later. Professer Hervé Harant. It is most unfortunate that
this program did not corne to a climactic synthetic treatment, because the descriptions of new forms and
the beginiiings of formulation of concepts as to higher categories are at an absolutely first rank scientific
level. The most notable contributions of this group are the extensions of knowledge of the subfamilies
Enterocolinae and Haplostominae, both of which were proposed as formai taxa after a long period of
usage informally — “ enterocoliens ” and “ aplostomiens, ” after terms set forth by Canu (1886a,
p. 373-374). The formai diagnoses of Enterocolinae and Haplostominae (Chatton & Harant, 1924 b,
p. 360-363 ; 1924e, 416-416) included the explicit statement that latter subfamily was placed in the
family Ascidicolidae. Various more recent authors hâve apparently held a concept that included these
taxa within a family Enterocolidae, although it is the usage that has furnished this indication, not
explicit statements. The formulations of Chatton & Harant helped to lead us to our conclusion that
a réconciliation of the many related but diverse gnathostomes in two major families, Notodelphyidae
and Ascidicolidae is possible. It was the inspiration of the work of M. Chatton and his colleagues and
the opportunity to study his specimens and to collect more material from his localities which led to the
présent paper.
Source : MNHN, Paris
g
PAUL L. ILLG AND PATRICIA L. DUDLEY
ACKNOWLEDGMENTS
We wish to express our appréciation and hearty thanks to the institutions and persons mentioned
in our account of the Matériel and Methods. Many others hâve generously contributed information
and specimens and shared their time in some manner in connection with our studies. We must acknow-
ledge the continuing support of the Department of Zoology, University of Washington ; the number
of helpful contributions from our colleagues and friends of the faculty and among the students, past
and présent, hâve been too numerous to Hst individually. The Department of Biology, Barnard College,
Columbia University was a similar source of encouragement. We owe thanks in a variety of connections
to : Dr. Takasi Tokioka, Seto Marine Biological Laboratory ; M''® Françoise Lafargue, Laboratoire
Arago ; Professer Hervé Harant, Université de Montpellier ; Dr. D. L. Abbott, Hopkins Marine
Station ; Dr. Shigeko Ooishi, University of Mie ; Department of Living Invertebrates, American Muséum
of Natural History ; Station Biologique, Roscoff ; Dr. J. F. L. Hart (Mrs. G. C. Carl) ; Mr. David
Montgomery.
The base of our long-continuing studies was furnished by the direct collection of specimens and
the obtaining of life history material by rearing procedures by Dudley at the Laboratoire Arago.
By the courtesy of M. C. Delamare Deboutteville and M“® Édouard Chatton there became avai-
labié to us at the same time the suite of specimens left by Professer Chatton, including many of this
type specimens. The study of Dr. Dudley was supported by a grant of the National Science Founda¬
tion, GB-6474X2, to the senior author. Subséquent grants provided him the opportunity of several
years of coUecting, anatomical studies and the drawing and drafting of the preliminary présentation
of our ôndings. Much of this work was done at the Friday Harbor Laboratories, University of Was¬
hington, then under the direction of Professer R. L. Fernald.
Some specimens were collected at the Stazione Zoologica, Naples and at the Marine Biological
Laboratory, Plymouth by Dr. Dudley. We were able to examine some type specimens and other
materials during brief visits at the British Muséum (Natural History) and at the National Muséum of
Natural History (formerly United States National Muséum). Many specimens and much advicc on
the taxonomy of hosts and on symbiotic interactions of copepods and ascidians were furnished by
M. C. and M®* F. Monniot. Opportunity for the senior author to compile the final draft of this work
was made by an invitation from the Muséum National d’Histoire Naturelle, with the sponsorship of
Professors C. Lévi and C. Delamare Deboutteville, through the director, M. J. Dorst and his collea-
gues, as invited Associated Professer in this institution ; the sejourn thus made possible was mainly
passed in the laboratory of M. C. and M™® F. Monniot. We are much indebted to ail those mentioned.
M.\TERIAL AND METHODS
Details hâve been provided in previous publications (Illg, 1958; Dudley, 1966; Ooishi &
Illg, 1977) on the collection and processing of materials. Most anatomical studies were pursued on
the basis of observations and dissections made by use of the lactic acid method described in the above
référencés. A serious objection to this procedure has been the difficulty of préparation of permanent
slide mounts of the dissections. A variety of media hâve been tried, but none has proven entirely
satisfactory. Methods by which the dissected appendages arrive in a mounting medium by washing
and transfer procedures are likely to resuit in the loss of some of the minute structures, such as the reduc-
ed mouth appendages of many of the species treated here. Mainly for this reason we hâve tried always
to provide an entire animal as the holotype, preserved in alcohol, and we consider the préparations
from which most of the anatomical drawings hâve been made as paratypic material. Types, as will
Source : Mt'IHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
9
be designated below, are installed in the National Muséum of Natural History, Washington (numbers
are designaled by the traditional USNM rubric), in the Muséum National d’Histoire Naturelle, Paris,
and in the personal collection of the senior author.
Drawings were almost enlirely made by caméra lucida. We take the opportunity to présent
a few illustrations prepared from the records of Professor Chatton. Inking of the drawings, a number
of the caméra lucida renderings and most of the laborious task of composition of the plates hâve been
the work of Ruth Illg.
II. — FAMILY ASCIDICOLIDAE THORELL, 1859
Notodelphyidae, Thorell, 1859, p. 344-3589 (part) ; 1860, p. 117, 124-140 {part) ; 1862, p. 7-9, 14, 15-26
(part). — Claus, 1862, p. 102 (part). — Gerstaecker, 1863, p. 404 (part). — Claus, 1864, p. 379-
380 (part). — Buchholz, 1869, p. 100-101 (part). — Brady, 1878 (vol. 1), p. 18, 30,122-124 (part).
— Claus, 1880, p. 553 (part). — Carus, 1885, p. 341 (part). — Thompson, 1893, p. 189-190 (part).
— Graeffe, 1902, p. 39-40 (part). ■— Norman & Scott, 1906, p. 201-203 (part). — Norman &
Brady, 1909, p. 400, 402 (part). — Schellenberg, 1922, p. 219-220 (part). — Brehm, 1927, p. 490
(part). — Seweli, 1949, p. 174 (part). — Barnard, 1955, p. 237 (part).
Notodelphidae Gerstaecker, 1863, p. 4021 (part).
Notodelphyiden Claus, 1875, p. 350 (part). — Timm, 1894b, p. 396 (part).
Ascidicolidae Thorell, 1859, p. 338, 340, 344-358, subfamily of Notodelphyidae (+ Buproridae, p. 340,
358-359) ; 1860, p. 119 (+ Buproridae, p. 119, 139-141) ; 1862, p. 14, 18, 56-57(-l- Buproridae,
p. 7, 14, 17, 61). — Gerstaecker, 1870-71, p. 719 (part). — Kossmann, 1874, p. 2^-289. — Auri-
villius, 1882b, p. 96-97, 108, 109 (part) ; 1883, p. 92-93, 104 (part). — Giard, 1888, p. 505 (part).
— Canu, 1891a, p. 472, 475 (part) ; 1892, p. 25-30, 46, 54, 64-95, 107-108, 116-119, 127-133,
186-224 (part). — Giesbrecht, 1901, p. 76-77 (part) ; 1902, p. 455 (part). — Thompson
and A. Scott, 1903, p. 273, 255 (part). — T. Scott, 1906, p. 363. — Caïman, 1908, p. 177, 182
(part) ; 1909, p. 103 (part). — Chatton, 1909, p. 14 (part). — Chatton & Brément, 1909a, p. 12
(part) ; 1909b, p. 196 (part). — Smith, 1909, p. 66 (part). — Stebbing, 1910, p. 550 (part). —
Gravier, 1912a, p. 70 (part) ; 1913, p. 66 (part). — Chatton & Brément, 1915, p. 143-144 (part).
— Chatton & Harant, 1924b, p. 361 (part) ; 1924e, p. 416 (part). — Pesta, 1934, p. 8, 9,
11 (-[- Buproridae, p. 8). — Cang, 1948, p. 25-27 (-|- Buproridae, p. 6, 25, 26, 27 ; + Botryl-
liphilidae, p. 23, 25) ; 1949, p. 6. — Sewcll, 1949, p. 153 (part). — Changeux & Delamare
Deboutteville, 1956, p. 155 (part). — Gotto, 1957, p. 282. — Stock, 1959, p. 74 (part). —
Bresciani & Lützen, 1962, p. 373. — Bocquet & Stock, 1962, p. 293. — Monniot, 1965, p. 158. —
Stock, 1966, p. 211, 215. — Lützen, 1968, p. 96-97. — Gotto, in Anderson & Rossiter, 1969,
p. 464. — Ooishi & Illg, 1977, p. 147.
Ascidicolîdés, Gravier, 1913, p. 28. — Chatton & Harant, 1922c, p. 251.
In many regards the characters of the family still can be particularized in terms of the excellent
discussion by Schellenberg (1922, p. 277-281), although he refers to the assemblage as the subfamily
Ascidicolinae and there are certain corrections to be applied to his terminology of the mouthparts.
He does not include Buprorus, but the range of his discussion accounts for the généra Ascidicola, Botryllo’
philiis, Enterocola, Enteropsis, “ Aplostoma ” and Mychophilus.
His approach was to construct a range of variation for each appendage and to show for this range
two important considérations ; 1) that the characters as they appear through the varions généra, form
the graduation ; and 2) that it is the graduation in variation that is the essential character for the fami¬
lial définition. To a considérable degree he presses the points that the graduations consist of réductions
and that these represent adaptations to progressive degrees of parasitic (or otherwise symbiotic) existence
Source : MNHN, Paris
10
PAUL L. ILLG AND PATRICIA L. DUDLEY
ScHELLENBEBG (Icmonstrated his conviction as to the close afTinity of the ascidicolids and noto-
delphyids by including them in a single family. This view, it woiild appear, also led him to develop
an impression of great significance of prehensile adaptations in the head appendages of the whole
assemblage. A major defming îeature of the Notodelphyidae then emerged in the very persistent
presence of the prehensile antenna. By adding the incubatory cavity of the female, a diagnostic
morphological criterion for this family, has been formulated which has very successfully confronted
the addition of a great number of newly described forms (Illg, 1958; Illg & Dudley, 1961 ; 1965,
Stock, 1967), and has also held good in a scrutiny of developmental evidence (Dudley, 1966).
Unfortunately to extend this first phase of the approach to the formulation of a définition of
the Ascidicolidae involves the intellectually very unsatisfactory procedure of invoking négative crite-
rian — the absence of prehensile modification of the antenna, and the absence of the incubatory cavity
in the female. Indeed, even the négative criterian are not thoroughly applicable since the speeies
of Styelicola and copepodids and males of Enterocola speeies hâve antennae with somewhat the aspect
of prehensile modification, and females of Buprorus hâve the incubatory cavity (Schellenberg, in
fact, was apparently puzzled by the indicated afTinities of Buprorus, since he indicated in his treatment
that he considered the relationship of the genus as unknown).
In developing his thesis on modification of appendages in ascidicolids, Schellenberg pointed
out that his base forms — Ascidicola and Botryllophilus — show a very fair conformity in the head
appendages to the notodelphyids, except for the antennae and the maxillipeds. The latter appendage
shows a very strong modification for préhension in Botryllophilus (indeed, in ail botryllophilins and
haplostomins) in very distinct contrast to ail notodelphyids. The conformity to this prehensile pattern
in the maxilliped, or some degree of dégénération from it, formed a concept of emphatic significance
in Schellenbebg’s définition of the .\scidicolinae. To some degree he was led into error by consider-
ing the maxillae of enteropsins and enterocolins to be maxillipeds and thus to conform to his sériés.
In fact the latter two subfamilies lack maxillipeds entirely. But, this situation might in a way be consi¬
dered to conform to a concept of modification of mouthparts as the essential prehensile device, with the
transfer from one appendage to the next anterior of this powerfully expressive adaptive fonction for-
ming part of a single evolutionary trend.
A character which Schellenberg also stressed as one unifying the ascidicolin (his term) lineage
is the structure of the fîfth legs. This can also be very reasonably endorsed in light of modem informa¬
tion. The pattern here is very emphatically a complex one, involving on the one hand trends to forma¬
tion of very complicated structures including body éléments beyond the mere appendages, and on the
other hand proceeding in réduction to the point of complété disappearance.
Schellenberg discussed ail the appendages in his diagnostic treatment. The range of material
available to him from literature and from collections was distinctiy limited, so that his présentation
amounts essentially to a sketch. In adopting his basic viewpoint on the composition of the group and
his approach in defining it, we propose to develop the treatment of the appendages, after his method,
to the extent we can do so with our information.
The pronounced sexual dimorphism of the ascidicolids and the variety of habituses assumed by
the females in relation to the varions modes of symbiotic existence are reflected in great superficial
anatomical diversity. Thisisdealt withbelowby describing the range of variations foreachof thefeatures
of the body and the divergences found in the two sexes, as far as we can judge.
Body form is so variable in the Ascidicolidae as to defy formulation of an unequivocal statement
in positive terms. Segmentation varies from clear-cut with well-demarcated units making up the
greater part of the body to a condition of grub-like obsolescence of segmentai boundaries and cuticular
démarcations. The appendages demonstrate that there are as usual an apical piece bearing antennules
and eyes and typically this is incorporated into a cephalosome including as well the antennae, the gnathal
appendages. and the segment of the maxillipeds. The latter appendages are lacking in several instances
but there are some developmental grounds for assuming the persistence of the segment itself. The
first to fourth legs may originate on clear-cut segments, with good articulation of the protopodite on
the body ; alternatively there is a degree of graduation to a metasomal section without segmentai démar¬
cations. The substance proper of the fifth legs seems to participate in the formation of pediform
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMIMES
11
projections, including these legs themselves as well, often serving to envelop the anterior portions
of the egg sacs. There is often no clear démarcation between the metasome, the element of the pediform
processes and the segment bearing the génital apertures ; as a resuit a complex not often found in other
copepods makes its appearance here and suppresses the usual copepod major body articulation.
The urosomc typically incliides true abdominal segments, up to three, and the anal somite.
There are two lines of modification. In many forms abdominal segmentation is reduced or obsolescent.
In the hotryllophilins a phenomenon of something like cuticular annulation either gives rise to super-
numerary segments or results in the appearance of doing so ; exact morphological information for pré¬
cisé opinion in the matter is so far lacking.
Grcat inflation of the body and development to gigantic size, 8 mm body length in species of
Enleropsin, are frequent among the trends in body modification. The céphalothorax tends somewhat
less to modification and réduction than does the urosome. There are typically paired external ovisacs ;
these may be fused, as in species of Botryllopkiliis ; in Biiprorus there is an internai incubatory cavity
and it is not known whether the egg mass is subdivided. The appendages are ail modified ; in the females
noue retains the basic cyclopoid habitus, and a variety of modifications at the generic level is found.
The basic form of the males is certainly of generalized cyclopoid habitus, but there are modifications
of appendages. In most of the généra for which they are known, the males are actively swimming
forms, and indeed this propensity may be the reason for the rarity of discovery of males in the hosts
of the females.
There seems to be a rostrum basically, but it is reduced to absent in many représentatives.
The antennule throughout conveys the impression of réduction and specialization. Among
the females, of the long known species that of Ascidicola has the most developed segmentation, with
five segments. There is a great number of setae borne on these segments, possibly indicative of a
suppression of segmentation. Heavy sclerotization also suggests a considérable degree of specializa¬
tion. In Styelicola there are seven or eight antennular segments. It would be easy to visualize a more
numerously segmented antennule of quite reasonable correspondence to the cyclopinid type as the
forerunner of this appendage in either case. One of the least well-known of the species we include is
Buprorus nordgaardi Sars 1921. In the few figures he gives for the features of this animal, Sars depicts
(pl. 37, fig. la 1), and corroborâtes by statement in the text, an antennule of seven segments. The
appendage has a distinctive flattening and some slight suggestion of sclerotization. The general
pattern of the segmentation and the setation suggest great correspondence with the antennules of the
other species of Buprorus, which, however, hâve reduced segmentation (3 segments). A similar corres¬
pondance of the basic éléments of the profusely setiferous 4-segmented antennules of some Botryllo-
philus females could also be surmised without too great exaggeration. In the other subfamiiies of
the présent treatmenl the pattern of réduction of the antennule reaches such a degree that morpholo¬
gical continuities cannot be reasonably traced out. The antennule, or possibly the antenna, disappears
in Lequerrea.
Although the most developed segmentation of the antennule is in the males and evidently the
dimorphism is considérable, no male has been found with prehensile antennules. Unfortunately, the
male of Styelicola is still unknown. The greatest number of antennular articles foundis in the male of Entera-
cola, with seven or eight articles. These actually hâve fewer setae than the five-segmented antennule
of male Ascidicola. The indications are that the males of enteropsins hâve nearly as full antennular
segmentation and setation as in Enterocola. In males of Botryllophilus, as in the females, there are
only four antennular articles, and a reduced setation, but on the basal article there is developed an
enormous number of aesthetes. The male of the genus Haplostoma shows the same feature (Ooishi
& Illg, 1977). The male is not known at ail for buprorins.
The antenna strongly suggests affinity with notodelphyids in basic construction but difîers
in adult females of ail généra but Styelicola by absence of a terminal, articulated, prehensile hook.
The trends in réduction and specialization are distinctly different from any such in the notodelphyids.
Our studies of development show that the basic antenna consists of a single protopod segment and a
two-segmented endopod.
In the unfortunately little known genus Buprorus there is an antenna of such generalized cyclo-
Source : Paris
12
PAUI. L, ILLG AND PATRICIA I-. DUDLEY
poid aspect, with three segments, the terminal one bearing numerous setae, that il could be encoun-
lered on a species of cyclopinid. In several lineages — the botryllophilins, the haplostomins, entero-
colins and enteropsins, although the modifications of the antenna are varions, there is a strong trcnd
to flattening of the whole appendage and to réduction in segmentation. In inost of the subîamilies
the antenna of the male is similar to that of the female. A most interesting exception is in Enlerocala,
where the antenna in the male bears a prehensile seta and is of a very different general aspect from that
of the female. Tliis is interesting also in reference to the presence on the early copepodids of E. fulgeiis,
as depicted by Canu (pi. XIX), of somewhat similar antennae, exlremely reminiscent of the appendage
as it occurs in copepodids throughout the Notodelphyidae. In Haplostomella the males and females
bear comparable highly distinctive antennae, witli a distal curvature and stout terminal seta, sugges¬
tive of a possible prehensile adaptation.
There is typically a prominent labrum, and in inany forms it exhibits distinctive ornamentation,
but there does not seem to be an available generalization in ihis regard, serving as a family charac-
teristic. Paragnaths and labium hâve not heretofore been found in any représentative, this in contra-
distinction to the persistent occurence throughout the notodelphyids. However, in some ascidicolids
there is a somewhat sclerotized, or even spinulose, postoral area on the integument, and for EnUrocola
pelili Guille, there is depicted (1964, fig. 1, d) a postoral area with seta-like structures inserted bila-
terally. We présent below a figure (fig. 1, g) of the paragnath of Ascidicola ; it is notably like thosc
of a number of notodelphyids. Enlerognathus has been known since the original description to possess
well-developed paragnaths.
The mandible tends to show clearly the basic gnathostome dérivation of the whole stock. The
masticatory base in Ascidicola, Buprorus, and Botryllophilus is typical. The réductions in mandibles
in a somewhat graded sériés of examples we can also find paralleled in sequences of forms of notodel¬
phyids. However, the mandibular palp, in such comparison with notodelphyids, is reduced even
in the most generalized of examples, such as Botryllophilus. In some ascidicolids (enteropsins, entero-
colins] the mandible disappears in the adults ; the comparable cases known among notodelphyids are
in extremely modified forms only. Even in généra in either family lacking the adult mandible the
persistence of the appendage in the early copepodids shows the gnathostome affmity, as constrasted
with poecilostome development. Some of the trends in réduction in segmentation and setation seem
to parallel somewhat those in cyclopinids. In the généra Botryllophilus and Haplosloina there is a
sexual dimorphism in regard to characters of the mandible and in most cases for which the male is
so far known, if the appendage is lacking in the female it does not occur in the male.
The maxillule is typically a modified appendage, and remarkably enough, is very rarely absent.
The basal endite is présent in botryllophilins, ascidicolins and buprorins, where it is well developed,
with numerous setae, and overall strongly resembles that of notodelphyids and thus also cyclopinids
and archinotodelphyids. The haplostomins show a graduation in this feature, since some généra lack
the maxillule, but Haplostomides has a maxillule resembling that in botryllophilins ; the endite is pro¬
minent. The palp in these forms shows réduction of the exopodite to obsolescence and a general
coalescence of the distal éléments of the appendage. In enteropsins and enterocolins there is a distinc¬
tive modification. The bilobed appendage appears to préservé some biramous indications, although
modified by coalescences. It remains probable also that one of the conspicuous lobes is actually the
modified endite. Such a modified appendage suggests an aspect of arrest in a developmental stage.
The maxilla is remarkable in being almost typically cyclopoid in general aspect in botryllo¬
philins and with liltle more modification in ascidicolins and buprorins. In none of these, however,
will there be found the full development of endites or setae of the generalized appendage. The basic
cyclopoid éléments are more strongly modified, strongly suggesting prehensile adaptation, in enteroco¬
lins and possibly Enlerognathus and extremely so in enteropsins. In haplostomins the appendage
is suppressed to absent and when présent is a much rediced setiferous lobe. The appendage in botryllo¬
philins, ascidicolins and buprorins is remarkably similar to that in some of the modified notodelphyids
living in compound tunicates. In enterocolins and enteropsins the maxilla is massive, seems to approach
a prehensile condition and is very prominent in the mouthpart sériés. It resembles strikingly the maxil-
liped in botryllophilins and haplostomine.
Source : AlfJHW, Paris
ASCIDICOLIDAH AND SUBF^VMILIES
13
Maxillules and maxillae are the same in the male as in the female of ascidicolins. There is
good evidence to suggest sexual dimorphism in the maxillules and maxillae in botryllophilins and
haplostomins, and the features involved in turn reinforce the impression of rather close relationship
of these two subfamilies. In the male of some species of BotryÜophilus the maxillules and maxillae
are so suppressed in the adult stage that they hâve been overlooked in some examinations. In the
enteropsins, based on what little knowledge we hâve of males these appendages may be expected to
show much less dimorphism, but ail indications are of some degree of réduction in males. The males
of Enlerocola show a distinctive trend of modification, with élongation of the appendages and their
setae and loss of the gnathal processes.
The maxillipeds exhibit great diversity and the aspect in ail cases is of strong modification from
the basic cyclopinid appendage. In ascidicolins and buprorins the appendage is a reduced setiferous
lobe like that in some modified notodelphyids from solitary tunicates. Enteropsins and enterocolins
hâve lost the appendage. In Enterognathus either the maxilla or maxilliped is lost. In the botryllo¬
philins and haplostomins there is an exaggerated modification to render the maxilliped the most salient
appendage of the forebody. It becomes massive, conspicuously prehensile, and acts in a variety of
functional applications. It remains remarkably consistent in structure throughout both subfamilies.
From indications so far, there is little dimorphism in the maxilliped ; the males of haplostomins
and botryllophilins seem to hâve the appendage as well developed in proportion to the other appendages
as do the females. In the family in general when the appendage is lacking in females it is also lacking
in the males.
The four pairs of appendages of the second to fifth thoracic segments, usually referred to in
copepods as the swimming legs, always show distinct modifications toward other functional applica¬
tions in the females throughout the ascidicolids. The patterns are essentially generic and details
will be provided below in the subfamilial or generic diagnoses. The sexual dimorphism of these appen¬
dages is pronounced and males tend to hâve the aspect of cyclopinids, with legs of actual swimming
function, except perhaps the first leg of the male of BotryÜophilus, and some species of Flaplosloma,
in which the endopodite is modified into a clasper-Iike structure.
The fifth legs in female ascidicolids show extremes of modification and, alternatively, réductions
to disappearance. There are strong indications that protrusions of the actual body participate in the
structure, particularly in the notably expanded legs. In the literature the varions protrusions of the
corresponding somite might be found referred to as pterostegites, oostegites, or fifth legs. In particular
the school of Cratton tends to term the protrusions oostegites, doubtless because of the usual close
association with the ovisacs. An early viewpoint suggested there was direct homology of the lamellar
fifth legs of ascidicolids with the dorsal excrescences of varions notodelphyids, such as Notopleropkorus,
and with the incubatorium of notodelphyids. This latter idea, particularly, has beenrefuted by
Chatton and his associâtes. We are much înclined to agréé in large part, except for designating the
structures as oostegites, with the position of Chatton’s group as stated in Chatton & Habant, 1924c,
p. 405-406 : “ Thus the oostegites of the Ascidicoles must be considered, not as being themselves the
hypertrophied pereiopods of the fifth pair, but as dorso-lateral duplicatures homologous to those observ-
ed in Notopleropkorus, some Enlerocola species and in Haplostomeüa tuberculala, and carrying the pereio-
pocls of the fifth pair, or their vestiges, on the free margin. They thus are not to be considered as simple
appendages but as complexes formed of an appendage of ventral origin and of an integumental dupli-
cature of dorsal origin. ” We cannot go entircly as far as this statement. There are quite possible
cases where the fifth legs hâve not appeared at ail, in which case the protrusion on the sixth thoracic
somite would be only a pterostegite. The homologies involved cannot be thoroughly traced until
full anatomical information and the developmental sequences in the late copepodid stages hâve been
worked out. It is pertinent to note that there are very few descriptions available of male ascidicolids,
but those known are comparable in urosomal segmentation and in development of ventrally placed
fifth and so-called sixth legs with those known from the notodelphyids.
We feel the term oostegite should not be used in application to any protrusion of the thorax,
because of confusion with the term as otherwise applied in the higher Crustacea. There are undoubted
cases of occurrence of pterostegites (i.e., Enlerocola pterophora). There are perhaps instances of plate-
Source : Mt-JHN, Paris
14
PAUL L. ILLG AND PATRICIA L. DUDLEY
like fifth legs [Buprorus spp.). But in light o( présent knowledge we prefer to use throughout the ascidi*
colids for the females, the term pediform projection wherever the structure seems to support setae
referrable to the cyclopoid fifth leg, and where the amount of involvement of additional structure is
uncertain. In many cases in the ascidicolids these pediform projections are lamellate and, although
they dérivé structurally from the somite of the fifth legs they associate very consistently wilh the ovidu-
cal apertures and with the ovisacs. The position may be ventral, latéral, or considerably dorsal, but
they essentially take origin on the metasome. The pediform projections in botryllophilins, which are
supportive of the dorsal ovisacs, fused into a single globular mass in most cases, are seen to originate
more anteriorly from the metasome and thus apparently protrude from the segment of the fourth legs,
hecause the somites are fused. In Ascidicola and Styelicola the body articulation is complicated by
the great extension of the sixth and seventh thoracic somites and the accompanying enlargement of the
pediform projections. A parallel situation appears in the Botryllophilinae', where in Schizoproctus
inflatus and its near allies the pediform projections enclose the ovisacs and the body articulation is
obscured by the related fusions and complexities. The Enteropsinae are distinctive in lacking fifth
legs or pediform projections.
The pediform projections, in relation to the ovisacs, may envelop them completely, as in Ascidi¬
cola and Schizoproctus. They may tend to encircle the egg masses (or mass) extensively, without com-
pletely covering them, as in Botryllophilus. In many species of haplostomins and enterocolins the
pediform projections form caps over the proximal portions of elongate egg strings and may almost
encircle the urosome. In Buprorus, which has developed an internai incubatory cavity, the pediform
projections are ventral and extend prominently, with no relation to egg protection, and they are perhaps
no more than truly enlarged fifth legs.
On the seventh thoracic somite of female ascidicolids there may also be found setiferous projec¬
tions, located at the oviducal apertures [Ascidicola, enteropsins, Schizoproctus, and others). These
were referred to as vulvae in some of the descriptions of Chatton and associâtes, but this usage does
not seem appropriate. Such structures, when there are prominent setae, hâve often been referred
to as sixth legs in the hterature on cyclopoids. There is a possibility that some of the claimed occur¬
rences of fifth legs in ascidicolids may dérivé from misidentification of these structures (see p. 96
below, in the discussion of Enteropsinae). These setiferous projections vary greatly in aspect through
the various species and are undoubtedly complicated in anatomical composition. There is insufficient
evidence to attempt further analysis of them at présent.
The great complication in formation of the metasomal-urosomal articulation in the female
ascidicolids to some degree parallels the phenomenon in notodelphyids, but only in the case of Buprorus
does the ascidicolid body develop an inclosed incubatory cavity. It is rather more usual for the ascidi¬
colids to form a genital-urosomal complex, but the diversity of arrangements is so extensive that no
further general statement can be made. There is a great tendency for the oviducal apertures, suppo-
sedly diagnostic of the seventh thoracic somite, to open near the posterior basal juncture of the pediform
projection with the body, just posterior to it, and on the urosome.
The notodelphyid and ascidicolid females evoke an impression of considérable similarity in the
structure of the female insémination apparatus, and indeed the characteristic médian insémination
pore, also widely referred to as the vulva, is considered by Canu to be found throughout gnathostome
cyclopoids, and still other major copepod stocks. The médian pore of the female, the site of attach-
ment of the spermatophores, is usually conspicuous and may hâve accessory sclerotizations ; it coramu-
nicates by diverging branches with paired séminal réceptacles lying laterally close to the terminations
of the oviducts. Here at the apertures will be found the structural features discussed above as often
referred to as sixth legs.
There is a great variety of modifications at the generic level of the caudal rami, particularly
in females. Many of the features can be found to resemble most interestingly similar spccializations
among some of the notodelphyids. Réduction, but with only extremely rares case of complété disap-
pearance perhaps forms one such trend. Sclerotization, as in Ascidicola, tends also to progress to
conversion of the setae into hook-like éléments, as in Botryllophilus. In the males, the caudal rami
Source : iMfJHW, Paris
ASCIDICOLIDAE AND SUBFAMILIES
15
are much more cyclopoid, but too few instances are known to make broad generalization possible at
this point.
In summary we can say that no ascidicolid adult female bas a generalized cyclopoid habitus,
although the appendages are basically cyclopoid, so such females could only be confused with nono-
vigerous notodelphyids and certain vermicole cyclopoids of more or less uncertain taxonomie position
among the poecilostomes. The antennae of female ascidicolids almost always lack the terminal pre-
hensile hook which is présent throughout female notodelphyids. We find the configuration, segmenta¬
tion, and setation of the antennae of ascidicolins, haplostoinins, botryllophilins and buprorins to con-
form so consistently that we consider this a strong indication of the familial relationship, and diagnostic
of the family. In the remaining subfamilies the antennae are extremely modified but we conclude
they dérivé from this pattern. In a somewhat similar way the development of extensive expansions
involving the fifth legs associâtes ail the subfamilies except the enteropsins and contrasts with ail
notodelphyids. The enteropsins are so similar to the enterocolins in the remarkable modification
of the mouthparts, including loss of the mandibles and maxillipeds, that there is no difficulty in
assigning both to the ascidicolids. Although so few examples are known, the males hâve so far been
assignable to their généra by the structure of the mouthparts. In assigning a male to the family, only
the mouthparts would be satisfactorily diagnostic, and the whole sériés of these would hâve to be con-
sidered in combination.
KEY TO THE SUBFAMILIES
Mature females, preferably ovigerous, can be assigned to subfamilies by use of the following
key :
1. Internai dorsal brood-sack. Buprorinae, p. 30
1". No internai brood-sack. 2
2. No pediform projections or fifth legs. Enleropsinae, p. 94
2". Pediform projections and/or fifth legs varyingly developed. 3
3. Caudal rami ornamented terminally with clawed prehensile hook-like éléments.
Botryllophilinae, p. 136
3”. Caudal rami ornamented with spiniform or setiform éléments which may be much reduced. 4
4. Pediform projection-fifth leg lobate, set with few, reduced setae, tend to substantial réduction..
Haplostofninae, p. 132
4". Pediform projections substantially expanded, lamelliform or lobate, usually setiferous.... 5
5. Antennules well developed with 5 to 8 well-articulated setiferous segments.
Ascidicoîinae, p. 16
5'. Antennules expressive of substantial réduction, few-segmented, weakly setiferous. 6
6. Basipodites of legs 1 to 4 with setae reduced to absent. Enterocolinae, p. 35
6'. Basipodites of legs 1 to 4 with conspicuous long latéral setae. Enterognathinae, p. 151
We hâve felt it necessary to remove a species of ascidicolid from the genus in which it was
described, because the details furnished by the author départ substantially from what are now rather
well tested generic concepts. We suspect an actual confusion of parts of specimens was involved in
constructing the original description, as discussed below (p. 132). We are at a loss, with the informa¬
tion available as to which subfamily would accept the species as described, so must place as species
incerta sedis in the family Ascidicolidae Enteropsis pilosus Canu, 1886.
Source ; MNHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
SUBFAMILY ASCIDICOLINAE, THORELL, 1859
Notodelphyida» Thorell, 1859, p. 344-358 (part) ; 1860, p. 117, 124-140 (part) ; 1862, p. 7-9, 14, 15-26
(part). — Buchholz, 1869, p. lOO-iOl (part). — Claus, 1862, p. 102 (part) ; 1872, p. 418-419;
1880, p. 553 (part). — Thompson, 1893, p. 189 (part). — Graefle, 1902, p. 39-40 (part). — Norman
& Scott, 19W, p. 201 (part). — Norman & Brady, 1909, p. 400, 402 (part).
Cetochilidae White, 1850, p. 115-116 (part).
Notodelphidae Thorell, Gerstaecker, 1863, p. 404 (part).
Ascidicolidae, Thorell, 1859, p. 338, 344-358, subfamily of Notodeiphyidae ; Thorell, 1860, p. 117,
124-140 ; 1862, p. 7, 9, 14, 15-26. — Buchholz, 1869, p. 101. — Gerstaecker, 1870-1871, p. 719
(part). — Claus, 1872, p. 419 (part). — Kossmann, 1874, p. 288-289. — Brady, 1878, vol. 1,
p. 30, 144-145. — Délia Valle, 1883, p. 252. — Canu, 1891a, p. 20, 472, 475 (part) ; 1892, p. 64-
96, 107-108, 116-119, 127-128, 130, 133, 186-224 (part). — Scott, T., 1901b, p. 241-245 (part) ;
1902, p. 455 (part). — Thompson & Scott, 1903, p. 255 (part). — Caïman, 1908, p. 177, 182
(part). — Chatton, 1909, p. 14 (part). — Chatton & Brément, 1909b, p. 196 (part). — Smith,
1909, p. 66 (part). — Stcbbing, 1910, p. 550 (part). — Gravier, 1912, p. 70 (part). — Chatton &
Brément, 1915, p. 143-144 (part). — Sars, 1921, p. 63. — Chatton & Harant, 1924b, p. 361
(part) ; 1924e, p. 416. — Pesta, 1934, p. 8, 9,11. — Lang, 1948, p. 25, 26, 27 ; 1949, p. 6. — Chan-
geux & Delamare Deboutteville, 1956, p. 155 (part). — Gotto, 1957, p. 282. — Stock, 1959,
p. 74 (part). — Bresciani and Lûtzen, 1962, p. 373. — Stock, 1966, p. 211-215 (part). — Lûtzen,
1968, p. 96-97.
Ascidicolinae Brady, 1878, (vol. 1), p. 144-145. — Carus, 1885, p. 341. — Chatton, 1909, p. 14. — Chatton
& Brément, 1915, p. 143-144, 145,148 (part). — Schellenberg, 1922, p. 277-298 (part). — Brehm,
1927, p. 490 (part). — Barnard, 1955, p. 237 (part). — Bresciani & Lûtzen, 1962, p. 373. — Gotto,
1966, p. 192.
Ascidicolinen Thorell, Claus, 1875, p. 350.
This taxon has been known for a very long time as monotypic, often under rather confusing
alternative désignation of Ascidicolidae, family, or Ascidicolinae, subfamily. The discovery by Lût¬
zen, 1968, of a second genus, Styelicola, represented at that time by a single species of very different
habitus from that of A. rosea, produced a considérable expansion of the concept of the subfamily.
The new species to be described below conforms very well in most features to Lützen’s genus, but in
general habitus is very similar to A. rosea. The male is not known for Styelicola so the formulation
of a concept of the subfamily dépends on the characters of the female.
The form of the body is variable, but in A. rosea and the new species of Styelicola the aspect
is fusiform, with well-marked segmentation, obscured at the mid-body for formation of a complex
bearing the expanded lamelliform pediform projections involving the fifth legs. In Styelicola bakusia
the body might -well be said to be enterocoliform, but most of the appendages and in particular the
lamellate projections involving the fifth legs are readily diagnostic. The antennules are regularly
tapered, of fairly numerous segments and regularly and fairly abundantly setiferous. The antenna is
trimerous, the terminal segment with a prominent claw-like apical element in Styelicola ; in Ascidicola
with four setae and a stouter spine-like element with a suggestion of prehensile modification. The
mandible has a very strong, markedly toothed masticatory coxal lamella and a fairly prominent palp
bearing several éléments. The other mouthparts show distinct modifications, but form a part of a
graduated sériés of adaptations found throughout the family. The “ swimming ” legs one to four
show distinctive modifications, these varying most strikingly with the species. The fifth legs are
Source ; Mt-JHN, Paris
ASCIDICOLIDAH AND SUBFAHILIES
17
involved in expanded lamelliform projections, each of which shelters a part or the whole of an ovisac.
In Slyelicola the fifth leg is further represented by an articulated endopodite bearing several well-
developed setae. In Ascidicola the lamella is set with a single seta near its apex ; the conBguration
is not readily separated in description front the lamellate projections found in species of Schizoproclus
in the BotryUophilinae. The caudal rami in the ascidicolins are variously modified and bear apical
setae. Females of the two généra niay be diflerentiated by means of the following key.
KEY TO THE GENERA
1. Fifth legs conspicuous expanded lamellae ; ornamentation a single seta. Ascidicola Thorell, 1859
1'. Fifth legs include a conspicuous expanded lamella, this supporting an articulated endopodite
bearing several long setae. Styelieola Lützen, 1968
Ascidicola Thorell, 1859
NotoiUlphys Allman, 1847, p. 2 (type, by monotypy, N. ascidicola Allman, 1847 [indéterminable species])
(part). — White, 1850, p. 116 (part). — Leuckart, 1859, p. 247 (part).
Ascidicola Thorell, 1859, p. 337, 340, 356, 347 (type, by monotypy, A. rosea Thorell, 1859) ; 1860,
p. 116, 119, 127, 128 ; 1862, p. 6, 14, 57-59. — Claus, 1862, p. 102 ; 1864, p. 380. — Hesse, 1866,
p. 52. — Buchholz, 1869, p. 149. — Gerstaecker, 1870-1871, p. 719. — Claus, 1872, p. 419. —
Kossmann, 1874, p. 288-289. — Claus, 1875, p. 350-351, 360. — Brady, 1878, p. 21, 30, 145.
— Kerschner, 1879, p. 20. — Aurivfflius, 1882b, p. 95-97, 109, 110 ; 1883, p. 91-93, 105, 106 ;
1886, p. 45-46. — Canu, 1891a, p. 475 ; 1892, p. 29, 108, 204, 208-209, 214. — T. Scott, 1901a,
p. 352. — Graeffe, 1902, p. 40. — T. Scott, 1907, p. 367. — Caïman, 1909, p. 103. — Norman &
Brady, 1909, p. 402. — Smith, 1909, p. 66. — Hartmeyer, 1909-1911, p. 1734,1735. — Chatton &
Brément, 1915, p. 144,145,147. — Sars, 1921, p. 64. — Schellenberg, 1922, p. 280, 281. — Brehm,
1927, p. 90. — Harant, 1931, p. 370. — Wilson, 1932, p. 599, 600, 626. — Pesta, 1934, p. 11-12.
— Neave, 1939, p. 314. — Lang, 1949, p. 6. — Gotto, 1957, p. 281-289. — Monniot, 1959, p. 158 ;
1965, p. 158-159, fig. 43A-43U. — Gotto, 1966b, p. 162. — Lützen, 1968, p. 97, 101.
Coiliacola Hesse, 1862, p. 343 (type, by monotypy, C. seligera Hesse, 1862). — Neave, 1939, p. 794.
Coiliacea Hesse, 1863, p. 110 (misspelling for Coiliacola Hesse).
Coeliacola Sars, 1921, p. 64 (in synonymy, misspelling for Coiliacola Hesse).
We consider Ascidicola to be monotypic, so the characters of the genus are the same as those
presented for the single species below. To assure the identity of the example from the Pacific Océan
studied by us with the species as long known from Europe we hâve had to study the anatomy of Euro¬
péen représentatives in detail We illustrate below mainly from specimens from France and find that
the features correspond in our local specimens. We hâve collected here only females and cannot give
new information about males.
Ascidicola bosea Thorell, 1859
(figures 1, 2, 3)
Notodelphys ascidicola Allman, 1847, p. 2-6, pl. 1, fig. 14, pl. 2, figs. 15-21 (type locality, Ireland, British
Isles, in Ascidia communie) (part). — White, 1850, p. 116 (part).
2
Source : Paris
Fig 1. — AscidUola rosea Thorell, fetnale : a, habitus, latéral view ; b, antennule ; c, antenna ; ‘J- ««S*
ment of antenna ; e, en face view of some cephalosomic appendages : L = labnim, Md = mandible. Mxl - maxd-
lule, Mx2 = maxîUa, Mxp = maxilliped ; f, mandible ; g, paragnath ; h, maxillule ; i, maxüla ; j, maxilliped.
Scalc for a = 0.2 mm j other scales = 0.1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Fio. 2. — Ascidicola rosea Thorell, female : a, fiwt leg ; b, second leg ; c, third leg ; d, fourth leg ; e, pediform projection-
fifth leg, latéral view ; f, surface ornamentation of " fifth leg ” ; g, internai surface of fifth leg enclosing sac of
ova ; b, génital aperture ; i, caudal ramus. Soales for e and g = 0.2 mm ; other scales = O.l mm.
Source ; MNHN, Paris
20
PAUL L. ILLG AND PATRICIA L. DUDI.KY
Ascidicola rosea Thorell, 1859, p. 335, 337, 340, 343 (type locality, Skagerrak, Sweden, in Ascidia
intestinalis, A. parallelogramma, A. canina, A. aspersa) ; 1860, p. 114, 115, 119, 123,138; 1862,
p. 5, 6, 7, 59-61, pl. 9, 10, fig. 13. — Buchholz, 1869, p. 101, 149-150. — Gerstaecker, 1870-1871,
p. 775, 776, 777, 801. — Clans, 1872, p. 419. — Brady, 1878, p. 145-146, pl. 30, fig. 1-10. — Auri-
villius, 1882a, p. 46 ; 1882b, p. 87-95,111, 113, pl. xvi, fig. 13-22 ; 1883, p. 83-91,106,109, pl. 7,
fig. 13-22 ; 1886, p. 45, 46. — Herdman, 1891, p. 210. — Canu, 1892, p. 67, 209. — Timm, 1894a,
p. 159 ; 1894b, p. 396. — T. Scott, 1901a, p. 352. — Graeffe, 1902, p. 40. — Norman & Scott,
1906, p. 202. — T. Scott, 1906, p. 363 ; 1907, p. 367. — Norman & Brady, 1909, p. 402. — Riddell,
1909, p. 140. — Smith, 1909, p. 66. — Hartmeyer, 1909-1911, p. 1734, 1735. — Chatton & Bré-
ment, 1915, p. 145. — Sars, 1921, p. 64-66, pl. 31. — Schellenberg, 1922, p. 293. — Harant,
1931, p. 370. — Pesta, 1934, p. 12, fig. 8. — Leigh-Sharpe, 1935, p. 48. — Oorde-de-Lint &
Schuurmans Stekhoven, 1936, p. 121, fig. 78. — Lang, 1948, p. 3, 19, 20-21, fig. 17 ; 1949, p. 6.
— Sewell, 1949, p. 182, 184, 188, 190. — Millar, 1953, p. 78. — Gotto, 1957, p. 281-290 ; 1960,
p. 216, 221. — Monniot, C., 1961, p. 98-99, fig. 1. — Bresciani & Lützen, 1962, p. 373. —
Monniot, C., 1965, p. 158-159, 160, 161, fig. 43, A-U. — Gage, 1966, p. 229. — Gotto, 1966b,
p. 162. — Lützen, 1968, p. 97, 101. — Hamond, 1973, p. 350.
Coiliacola setigera Hesse, 1862, p. 343-349, pl. 18, figs 1-15 (type locality, Finistère coast, in Phallusia
canina and P. intestinalis).
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
21
Asidicola acuUoretusa Lang, 1949, p. 6, fig. 14-16 (type locality ,South Georgia, in Pyura georgiana
Michaelsen). — Gotto, 1957, p. 283. — Lützen, 1968, p. 97, 101.
Distribution : Mediterranean to Norway and Sweden, British Isles, South Georgia.
Hosts : .^scidio commuais ”, Ascidia aspersa, A. canina, A. intestinalis, A. mentula, A. paralle-
logramma, A. sordida, A. oirginea.
Ciona intestinalis, C. canina.
Corella paraUslogramma.
Cynthia papillata.
Ualocynthia papillosa.
Microcosmus, M. claudicans, M. nudisligma, M. polymorphus, \1. sabalieri, M. pulgaris.
Phallusia canina, P. intestinalis, P. mentula, P. obliqua, P. virginea.
Pyura georgiana, P. micrososmus, P. papillosa, P. squamulosa.
We hâve listed the names of hosts above as we encountered them in the records of the various
authors, leaving the task of compiling the proper synonymies to the authorities on ascidians. Our
new records add some interesting host associations.
Speciniens exaniined :
From Microcosmus sulcatus (Coquebert), s. lat. :
From Gaiola, Bay of Naples, Italy, 30-40 m, October 7, 1957, 2 females. Near Port Vendres,
near Banyuls-sur-Mer, France, 50-25 m, May 13, 1958, 1 female.
From Ualocynthia papillosa (Linnaeus) :
Gaiola, Italy, 30-40 m, October 8, 1957, 1 female.
Gaiola, Italy, 40 m, November 29, 1957, 1 female.
Cap l’abeille, near Banyuls, France, 25 m, May 7, 1958, 1 female.
From Ascidiella aspersa
Pozzuoli, Bay of Naples, Italy, 35 m, January 25, 1958, 1 female.
From Rhopalaea neapolitana
Gaiola, Italy, 30-40 m, October 7, 1957, 1 female.
From Phallusia nuimmillata
Stazione zoologica, Naples, Italy, physiological laboratory, January-February, 1958, 3 females.
Near Port Vendres, France, 70 m, 60-30 m, mixed trawl, May 6,1958,1 female ; 50-25 m, May 13,
1958, 1 female.
From Corella eumyota Transtedt
ELTANIN Cruise 12 station 1078, 61“27'S-61«26'S ; 41o55'W-41‘'55'W, 604 m, 5' Blake
Trawl, April 12, 1964, 1 female.
From Molgula euplicata Herdman
ELTANIN Cruise 27 station 1878, 72o57' S-72o58' S ; 171035' E-17lo38' E, 576-573 m, 5' Blake
Trawl, January 15, 1967, 1 female from esophagus of host.
We are indebted to M. C. Monniot for the indentiScation of the ascidians in the aiitarctic material
listed just above. From his collections from the coasts of Europe he has also provided us with the
following host records for
Ascidicola rosea : Diazona oiolacea Savigny. Banyuls-sur-Mer ; Ascidia obliqua O. F. Müller, Kristi-
neberg; Polycarpa pomaria (Savigny), Banyuls-sur-Mer; Styela parlita (Stimpson), Banyuls-
sur-Mer. We hâve also :
From Polycarpa obtecta Traustedt
Gulf of Mexico, Albatross Station 2405, 30 fathoma, USNM 452, March 15, 1885, 1 female.
From Styela coriacea hemicaespilosa Ritter :
Off Cerros Islands, Lover California, Mexico, .\lbatross Station 2838, ÜSNM 92218, May 5,
1888, 2 females.
From Ascidia paralropa Huntsman :
Satellite Channel, British Columbia, Canada, August 7-10, 1963, 4 females.
Northwest coast of British Columbia, exact locality unknown, no date, 2 females.
Source : MNHN, Paris
22
PAUL L. ILLG AND PATRICIA L. DUDLEY
Unidentified pyurid tunicate :
Ofl coast of Southern California, University of California collections : “ Channel Dredge, ”
June 18, 1901, 2 females.
Description :
Female : The overall length in a représentative spécimen, to the ends of the caudal rami is
3.6 mm. The body (fig. 1, a) has been frequently figured and described. It is inarkedly cyliudrical
with a slight posterior taper. The cephalosome has a strongly developed shield, which overlaps the
first segment of the metasome. The composition of the metasome is complicated by the coalescence
of the segments of the fifth legs and of the génital apertures and possibly one abdominal segment,
The body articulation has been variously interpreted, but descriptively, the metasome can be stated
to consist of 4 clearly delimited leg-bearing segments and a long complex région bearing the fifth
legs and the génital apertures. The usual major body articulation is not formed. The urosome then
consists of 3 clearly articulated segments, 2 lacking appendages and terminating with the anal somite
and its caudal rami. It was only lately pointed out (Gotto, 1957 ; Monniot, 1965) that the anal
somite bears at its antérolatéral margin a wide transverse pad set profusely with cuticular projections
(Gotto, 1957, fig. 1). This structure is not depicted in the figures here and is rarely observed on preserved
material. Fixation causes a slight contraction, sufiieient to telescope the urosomal articles and retract
the pad. \Ve were struck by the observation that a similar pad is developed on the dorsal surface
of the urosome of Slyelicola lighli (figs. 4a, fie).
The antennule (fig. 1, b) is short, thick, 5-segmented, and the articles are heavily sclerotized.
Ail the setae of the appendage are stiff and most are short. The armature of the 5 articles, I-IV,
is as follows : I — 8 setae ; Il — 7 setae ; III — 5 setae, 1 asthete ; IV — 3 setae, 1 aesthete ; V —
8 setae, 2 aesthetes. Further ornamentation consists of ridgings of the cuticle.
The antenna (figs 1, c, d) is trimerous, the articles decreasing in width distally. Ail the articles
are heavily sclerotized. The basal article has a stout seta at the distal médial corner. The second
article has a flattened spine on the surface ; there is a sclerotized lobe on the opposite surface at the
proximal third. A row of spinules ornaments the distal médial corner. The terminal article is very
narrow, its length about equals the combined lengths of the 2 basal articles. There is a seta in an emar-
gination at the middle of the médial margin. Distally there is 1 spine with a slightly hooked end,
2 spiiuilose éléments of the same length as the spine and 2 cirved, stilTened setae, one about one third
longer thaii the other. A curved row of spinules ornaments the face of the segment at the distal fourth.
The remaining cephalosome structures are described in the orientation of an en face view (fig. 3, b).
The labrum has a more or less simple posterior margin, but bilaterally placed dorsal sclerotizations bear
on either side a row of spiniform processes.
The mandible (figs. 1, e, f) consist of the unimerous protopodite and bimerous palp. The proto¬
podite extends medially as an extremely large masticatory lamella, the médial margin of which is
formed nf a sériés of teeth of very characteristic form, some bipartite and some tripartite. The basal
article of the palp bears no armature but is ornamented with a single row of spinules. The distal
article has an emargination at the latéral fourth and below this there is articulated a long seta. The
apex bears 2 setae and there is an additional seta at the distal médial fourth. The setae are graduated
in length from médial to latéral around the apex. A row of spinules is set at the bases of the 2 terminal
asetac. A pair of small paragnaths (fig. 1, g) is set on a transverse postoral sclerotization at the bases
of the mandibles. The paragnath is a lobe with sclerotizations at the base and along the latéral margin.
The much thinner distal portion has a rounded contour.
The maxillule (fig. 1, g) is a bilobed structure, the segmentai composition obscure. The médial
lobe, probably the basal endite, bears 7 setae. The latéral, more ventrally placed lobe bears 7 spines
and gives no indication of possible composition as to segments or rami.
The maxilla (fig. 1, h) is bimerous, the proximal article longer and very much wider than the
distal. At the distal médial corner of the basal article there is a small endite. Distally this endite
Source ; MhSHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
23
tapers as a spiniform process. There is a setule inserted at the base of the process. The distal article
narrows into a slightly hooked process, at the base of which on the anterior face there is a closely appres-
sed seta. Five small setules articulate on the ventral margin proximal to the hook.
The maxillipeds (fig. 1, i) are joined together by a médial plate so that the pair has a common
articulation on the body. Each consista of a long, fiat lobe, with three apical setae and three médial
setae.
The first legs {fig., 2, a) are biramous with bimerous protopodites and rami. The coxopodites
unité medially to form a common articulation with the body. There is a médial apparatus, consisting
of an elongate bar distal to the common base. There are distinctive sclerotizations. Rows of spinules
ornament the sclerotized and unsclerotized areas. The basipodite is much extended medially so that
the bases of the rami are at markedly different levels. There is a long seta at the distal latéral corner
of the basipodite, and a spine at the distal médial corner, The basal article of the exopodite bears
a spine at the distal latéral corner. A patch of spinules is set just proximal to this. The distal article
articulâtes obliquely on the basal and bears 3 latéral spines, 3 terminal spines and a small spine, set
in front of 2 of the terminal spines. There are 2 spinules at the base of each spine. The basal article
of the endopodite has no armature but is very spinulose. The terminal article has 3 apical spines.
Articulating at the middle of the posterior face there is a single very long seta (.34 mm long in the spéci¬
men examined).
The second leg (fig. 2, b) is biramous with bimerous protopodite and rami. The protopodite,
exopodite and basal article of the endopodite are as in the first leg except that the basipodite lacks the
médial distal spine. The terminal article of the endopodite has 4 distal spines and 3 very long setae
articulating on the posterior ace. The most proximal end médial of these setae is .97 mm long. The
middle seta is .67 mm long, the latéral is .39 mm long.
The third leg (fig. 2, c) is biramous with bimerous protopodite and rami. It is just like the second
leg. The very long setae measure .91 mm, .53 mm, and .4 mm, respectively, from médial to latéral.
The fourth leg (fig. 2, d) is biramous with bimerous protopodite and rami and the protopodite
is much as in the preceding 2 pairs of legs. The basal article of the exopodite has a single spine at the
distal latéral corner. The terminal article bears 2 latéral spines, 3 terminai spines and a short spine,
articulating somewhat on the face. The basal article of the endopodite bears 1 extremely long seta
at the distal médial corner. This measures 1.12 mm long. The terminal article has 2 terminal spines
and 2 spiniform projections apically. Two long setae articulate on the posterior surface ; the more
médial measures .98 mm, the other is .84 mm long.
The fifth leg (figs 2, e, g) is greatly enlarged and swollen and reaches to the end of the elongate
genital-metasome complex. The rounded posterior margin bears a single seta. The 2 fifth legs overlap
dorsaily and each covers the whole side of the génital complex. The outer surface bears a general
covering of raised rings (fig. 2, f). Ventral to the dorsal surface of each leg the extruded ovisac is
accommodated in a dorsal concavity of the génital complex. There is a cuticular line ventrally, on
the body proper apparently delimiting the segment of the fifth legs. Just posterior to this line there
is on each side a complex of sclerotizations surrounding the oviducal aperture (fig. 2, h). There is
1 setule on a membrane at the margin of the aperture. The insémination pore is midventral just sligh¬
tly posterior to the level of the oviducal apertures and diverging séminal tubes pass from the pore toward
the apertures.
The caudal ramus (fig. 2, i) is shorter than the anal somite, and tapers slightly. There is a stiffe-
ned seta at the middle of the latéral margin, and 4 apical setae. The longest seta is about two-thirds
the length of the ramus.
The copepod, as coUected in the locations in British Columbia recorded above, is whitish but
light rosy pink ova and embryos give a distinctive coloration to the oviducts or to the ovisacs. The
pecific name indicates that most specimens observed in Europe were distinctly pink.
Détection of the paragnath is of substantial interest, as indicative that this structure can occur
among the Ascidicolidae. It has been known since the original description that a paragnath occurs in
lùiterognathus. In this form and in Ascidicola the structure is substantially of the same form as found
widespread among the Notodelphyidae. In notodelphyids with much modified mouthparts the para-
Source : MNHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
gnath often is still included. No such case has yet been reported for the more modîBed ascidicolids
and we bave not satisfactorily identified a paragnath in the wide sampling of représentatives of ail
the suhfamilies we hâve studied.
We call attention here to the specimens from the Antarctic we hâve listed above. We examined
tbese in an attempt to confirm the existence of modified spines in the first legs, as invoked by Lang
in his description of A. aculeoretusa. We failed to find any unusual development in our specimens and
accordingly cannot support establishment of a separate species for the Antarctic population.
We hâve not had opportunity to study the male, but there is information concerning it in the
literature. In the original description, Thorell included an account of what he considered to be a
male. Canu (1892) made the supposition that Thorell’s specimen was an immature female, as has
tumed out to be the case. Sabs (1921, p. 65-66, pl. 31) briefly characterized the male and figures the
habitus, the antennule, first and second legs and the caudal ramus. Monniot (1965, p. 158-159,
figs. 43, b, c, j. 1-n, q, s, t) treated the dimorphism of the species. Gage (1966) presented most interest-
ing information on the incidence of adult males and subadult stages.
The male is remarkable in that while being much smaller and more cyclopoid in habitus than
the female, it has the head appendages almost exactiy similar to those of the female. In this regard
it is distinctly different from ail the other ascidicolid généra for which the male is known. It is inte-
resting that in some of the notodelphyids {Pygodelphys spp., some Doropygus spp.) a similar correspon-
dence of the head appendages in the two sexes occurs.
The legs of the male Ascidicola differ strongly from those of the female. It could be roughly
said that they are entirely like the legs of some modified notodelphyids. There are discrepancies in the
figures of Sars and Monniot, as to the setation of the first 2 legs, but there is a possibility of a considé¬
rable degree of individual variation. We hâve found such variation in the legs of male notodelphyids.
The fifth and sixth legs and caudal rami are remarkably like those of a number of notodelphyids.
In the dimorphism and in the characters of the head appendages of both sexes and the legs of
the female, Ascidicola is a thoroughly distinctive copepod type. On the other hand, as we hâve pointed
out above, most of the basic features of the appendages in Ascidicola show remarkable conformities
to what might be considered prototypes in the Ascidicolidae in the wide sense, and often also in the
Notodelphyidae. The dimorphism is remarkable in the light of the knowledge now available on the
biology of the species (Gotto, 1957 ; Gage, 1966). The feeding process of the female has been tho¬
roughly described. The anatomy of the male présents distinct problems in attempts to reconcile the
feeding biology in the 2 sexes. Since the male has ail the mouthparts developed to the same degree as
in the female it seems reasonable to assume it feeds, and very likely, on food similar to that of the female.
However, the male thoracic appendages totally lack the exaggerated development of the setae which
is crucial to the highly characteristic feeding process of the female.
The species is now rather well known, as might be expected from its wide géographie range.
Its occurrence has doubtless been missed in many additional localities because of its particular biology.
Gotto (1957) and Gage (1966) présent many details of the biology, development and seasonal cycles
of this species in the British Isles, showing particularly a coordination of developmental stages in the
copepod with stages of development of the ascidian hosts. These ûndings serve well as a hint as
to the extremely interesting possibilities remaining in the study of the natural history of the ascicicole
copepods, individually and as an assemblage.
Styëlicola Liltzen, 1968
Styelicola Lützen, 1968, p. 97 (type species, by monotypy, S. bahusia Lützen, 1968).
The type species of the genus, discovered by Lützen, is shown by the discovery of our new
species to be a rather aberrant member of its genus, and of the subfamily. Our new species in preli-
minary superficiel examination was listed as another record of Ascidicola cf. rosea. Gloser inspection
and ultimately, dissection disclosed a very good conformity to the generic characters of Styelicola,
Source : MNHI\I, Paris
ASCIDICOLIDAE AND SUBFAMILIES
25
with a new, unsuspected set of very elaborate adaptations of the legs, but retaining an équivalent degree
of development of the mouthparts. Our new anatomical data lead us to emend the generic diagnosis
slightly.
Diagnosis :
Female : The body form is ascidicolin or short and with various tendencies to réduction in segmen¬
tation and development of the appendages. The antennules are short, compact, 7- or 8- segmented ;
the antennae are trimerous, and the terminal article bears a hooked seta or claw. The mouthparts
show reduced segmentation and ornamentation. The first 4 pairs of legs show distinctive adaptations,
either toward prehensile modification, or by réduction of segmentation and ornamentation. The
fifth pair of legs consist in part of an expanded structure which bears a free endopodite, this bearing
several setae.
The male remains unknown.
KEY TO SPECIES, BASED ON FEMALES
1. General body form trim, fusiform, forebody and hindbody of about equal length, liindbody with
slight but regular posterior taper. ligkti, new species
1'. Body short, thick, broad, approaching eruciform habit, forebody much exceeding hindbody..
bakusia Lützen, 1968
Styelicola bahusia Lützen, 1968
Styelicola bakusia Lützen, 1968, p. 97-102, figs. 1-2 (type locality, Skagerrack, Bohuslan coast, Swedcn,
in Styela alUintica (Van Name) and S. gelatinosa Traustedt).
Styelicola lighti new species
(Figures 4, 5, 6)
Type : Holotypic female, USNM (type locality, Amoy, China, from Harlmeyeria chinensis Tokioka,
1967 (identified by T. Tokioka), collected by S. F. Light).
Specimens examined : The holotype remains the only specimen so far discovered.
Female : The overall length of the single specimen, to the ends of the caudal rami is 3.4 mm. The
body (figs. 4, a, c, d) remarkably resembles that of Ascidicola rosea ; it is subcylindrical, blunt apically,
and with a slight posterior taper. The well-marked shield of the cephalosome is sharply delimited
dorsally. It covers the head complex and this includes the segment of the maxillipeds. Each of the
thoracic segments corresponding to legs 1-4 is well developed and strongly marked off. The segment
bearing the fifth legs is very complex. At its anterior margin it is slightly narr'ower than the segment
of the fourth legs. Laterally the unit (because it doubtless incorporâtes more than one segment) is
involved in the large expansions which are the basal portions of the fifth legs and cover the proximal
portions of the paired egg sacs. Posterior to the origins of the expansions the unit tapers to somewhat
more than half of its anterior width by a sériés of 2 or 3 successive constrictions. Because the fifth legs
clearly originate on this unit, which also supports the oviducal apertures, it is probably a fused complex
of the last 2 thoracic segments. An interesting and unusual corollary feature is the apparent lack of
the usual major body articulation between a forebody and the hindbody ; this arrangement is also
typical of Ascidicola rosea. Four long urosomal articles apparently comprise 3 true abdominals seg-
Source : A1NHN, Paris
Fio. 4. Slyelicola Ughti, new species, holotypic female : a, habitus, dorsal
d, habitus, latéral ; e, antennule ; f, antenna ; g, mandible ; h, maxiliule ;
in place, ventral vievr. Scales for a and c = 1.0 mm ; other scalcs = 0.1
view; b, egg sac; c, habitus, ventral;
i, maxitla ; j, maxilliped ; k, legs 1 to 3,
Source : MNHN, Paris
Fie. 5. — Styelicola lighti, new specie*. holotypic female : a, flrst leg ; b, second leg. anterior view ; c, second leg, poste-
rior view ; d, second exopod ; e, third leg, anterior : t, third leg, posterior ; g, fourth leg, anterior. Scales = O.I imn.
Source ; MNHN, Paris
Fig. 6. — Styelicola ligkti, new species, holotypic female : a, fourth leg, potterior view ; b, pediform projection-Afth
leg, in position on body, ventral view ; c, fifth leg -, d, endopod ot fiftb leg ; e, anal somite and caudal ramue.
Scalee = 0.1 mm.
ments and the anal somite. The anterior articles are rather similar, participating in the general body
taper. Each shows a constriction at about the posterior fifth. At the articulation of the anal somite
there is borne on it a curious pad, located dorsally (figs. 4, a, 6, e). This shows a remarkable corres-
pondence to the spinose pad of the abdomen of Ascidicola rosea but is dorsal rather than ventral. The
many spinose éléments arranged in a very regular pattern on the pad seem to difïer basically from those
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
of Ascidicoki. The short anal somite supports the caudal rami, to be described and figured below.
The anterior urosomal articles and the posterior portion of the fifth leg-genital complex are oriiainented
ventrally with regular rows of fine spinules. The elongate egg sacs are firmly attached to the oviducal
apertures, each sac with its anterior end shielded by the flap-like expansion of the pediform projection.
Each egg sac {fig. 4, b) is about 0.6 the length of the body and contains considerably over 200 eggs, these
more or less arranged in about 10, or perhaps more, longitudinal rows.
The antennulc (fig. 4, e) is stout and well sclerotized. It is composed of 7 articles; the arti¬
culation is somewhat weaker between articles 4-5 and 6-7. The setae are stiff and none is notably
long. They seem in general to be lacking in plumosity but this feature could not be clearly made
out owing to the State of our specimen. The présentation of the armature here must be regarded as
an approximation, giving some minimal expectations, at least, for the 7 articles (I-VII) : I — 2 setae ;
Il — 7 setae and a short round-tipped élément ; III — 3 setae and 2 short stout éléments, one marginal
and one on the surface of the article ; médial to this element a short row of spinules ; IV — 1 seta ;
V — 1 seta ; row of spinules ; VI — 2 setae, row of spinules ; VII — 8 setae of varions lengths made
out on this specimen.
The antenna (fig. 4, f) is S-segmented. The articles show the foUowing ratio of lengths, proximal
to distal : 1:1:2.5. The basal article bears a distal spiniform seta ; this element is stout and thick with
distal fine spinose ornamentation. The second article bears a marginal short, stout spiniform element
at about midway on its length. The terminal article bears proximally 2 short stout éléments inserted
close together near the margin ; there are 4 subapical éléments : a long setule and 3 short, rather spini-
forni éléments ; one of the latter takes origin on a stout tapered claw-like element broadly articulated
on the apex of the article.
The mandible (fig. 4, g) much resembles that of Ascidicola in general configuration. The basal
portion extends as a masticatory lamella. This is heavily sclerotized and the gnathal margin consists
of 4 subequal tooth-like lobes, each acute terminally. The margins of these lobes are irregularly serrate.
The palp appears to comprise a single element, although basally this is complicated by a large marginal
lobe. The lobe bears a long seta, and there are 6 more setae inserted rather regularly along the apex
and médial margin of the body of the palp.
The maxillule (fig. 4, h) is basically bipartite. A proximal element, conforming well to the
usual endite, bears 7 stout marginal spiniform setae. .An articulated palp, indicating by its lobing
that some of the hasic éléments of the generalized maxillule are possibly represented, bears 5 setae
on a médial lobe, 6 setae on a terminal lobe and 2 setae on a latéral lobe.
The maxilla (fig. 4, i) is tapered markedly and apparently is composed of 2 articles. The massive
basal article bears a process basally which furnishes insertion apically for 3 subequal stout setae.
Distally on the article 2 additional longer setae are inserted close together. The remainder of the appen-
dage forms an unsegmented apical piece bearing 6 graduated setae ; not ail of these are clearly articu¬
lated on the appendage.
The maxilliped (fig. 4, j) is fiat with marginal sclerotization. It bears one stout seta about
midway on the medial margin, 2 inserted together somewhat distal to this, and a single apical seta.
There are some rows of spinules on the surface of the appendage.
The 4 pairs of legs of the metasome form a sériés showing an unusual prehensile adaptation
(fig. 4, k). The first legs (fig. 5, a) are the least modified along this line, but are still far from generalized
appendages. The protopodites are massive with sclerotizations and surface ornamentations of fine
spinules. The eoxopodites are linked by a lamella, but there is no medial coxal seta. The massive
basipodites furnish insertions for the rami. Laterally each basipodite develops a prolonged base which
supports a very long, basally stout, medially directed seta. On the distal medial corner the basipodite
bears a short stout spine, as is usual among related copepods. The 2-segmented exopodite seems to be
a broadened flattened plate. The basal segment bears a stout flanged spine at its distolateral corner.
The distal article is about as massive as the basal, rounded apically. There are 3 large marginal spines
inserted in emarginations ; a distolateral slightly longer spine with 2 small accompanying spinules,
one inserted to either side of its base ; and a slender spine borne at the distal medial corner.
The endopodite is entirely distinctive. It is 2-segmented ; the basal article without spines
Source : Mt-JHN, Paris
30
PAUL L. ILLG AND PATRICIA L. DUDLEY
or setae. The second article is wider than long, with its liroad apex produced into 4 lobes, each furnish-
ed insertion for a short stout spine. Each of the 3 more latéral lobes has a characteristic ilange enve-
loping the base of the spine. There are some characteristic cuticular ornarnents on the surface of the
ramus.
The second leg (fig. 5, b, anterior view) althoiigh highly modified is readily secn to be dérivative
from the basic plan of the first leg. The protopodite remains massive with almost complété fusion
of coxopodite and basipodite. An exceptionally long seta is insertcd laterally on the portion corrcs*
ponding to the basipodite. The exopodite (fig. 5, d) takes a very complex articulation so that it can
swing medially and fold against the basipodite and endopodite (fig. 5, c, posterior view). The basal
segment of the 2'segmented exopodite furnishes the complex articulation of the ramus and its anato*
mically latéral surface bears a short stout spine which in the usual posture of the appendage is directed
distally. The entire apical portion of the appendage is converted into a fiat, curved, tapered claw
with very heavily sclerotized margins. When the ramus is flexed the apical claw is enveloped in a fold
formed in the endopodite. The endopodite remains bimerous, with the basal article broad and short.
It supports a distolateral spine. The posterior surface of the terminal article participâtes in a longi¬
tudinal fold of the entire ramus which reçoives the claw of the exopodite. The latéral lobe of the ter¬
minal article bears 2 short spines inserted more or less apically on the lobe.
The succeeding third (figs. 5, e, anterior view ; 5, f, posterior view) and fourth (figs 5, g, anterior
view ; 6, a, posterior view) legs show only slight déviations from the basic plan of the second legs.
The fifth legs somehow participate in the lamellate projections of the sixth thoracic segment
(figs. 6, b, c ; 4, a, c, d). The expansions, which serve as covers for the anterior part of the egg sacs,
show by the 2 marginal setae borne on each that there is probably an élément of the appendage incor-
porated, or alternatively fully forming the projection. The médial and latéral setae could possibly
be taken as indications of the représentation of parts of a coxopodite and a basipodite. There is no
very substantiel evidence for assuming an exopodite is involved. The free segment (figs. 6, c, d),
with its 6 setae could best be accounted for as a modified endopodite, as LUtzen concluded in the
case of S. bakusia. Four of the setae are very long, subequal, with some indications of ornamentation
on some of them. The second from the médial margin is accompanied by 2 subequal short setae inserted
near its base.
The caudal ramus (figs. 6, e, f) is somewhat flattened dorsoventrally, with sclerotized margins.
A relatively normal setal complément is accounted for by the 2 latéral and 4 apical éléments, but these
tend to be spiniform and shortened. The largest of the apical éléments is only slightly more than half
as long as the ramus.
Unfortunately, tins most interesting species is known only from a single specimen. The disco-
very of the male could well provide some very suggestive information relative to the morphological
sequence of adaptations in the Ascidicolidae.
SUBFAMILY BUPRORJNAE Thorell, 1859
Buproridae Thorell, 1859, p. 340, 358-359 ; 1860, p. 119, 139-141 ; 1862, p. 7, 14, 17, 61. — Gerstaecker,
1870-1871, p. 719. — Kossmann, 1874, p. 281. — Brady, 1878, vol. 1, p. 18, 21, 30, 146-147
(part). — Aurivillius, 1882b, p. 108,109 ; 1883, p. 104, 105. — Délia Valle, 1883, p. 252. — Auri-
villius, 1885a, vol. 4, p. 236. — Sars, 1921, p. 61. — Pesta, 1934, p. 8. — Lang, 1948, p. 6, 25,
26, 27. — Bocquet & Stock, 1961, p. 225. — Bresciani & Lützen, 1962, p. 373.
Buproriden Claus, 1875a, p. 351, 352 (part).
Ascidicolidae Canu, 1891a, p. 472, 475 (part) ; 1892, p. 107 (part).
Boporidae Bocquet & Stock, 1963, p. 293.
Notodclphyidac Claus, 1862, p. 102 (part).
Source : Mh}HI\I, Paris
.\SCIDlCOLIDAE AND SUBFAMILIIÎS 31
Silice this subfamily is monotypic, at least as far as including a single genus, ils charactcrs are
ihose of Buprorus, discussed below.
Buprohus Thorell, 1859
Buprorus Thorell, 1859, p. 340, 359 (type, bv monotypy, B. looeni Thorell, 1859) ; 1860, p. 119, 140 ;
1862, p. 7, 11, 13, 14, 61-63. — Claus,'l862, p. 102. — Gerstaecker, 1870-1871, p. 719. — Claus,
1872, p. 419. — Kossmann, 1874, p. 283, 289. — Claus, 1875, p. 351, 352. — Aurivillius, 1882a,
p. 63, pl. 5, fig. 13 ; 1882b, p. 109,110,113 ; 1883, p. 105,106, 109 ; 1885a, p. 236 ; 1885b, p. 282 ;
1886, p. 44. — Canu, 1891a, p. 475 : 1892, p. 28. — Hartmeyer, 1911, p. 1735. — Sars, 1921,
p. 61. — Schellenberg, 1922, p. 224. — Hansen, 1923, p. 23. — Wilson, 1932, p. 602. — Pesta,
1934, p. 8. — Neave, 1939, p. 502. — Sewell, 1949, p. 191, 192, 193 (part). — Bocquet & Stock,
1961, p. 225. —
nol Buprorus Sewell, 1949, p. 192, for Buprorus pranizoides = Anomopsyllus pranizoides Sara, 1921.
The taxonomie characters of the genus are remarkably consistent through the component species,
these being differentiated by rather minor features. The body form is very distinctive, with almost
complété réduction of the urosome. The incubatory cavity strongly suggests affinity with the Noto-
delphyidae, but we would interpret this as a convergent fealure and we feel that relationship with the
olher ascidicolids is more strongly indicated by the appendages. The antennule is distinctive for the
genus in configuration, but the ornamentation and the segmentation show great resemblance to Bolryllo-
pkilus. In B. nordgaardi the antennule is 7-segmented, with profuse setation, the exact details of which
are not available from Sabs’ présentation. The pattern is not at ail notodelphyid, but could serve as
an approximate model of the prototype of the ingrédient segments and setae of the whole sériés that
we are proposing to include in the Ascidicolidae. In the antenna there are more setae than occur in any
of the ascidicolids and, again, the segmentation and setation départs very definitely from the noto¬
delphyid pattern. AU the antennae in the whole sériés we consider as ascidicolids could dérivé from
an appendage with very much this basic anatomy, which could also very possibly be found to occur
among cyclopinids. The mandible, maxillule, and maxilla are ail reduced appendages. In ail cases
they are very like the corresponding appendages of Ascidicola, which are in turn a bit more generalized.
Among the ascidicolids probably the closest approaches to the most generalized représentatives of these
appendages occur in Botryllophilus. These, in turn could ail dérivé from the most basic condition in
the Notodelphyidae and very likely reflect a common ancestry with that family. The maxilliped
of Buprorus is so reduced as to offer very little information to comparative morphology. It is, however,
almost exactly like that of Ascidicola. Legs 1 to 4 strongly suggest those of either Ascidicola (in segmen¬
tation, not ornament) or Botryllophilus, but with some definite modifications from either condition.
Diagnosis :
The body form in the female is highly distinctive, with a fusion of the segments of the metasome,
extension throughout the metasome of the incubatory cavity, and with strong suppression of the uro¬
some, essentially eliminating the caudal rami. The antennules are flattened, of 3 to 7 segments, with
many setae, these distributed along the anterior and terminal margines. The antennae are trimerous,
non-prehensile. The mandible is reduced to a protopodite article with masticatory lamella and with
the palp reduced to a small setiferous lobe or a single seta. The maxillule is reduced, the protopodite
consisting of an article with a prominent endite with numerous marginal spines. The uniramous distal
article bears 4 or 5 éléments of armature. The maxilliped is bimerous or trimerous, the basal segment
much exceeding the others and with a single distal endite. The terminal article or articles bear a medial
hook and accompanying setae. The maxilliped is reduced to a simple lobe with 4 distal marginal
setae. The first to fourth legs hâve bimerous protopodites, bimerous exopodites and unimerous endo-
podites. Setation varies in the species. The fifth legs are expanded lobes with 4 to 5 terminal and
latéral setae.
The male is not known.
Source ; MNHN, Paris
32
PAUL L. ILLG AND PATRICIA L. DUDI.MY
KEY TO FEMALES OF BUPRORUS
1. Antennule 7-segmented. nordgaardi Sars, 1921
1". Antennule trimerous. 2
2. Urosome reduced, no setae or spines présent. looeni ThoreD, 1859
2'. Urosome reduced to anal somite tipped with three spines. caudatus, new species
Excluded species : Buprorus pranizoides Sewell, 1949, p. 192, for AnomopsyÜus pranizoides, Sars,
1921.
Büphorus loveni Thorell, 1859
Buprorus loveni Thorell, 1859, p. 342, 343, 358-359 (type locality, Swedish Coast in Ascidia merUula,
A. canina, A. aspersa) ; 1860, p. 19, 122, 123, 139-141 ; 1862, p. 7, 8, 61, 63-64, pl. 10, fig. 14. —
Gerstaecker, 1870-1871, p. 775, 776, 780, pl. xi, figs. 27-30. ~ Claus, 1872, p. 419. — Aurivilius,
1882a, p. 63, pl. 5, fig. 13 ; 1882b, p. 110, 113 ; 1883, p. 33,106,109, pl. 1, fig. 13 ; 1885a, p. 236 ;
1885b, p. 282 ; 1886, p. 44. — Hartmeyer, 1911, p. 1735. — Sars, 1921, p. 62-63, pl. 30. — Han¬
sen, 1923, p. 23-24. — Lang, 1948, p. 3. — Sewell, 1949, p. 191, 192, 193. — Gotto, 1960, p. 225.
— Bresciani & Lützen, 1962, p. 373.
Distribution : Swedish west coast, BohUslan, Gullmarsfjord, N. Finmark, W. Finmark, Norway, Vest-
manhavn, Faëroes, SmôrkuUen, Loken.
Hosts : (Names as recorded by the various authors ; we hâve not attempted to form a synonymy for
the ascidians) : Ascidia aspersa, A. canina, A. mentula, A. obliqua ; Ascidiella aspersa ; Phallusia
menlula, P. obliqua.
Buprorus nordgaardi Sars, 1921
Buprorus nordgaardi Sars, 1921, p. 80-81, pl. 37, fig. 1 (type locality, Trondhjem Fjord, in Amaroecium
sp.). — Sewell, 1949, p. 192. — Gotto, 1960, p. 225. — Bocquet & Stock, 1961, p. 225.
Distribution : Trondhjem Fjord, Norway.
Host ; Aplidium sp.
Buprorus caudatus new species
(Figure 7)
Types : Holotypic female, UNSM 92537 (type locality, off Southern California, Albatross Station 2972,
34®, 18', 30" N, 119®, 14" W, 1889, from Styela coriacea (Aider & Hancock) (= Styela hemicaespi-
tosa Ritter, cat. no. 5684) ; and paratypes below :
Specimens examined :
California : From Styela coriacea (Aider & Hancock) (= S. hemicaespitosa Ritter) off Southern
California, 34®, 18' 30" N, 119® 41' W, Albatross Station 2972, 1889, 4 adult females, holotype, one
paratype dissected, 2 undissected paratypes.
Description :
Female ; Overall length 1.1 mm. The habitus (fig. 7, a) is as previously figured for B. lovein
by Thorell and by Sars an ovoid body with a projecting head and ventral row of appendages. The
Source : MNHN, Paris
ASCiDICOLIDAE
SUBPAMILIES
33
Fig. 7. — Buprorus caudatus, new species, paratypic female ; a, habitus, latéral ; b, antennule and rostrum ; c, antenna ;
d. labrum (L), maodible (Md), and maxillule (Mxl) ; e, mandible ; f, maxillule ; g, tnaxilla ; h, maxilliped ; i, firat
ieg ; j, second leg ; k, third leg ; 1, fourth leg ; m, fifth leg, latéral ; n, urosome. Scales = 0.1 mm.
inciibatory cavity can be seen to extend throughout the metasome. The body régions are a cepha-
losome, bearing as its last appendages, the maxillipeds ; unsegmented metasome bearing 4 pairs of
legs ; and much modified and reduced urosome bearing the large fifth legs and terminating in a conical
anal somite tippéd with 4 spines and lacking caudal rami. AU the appendages are heavily sclerotized,
The rostrum (fig. 7, b) is a simple lobe, somewhat sclerotized.
The antennule (fig. 7, b) is trimerous, flattened, and with a single row of setae along its anterior
and terminal margins. The basal 2 articles are markedly wider than the terminal article. The middle
article is much the longest. The armature is as (ollows : 1-2 setae ; II-IO or 11 setae ; III-7 setae,
1 aesthetc.
3
Source : MNHN, Paris
34
PAUL L. ILLG AND PATRICIA L. DUDLEY
The antenna (fig. 7, c) is trimerous, with the basal article nearly equalling the combined lengths
of the terminal 2. The basal article has a single seta at the medial distal corner. The second article
bears 4 heavily sclerotized spines on the medial margin along the distal half. The terminal article is
modified in outline by a small medial terminal projection. On this there are inserted 2 heavily sclero¬
tized spines. At its base are 2 curved, stiffened seta. Laterally, at the apex, there is a stout spine
considerably longer than the article.
The labrum (fig. 7, d [L]) terminâtes in 3 lobes bearing fine setules.
The mandible (figs. 7, d [Md] ; 7, e) consista of a protopodite drawn out medially as a toothed
lamella. From the middle of the appendage there emerges from a pit a conspicuous seta which is the
only remnant of the palp.
The maxillule (fig. 7, f) is bimerous. The protopodite is drawn out medially into a toothed
endite, bearing 6 spiniform projections, possibly homologuous with the setae of other forma. The
distal article, possibly representing the palp, has a medial terminal lobe bearing 2 short, stout spines.
At the latéral distal corner there are 2 curved, sclerotized spines, longer than the medial spines.
The maxilla (fig. 7, g) is bimerous. The basal article is about 3 times longer than the distal
and about twice as wide. At the distal medial corner of this article there is an endite which terminâtes
in a spinulose, spiniform process. At the base of this process there is a small articulated spine. The
distal article is contracted terminally by 3 emarginations and terminâtes medially in a curved hook.
In the emarginations there are articulated spines, 1 in the first, 2 in the second, 1 in the third at the base
of the hook.
The maxilliped (fig. 7, h) is a unimerous tapered lobe. Four spines form a row beyoïid the middle
of the medial margin.
The first legs (fig. 7, i) are biramous, with bimerous protopodite, bimerous exopodite and uni¬
merous endopodite. The coxopodites are united by a band of cuticle which is not developed as strongly
as the usual intercoxal lamella. There is a small setule on the latéral margin of each basipodite, which
is trapezoidal in outline, so that the articulation of the exopodite is latéral. The basal article of the
exopodite has a seta at the distal latéral corner. The terminal article has 3 terminal spines, 2 medial
spines and 1 curved, stiffened seta, articulated on the posterior surface at the distal fourth. The endo¬
podite is short, about as long as the basal article of the exopodite. It bears 1 distal latéral spine, 2 api¬
cal spines, and 1 distal medial curved spine. A long, curved, stiffened seta articulâtes on the posterior
face near the articulation of the distal medial spine.
The second legs (fig. j) are biramous, with bimerous protopodite, bimerous exopodite and uni¬
merous endopodite. The protopodites and endopodites are as in leg 1. The basal article of the exopo¬
dite has a seta at the ffistal latéral corner. The second article bears 1 apical spine, 3 medial spines,
1 medial curved, long seta, a setule and a long curved seta, the latter 2 articulated on the distal poste¬
rior face.
The third legs (fig. 7, k) resemble the second exactly except for the exopodites. The basal article
of the exopodite bears a setule at the distal latéral corner. The distal article bears 3 terminal spines,
2 medial spines, a medial, long, curved seta, 1 spine and 1 long curved seta, the latter 2 articulating
on the distal posterior face.
The fourth legs (fig. 7, 1) are biramous, with bimerous protopodites, bimerous exopodites and
unimerous endopodite. The coxopodites are united by a relatively poorly developed lamella. On the
anterior surface the articulation between the coxopodite and the basipodite are like those of the third
leg, except the second article of the exopodite lacks the long curved medial seta.
The fifth leg (fig. 7, m) is a large, swollen lobe, actually larger than the other legs. Its armature
consista of 3 terminal spines and 2 latéral spines spaced beyond the middle. Caudal rami are lackirig
but the body terminâtes in a conical anal somite tipped with 4 spines (figs. 7, a, n). The anus is subter¬
minal.
In almost ail its features, this species conforms very well with the figures presented by Sars
for B. hveni, except that the fifth legs and the urosome are much more developed and offer a diagnostic
différence. Sars’ figures of the head appendages are somewhat generalized, so it is entirely possible that
restudy of the European form will offer substantial différences. In the legs the exopodites are shown
Source : Mt-JHN, Paris
ASCIDICOLlDAIî AND SUBFAMILIES
35
by Saks to hâve many more éléments of armature than are found on our species. Buprorus nord-
gaardi is markedly distinct from both species. The name of our species is derived from the fact of
substantially greater représentation of the urosome than in the other species.
SUBFAMILY ENTEROCOLINA Délia Valle, 1883
Famille de lernéens, Tribu des dichélestiens van Benedeii, 1860, p. 160 (part) (for Enterocola n. g.). —
Dichelesthiina Kossmann, 1874, p. 288*289 (part). — Family Holotmeta Kossman, 1874, p. 285,
288-289 (part).
Ergasilina Claus, 1875, p. 352 (part).
Buproriden, Claus, 1875, p. 350-351 (part). — Buproridae, Brady, 1878, v. 1, p. 18*19, 21, 30, 146-147
(part).
Ascidicolidae, Gerstaecker, 1870-1871, p. 719 (part). — Giard, 1888, p. 505 (part). — Canu, 1891a,
p. 472 (part) ; 1892, p. 186 (part). — T. Scott, 1901, p. 351-352 (part). — Caïman, 1908, p. 177,
182 (part). — Chatton Brément, 1909c, p. 223 (part). — Smith, 1909, p. 66 (part). — Chatton
& Brément, 1915, p. 143-144, 148 (part).
Ascidicolinae, Chatton & Brément, 1915, p. 143-144, 148 (part). — Schellenherg, 1922, p. 219-220
(part). — Brehni, 1927, p. 490 (part). — Barnard, 1955, p. 237 (part).
Enterocolidi Délia Valle, 1883, p. 252.
Enterocolidés, Canu, 1892, p. 107.
Enterocoliens, Canu, 1886, p. 373-374 ; 1892, p. 117. — Chatton & Harant, 1922c, p. 245, 249-251 ;
1924e, p. 417-418.
Enterocolidae, Sars, 1921, p. 73-74 (part). — Blakc, 1933, p. 226 (part). — Gray, 1933, p. 523 (part).
- Pesta, 1934, p. 8 (part). — Lang, 1948, p. 23, 25-27 (part). — Rose <& Vaissière, 1953, p. 91
(part). — Gotto, 1954, p. 659 (part). — Bresciani Lützen, 1962, p. 376 (part). — Monniot,
1965, p. 160 (part). — Dudley, 1966, p. 155 (part). — Gotto, 1966a, p. 193 (part). — Stock,
1967a, p. 9 (part). — Lützen, 1968, p. 101.
Enterocolinae, Chatton & Harant, 1924a, p. 352-353 ; 1924b, p. 360-363. — Gotto, 1962a, p. 541.
Notodelphyidae, Norman & Scott, 1906, p. 201 (part). — Barnard, 1955, p. 237 (part).
Chatton & Habant (1924b, p. 360-363) concludod that the three généra so thoroughly studied
by them, Enterocola van Beneden, Enterocolides Chatton & Harant, and Lequerrea Chatton & Harant,
shonid form the nucléus of a distinct subfamily, Enterocolinae, in the family .\scidicolidae. They
provided an excellent diagnosis and we are essentially adopting this, adding characters derived from
our comparative studies of previously known and new forms. We propose to exclude from this siih-
family, the généra mentioned tentatively by Chatton & Harant as possibly of cventual inclusion :
Zanclopus Caïman ; Enterognalhus Giesbrecht ; Baciropus Gravier ; Enlobius Dogiel ; and VerUriculina
Bassett-Smith. They based their définition on the females of the généra and pointed out that the
characters of males must eventually be considered, but were not availalde at the time. We must also
acknowledge this necessity, but hâve little additional information concerning males, so the diagnosis
must still rest on characters of females. We hâve a few comments on males below (see p. 46).
The enterocolin body is eriiciform in the female but there is no exactly diagnostic detail déri¬
vable from the general habitus. Other ascidicolids and sonie other cyclopoids hâve the same approxi-
mate construction.
The cephalosome is broad, approaching semicirciilar outline, the convexity anterior and dorsal.
The cephalosome lacks pleural folds, apical plaque or rostrum. There are 4 or less pairs of cephalosomal
Source : MtJHN, Paris
36
PAUL L. ILI.G AND PATRICIA L. DUDLEY
appendages, the mandibles and maxilliped always are lacking. The labrum is présent, not particu-
larly salient and characteristically displaced anteriorly. A very consistent feature of the labrum is
the occurrence of 2 spinulose setae at the latéral corners.
The metasome may hâve the 5 component thoracic segments (which must commence with the
second segment ; the thoracic segment corresponding with tlie missing maxillipeds is apart of the head
complex) clearly demarcated, but there are considérable modifications and fusions. The sixth thoracic
segment is frequently coalesced with the fifth, and may fiirtber participate in a fusion with the urosome.
There may be paired pterostegites on any of the segments and the pediform processes of the last segment
are probably in part homologous.
The urosome is shorter than the metasome, stumpy, and its outward appearance may vary from
segmented to coalesced. The caudal rami are reduced, usually not clearly articulated with the urosome,
or may be totally lacking.
The antennule is reduced, tending to indistinct segmentation and sparse setation ; in Lequerrea
the appendage is possibly absent (alteratively, it is présent and the antenna is lacking).
The antenna is long and often much expanded. It is uniramous and mostly clearly bimerous.
There are no indications of prehensile fonction in the female and the setae tend to be reduced in number
and concentrated apically. Copepodids and the male of at least Enterocola fulgens hâve the antenna
prehensile.
The mandible is absent in ail copepodid stages.
The maxillule is massive and articulâtes on the head in a very coraplicated way. It appears
to be composed of a base and a palp but development and comparative morphology indicate the appen¬
dage may be biramous. The protopodite and presumed endopodite are fused to form the lamelliform,
heavily sclerotized base, which is produced medially with a lobed apex. There is an articulated seta
inserted on the surface. The free article is a setigerous lamella articulated on the base ; the apex of
this is entire to bilobed, with 5 or 6 setae, these variously grouped on the lobes.
The maxilla is uniramous, heavily sclerotized and with the apex produced medially as a hook-
process. There is always a complicated articulation of the base of the hook with the terminal part of
the appendage suggesting the composition is bimerous. There is a spinulose endite distal on the basal
segment, and a seta on the ventral margin of the hook.
The maxilliped is absent.
There are 4 pairs of subequidistantly spaced legs. These are subequal, always obscurely arti¬
culated on the body, and biramous. To a considérable degree the patterns of structure and oramenta-
tion of the legs are generic characters. The protopodites tend to articulate obscurely on the body.
The endopodites vary from unimerous to bimerous, setiferous to not so, the setae when présent apical
and not exceeding 2 in number. The exopodites are unimerous to bimerous, of a characteristic outline,
not bearing proper setae, but with a variety of curiously developed spine-like éléments.
The sixth thoracic segment always bears pediform projections which may be setiforous or not.
When présent, the setae are minute. The projections tend to be larger than any of the pterostegites
when these are présent and are always salient features in the general aspect of the animal.
KEY TO GENERA, BASED ON FEMALES
1. Thrce pairs of cephalic appendages ; antennules or antennae missing. Lequerrea (p. 37)
1'. Four pairs of cephalic appendages; antennules and antennae both présent. 2
2. Endopodites 1 to 4 without setae. Enterocolides (p. 42)
2'. Endopodites l to 4 with 2 setae, terminal or subterminal. Enterocola (p. 45)
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIF.S
37
Enterocolinae sp. incerlae sedis
Enterocolidae indet., Monniot, 1965, p. 160 (from Banyuls-sur-Mer, Golfe du Lion, France, from
Cratostigma graf>ellophila).
Lequerrea Chatton & Harant, 1924
Lequerrea Chatton & Harant, 1924a, p. 349 (Type species, by monotypy, L. perezi Chatton & Harant,
1924) ; 1924b, p. 362-363. — Wilson, 1932, p. 601. — Neave, 1939, p. 923. — Gotto, 1960, p. 226.
In adding a new species we extend somewhat the diagnosis of the authors of the genus. As
pointed out by Chatton & Harant, most of the appendages and the general features of the habitus
conform with the diagnostic characters of the subfamily Enterocolinae.
The general aspect is much like that of species of Enterocola, but the présent form is a very
large copepod. The size may be a reflection of the occurrence in solitary ascidians, both known species
so far found to occur in species of Polycarpa.
The urosome is markedly shorter than the remainder of the body, unsegmented, with the anus
dorsal, but at the end of the body.
Either the antennule or the antenna is missing, and the exact détermination will hâve to await
evidence from development. By comparison with the sériés of species of enterocolins, we tend to
concur with the surmise of the authors that the antennule is missing. There are 5 setae on the exopo-
dite of the maxillule, in contrast to the 6 setae usual in species of Enterocola and Enierocolides.
The 6rst to fourth legs are ail biramous, and it is characteristic, as in Enterocoîides, that the
endopodite lacks any articulated éléments of armature. Proportions of the rami are not distinctive,
since the endopodite may be longer or shorter than the exopodite. The pediform projections of the
sixth thoracic segment lack setae, but some species of Enierocola correspond in this feature.
The anatomical characters not specifically mentioned above can be assumed, as far as the 2 species
now known are concerned, to conform to the general features of the Enterocolinae.
The 2 species can be readily distinguished by the relative proportions of the rami of the legs.
In L. perezi, the exopodite slightiy exceeds the endopodite ; in L. canui, the endopodite greatly exceeds
the exopodite.
Lequerrea perezi Chatton & Harant, 1924
(Figure 8)
Lequerrea perezi Chatton & Harant, 1924a, p. 347, 349-352, fig. 1 (type locality, La Rade de Brest,
France, in Polycarpa sp., from intestine) ; 1924b, p. 363. — Harant, 1931, p. 371. — Gotto, 1960,
p. 226 .
Distribution : Atlantic coast of France ; Plymouth, England.
Host : Polycarpa sp., probably restricted to Polycarpa gracilis Heller.
Specimens examined :
Host undetermined : Brest, France, Chatton number 263, (possibly holotypic specimen).
1 female.
Host undetermined, probably Polycarpa sp.,;
Duke Rock, Plymouth Bay, Plymouth, England, 6 fathoms., September 8,1958, P. L. Dudley, 1 female.
From Polycarpa gracilis Heller :
Duke Rock, Plymouth, England, 1 female.
Source : MNHN, Paris
Fie. 8. — Lequerrea perezi Chatton & Harant, femate : a, habitua, ventral view ; b, habitus, latéral ; c,
tral ; d, antenna (?) ; e, labnim ; t, palp oî maxillule ; g. maxitlule, ventral ; h, maxilla, posterior ;
rior ; j, flrst leg ; k, third leg ; 1, caudal ramus. Scales for a, b, c, = 1.0 mm ; otber soales = 0.1
, urosome, ven-
i, maxilla, aote-
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Description :
Female (figs. 8, a-1), supplementîng the original description ;
Body (figs. 8, a, b), 3.17 mm total length, measured from anterior margin to the end of the
caudal rami. The body is somewhat sclerotized, relatively slender, with segmentation indicated ven-
trally by constrictions. Dorsally there are paired pterostegites on the second through fifth thoracic
segments. This feature départs from the statement of Chatton and Harant to the effect that there
are no dorsal duplicatures, except as involved in the fifth pedigerous projections. In frontal view
the cephalosome and thoracic segments are of about equal width, contrasting to the somewhat narro-
wer, tapering urosome. On the second thoracic (first leg-bearing) segment there is a sclerotized pro¬
jection anterior on the ventral midline, projecting somewhat anteriorly. The fifth pedigerous segment
is expanded laterally into conspicuous curved lamellae, unarticulated, and continuons ventrally as an
apron-like sheath covering the anterior part of the urosome (fig. 8, c). The insémination pore is covered
by this sheath. The lamellae bear no setae.
We use the orientation of an en face préparation to refer to the features of the head appendages.
Chatton & Harant considered that most likely the antennules were absent. We concur
in this conjecture, but developmental evidence will be necessary to provide definitive information.
The antenna (?) (fig. 8, d) is rather elongate but there is no clear indication of ingrédient segmen¬
tation. There is some feeble sclerotization and the setation is weak and apparently reduced.
The labrum (fig. 8, e) as we illustrate it, corresponds to the original description.
The maxillule (figs. 8, f, g) and the maxilla (figs. 8, h, i) correspond to the original description,
and we herewith présent the first illustrations of them.
We find the legs as described in the original description. The first (fig. 8, j) and second are
similar. The third (fig. 8, k) and fourth differ somewhat from the former and are similar to each other,
in bearing a long process with an articulated spine on the latéral margin of the basal segment of the
exopodite. In a similar locus on the first 2 legs there is a minute spinule. The caudal rami (fig. 8, 1)
were described generally in the original description but on our specimen each bears 2 terminal pro¬
jections.
The remarkably précisé description of the original authors leaves us in little doubt of the identity
of our specimens. We hâve been able to furnish some supplementary illustration and the distribution
is slightly extended by our records. For the first time a spécifie identification for the host can be
provided, namely Polycarpa gracilis, which is an interesting discovery in view of our procuring speci¬
mens of a new species of Lequerrea, described below, from Polycarpa rustica from approximately the
same locality. In our specimens the body was yellow and the embryos in the ovisacs were bright
orange.
Lequerrea canui, new species
(Figures 9, 10)
Types : Holotypic female (type locality, Salcombe, near Plymouth, England, intertidal, from Poly¬
carpa ruêlica (Linnaeus), August, 1958, paratypes, specimens listed below.
Specimens examined :
From Polycarpa rustica (Linnaeus) : Salcombe near Plymouth, England, intertidal, August,
1958, holotypic female and 1 other female.
Duke Rock, Plymouth Bay, England, 6 fathoms, September 9, 1958, 1 female.
Duke Rock, Plymouth Bay, England, 5-6 fathoms, September 12, 1958, 1 female.
Description :
Female (figs. 9, a-k, 10, a-e) :
Body (fig. 9, a) 5.78 mm total length, measured from anterior margin to the end of the caudal
rami of one specimen. The shape is eruciform, somewhat curved, with the segmentation obscured,
Source ; MNHN, Paris
Fig. 9. — Lequerrta «anui, new species, paratypic female : a, habitus, latéral view ; b, urosome and pediform projec*
tion*, ventral ; o, urosome and pediform projections, dorsal ; d, antenna (?) ; e, labnun ; f, postoral sclerotizations ;
g, maxillule ; h, basal portion of maxillule ; i, palp ot maxillule ; j, maxilla, posterior ; k, maxilla, anterior. Scalei
for a, b, c — 1.0 mm ; other acales = O.t mm.
Source ; Paris
ASCIDICOLIDAE AND SUBFAMILIES
41
indicated only by slight indentations. There are no apparent pterostegites. The body is more swollen
than in L. perezi, and as in the latter the third legs are separated so each is displaced somewhat dorsally,
the fourth legs are markedly latéral. The pediform projections perhaps corresponding to fifth legs
are markedly smaller than those of L. perezi. They are appressed to the proximal portion of the ovisacs
which are elongate and curved. The urosome (figs. 9, b, c) as in L. perezi, is unsegmented, and there is
no delimiting articulative line ventrally, although a dorsal articulation is présent. The anus is dorsal
near the end of the urosome.
The insémination pore is exposed on the ventral anterior surface of the urosome, since there
is no apron-like sheath, as in L. perezi. Diverging séminal canals pass from the pore dorsolaterally
to the vicinity of the oviducal apertures.
We use the orientation of an en face préparation to refer to thé features of the head appendages.
There is no rostrum and apparently the antennule is absent.
The antenna (?) (fig. 9, d) is elongate, with indication that there are 2 ingrédient segments, since
a strong sclerotization on the ventral face at the distal third perhaps represents an articulation, which
is incomplète dorsally. There are no setae proximal to the sclerotization. Terminally and medially
there are 10 short setae surrounding the distal margin. The dorsal face bears 6 rows of fine spinules
at about the distal third.
The labrum (fig. 9, e), as in L. perezi is displaced far anteriorly on the cephalosome to a position
at the bases of the antennae. As in the congener there is a terminal piece which is triangular and
somewhat denticulate, but without elaborate tooth-like structures. Rows of fine spinules ornament
the surface of this piece. On the anterior surface of the labrum there are médial rows of somewhat
coarser spinules. The usual latéral terminal setae of the labrum are covered with coarse spinules.
Posterior to the mouth on the surface of the cephalosome is a complex of sclerotization, includlng
raised médial lobes (fig. 9, f). This is emarginated by the heavy sclerotized ridges surrounding the
bases of the mouthparts. At the anterior latéral margin of the central sclerotization there is on each
side a lobe bearing a circular patch of heavy spinules.
The maxillule (figs. 9, g, h, i) is very heavily sclerotized, and there is evidently a fusion of the usual
éléments with the protopodite. The palp portion is also probably incompletely articulated, but it is
Source : MNHN, Paris
42
PAUL L. ILLG AND PATRICIA L. DUDLEY
represented by a flattened lobe-like element, which projects ventraUy over lhe remainder of the appen-
dage. It bears 5 marginal setae, in groupa of 2 and 3. Ail the setae are spinulose and there are patches
of spinules on the aurface at the baacs of the aetac. The basal portion is massive and heavily sclerotized.
Apically it is somewhat bilobed. On the anterior face there is inserted a very atrong aeta covered with
coarse apinules.
The maxilla (figs. 9, j, k) ia apparently Irimerous and very heavily sclerotized. The basal
segment is massive, somewhat truncate and with a distal médial protubérance that possibly représenta
one of the endites. This protubérance is dome-like and terminâtes in a spinulose thuinb-like process.
The articulation of the second probable segment is very complicated. On one surface it is very évident,
cutting directly across the appendage. The corresponding continuation on the opposite surface appears
to be obscured by a distal greatly extended expansion of the basal segment. There is no seta or déri¬
vative on the second segment. The terminal segment is triangular in outline, but is actually antero-
posteriorly flattened, and comprises 2 subequal appressed lobes. The lobes bear patches of spinules.
A seta, with a somewhat expanded base, is set on an emargination at about the basal fourth of the exter-
nal margin. A circulât sclerotization on the basal segment is possibly représentative of the aperture
of a glandular duct.
The first to fourth thoracic legs (figs. 10, a-d) are ail biramous, with lhe endopodites consistently
longer than the exopodites ; the reverse is the case in L. perezi. The articulations of the legs with the
body are not dclimited. The exopodites are distinctly articulated with the protopodites, but the arti¬
culations of the endopodites are obscure. Each protopodite bears a seta at tbe latéral distal corner,
a feature lacking in L. perezi. There are patches of fine spinules on the anterior surface of each proto¬
podite, and a row of 3 or 4 heavy spinules is found between the exopodite and endopodite on each leg.
Internally there are complex sclerotized skeletal pièces which serve as attachments for muscles. The
exopodites are very heavily sclerotized, short, conical, trimerous. In the first and second legs (figs. 10,
a, b) there is no latéral armature and the terminal element is regularly conical. In the third and fourth
legs (figs. 10, c. d), the basal segment is drawn out laterally as an elongate, unarticulated, spinose
process. The terminal element is modified in outline from a regular cône to a dôme sharply constricted
lerminally to a pointed apex. There is no surface ornamentation on the exopodites. The endopodites
are from 1 1/2 to 2 times longer than the exopodites and are digitiform, lacking articulations and arma¬
ture. There is a faint indentation on the posterior surface of each leg between the protopodite and the
endopodite. The anterior surfaces of the endopodites bear oval patches of fine spinules, and there are
a few heavier spinules on these surfaces near the latéral margins.
The pediform processes of the sixth thoracic segment are unarmed (figs. 9, b, c) and it is possible
that fifth legs are lacking. The processes cover the oviducal apertures which hâve accompanying
complex sclerotizations and are heavily muscularized.
The caudal ramus (fig. 10, e) is elongate, conical and bears a single terminal spiniform projection.
The surface is ornamented with patches of fine spinules and scattered, isolated, heavier spinules. This
ramus differs markedly from the ellipsoidal ramus of L. perezi.
Two of the 3 females collected were ovigerous. The animais were removed from the gut of the
host in each instance. The ova in the oviducts were purplish pink, the embryos in the ovisacs had a
mixture of orange and pink yolk. No males hâve been found.
Entbrocolides Chatton & Harant, 1922
Enterocolides Chatton & Harant, 1922c, p. 246-252 (type species, by monotypy, E. ecaudalus Chatton &
Harant, 1922} ; 1924a, p. 353. — Wilson, 1932, p. 598. — Neave, 1939, p. 241.
The characters of the one species of this genus serve also to furnish the generic diagnosis.
Source : MNHN, Paris
ASCICICOLIDAK AND SUBFAMILIliS
43
IOntekocoi.ides ecavdatvs Ciiatton & Harant, 1922
{Figure 11)
Eiüeiocolides pcaudatiis Chatlon & Harant, 1922e, p. 246-252, fig. 1 (type locaiity, Port Vendres, Golfe
du Lion, France, froni “ un Distomidc l)lanc ” [siibsequcntly idenlified as “ Didemnopsis crassa ”
Daumezon] ; Ciiatton & Harant, 1924a. p. 353) : 19241), p. 363. — Harant, 1931, p. 137. — Gotto,
1957, p. 268 ; 1960, p. 226.
Distribution : Golfe du I ion, France.
Host : Didemnopsis crassa, now known as Trididemnum inarmalum.
Specimens exaniiiied :
From D. inannata Drasclie :
“ distomidc blanc, giob. ” (Tunicate identified by M. Harant).
Ncar Port Vendres, ncar Banyuls-sur-Mer, France, Noveinber 11, 1910, Chatton number 151,
1 female. (Topotypic specimen, possibly holotype.)
From “ Distaplia sp. ” :
Near Port Vendres, ncar Banyuls-siir-Mer, France, 50-100 in, January 10, 1912, Ciiatton nunibor
58, 8 females.
We ofTer what we consider to bc pertinent supplementary information to the description of
Chatton & Harant and illustrate our specimens for comparison. The specimen, an adult female,
measured 1.42 mm, overall length, as compared with the range of 1.5 to 1.7 mm of the authors. The
proportions of our specimen, cephalosome : metasome : urosorne, are 3 : 14 : 4, as compared with the
piiblished ratio 1 : 5, 2.
The habitus (figs. 11 a, b) corresponds well to the figure and description of the authors. The
only additional feature shown by us is the insémination pore on the urosorne, with one of the diverging
séminal tubes, leading to the oviducal aperture under the pediform process, called the “ oostegite ”
by the authors. The antennule (fig. 11, c) agréés very well with the original with the exception that
basal rows of spinules were not seen, but we saw additional scattered spinules on the surface of the basal
article. The antenna (fig. 11, d) agréés well with the illustration but is uniincrous, contrary to the
statement of the authors that it is bimerous . The labrum (fig. 11, e) not figured by the authors, but
described, is a trapezoidal plate bearing rows of spinules on the ventral surface and a spinulosc seta
at cach distolateral corner. The maxillule (figs. 11, f, g) is bilobed. In the specimen we examined the
palp portion has only 4 terminal setac ; a knob in the center may indicate a vestigial seta. The authors
State the “ exopodite ” bears 3 setae on the distal margin of the external lobe and 1 on the external
margin of the external lobe and 2 on an internai lobe. The basal élément is massive, strongly sclero-
tized, bifid apically, and with a seta and a setule on the anterior margin. The maxilla, not previously
illustrated (fig. 11, h) is bimerous, with the basal article much longer and wider than the terminal.
An endite bearing a distinctly and articulated terminal claw protrudes medially from the distal médial
corner of the basal article. Tbe terminal article is large, claw-shaped, and extends parallel with the
endite but somewhat exceeds it. There is a small seta on the anterior face of the terminal claw and no
other surface ornamentation on the appendage. The 4 pairs of legs are biramous, each with a large
base carrying row's of spinules and with a single seta at the distal latéral corner. In ail the legs (figs. 11,
i, j, k, 1) the conical endopodite is unarmed and bears rows of spinules. The exopodite of the first leg
(fig. 11, i) is unimerous, has a patch of spinules at the middle of the latéral margin, a small hook-like
projection of the cuticle at the distal foiirth and is pointed apically. The exopodite of the second leg
(fig. 11, j), also unimerous, has spinules at the proximal fifth, a patch of spinules approximately at the
middle on a protubérance, a spinule at tbe distal fourth, and an apical cuticular minute claw accom-
panied by a small spine. In the third leg (fig. 11, k) the exopodite strongly shows the basic distinctive
structure characteristic of the entire subfamily. The ramus is a very heavily cuticularized, curved
stylet ; it is specifically distinctive in bearing a patch of spinules on a projection at the middle of the
Source ; MhiHN, Paris
44
PAÜL L. ILLO AND PATRICIA L. DUDLKY
Fig. 11. — Enlerocolides ecaudatué Chatton & Harant, lemale ; a, habitus, dorsal view ; b, habitus, latéral ; c, anten*
nule ; d, antenna ; e, lahrum ; f, palp ot maxülule ; g, médial process of basal portion ot maxillule ; b, maxilla ;
i, firat leg; j, second leg ; k, third leg; 1, lourth leg ; m, pediform projection. Scales = 0.1 mm.
latéral margin. The exopodite of the fourth leg (fig. 11, 1) is very similar to that of the first leg. The
pediform process (fig. 11, m) is a cupped plate with a curved terminal outline, so that it is somewhat
longer than broad. The authors show a single seta at the middle of the margin. We find 2 setules,
the more dorsal somewhat longer than the other.
The species has not been reported since the time of Chatton and his colleagues. AH the spéci¬
mens available for our study were from the Chatton collections.
There is some question about the host, but the authors seem to hâve settled on “ Didemnopsis
crassa ” which is now known as Trididemnum iTiarmatum von Drasche.
Source ; Mt-JHN, Paris
ASCIDICOLIOAE AND SUBFAMILIES
45
Enterocola van Beneden, 1860
Enlerocola van Beneden, 1860, p. 155-160 (type species, by monotypy, E. fulgens van Beneden, 1860).
— Gerstaecker, 1870-1871, p. 719. — Kossmann, 1874, p. 288-289. — Clans, 1875, p. 351-352.
— Brady, 1878, p. 21, 30, 147 (part). — Aurivillius, 1882a, p. 8, 92, 96 ; 1882b, p. 92 ; 1883,
p. 88. — Délia Valle, 1883, p. 245-247, 251. — Canu, 1886, p. 309, 311. — Giard, 1888, p. 505.
— Canu. 1891a, p. 468-475 ; 1892, p. 25, 29, 33, 39, 48, 50, 51. 54, 58, 66, 87,88, 90, 91,92, 93,
108, 117, 118, 119, 127, 130, 131, 133, 209, 213-218. — T. Scott, 1900, p. 386 ; 1901a, p. 352.
— Norman & Scott, 1906, p. 203. — T. Scott, 1907, p. 368-369. — Caïman, 1908, p. 182. — Chat-
ton & Brément, 1909c, p. 223-225. — Smith, 1909, p. 67. — Chatton & Brément, 1915, p. 144,
145, 148, 149, 153, 154. — Sars, 1921, p, 76-77. — Chatton & Harant, 1922a, p. 148-149 ; 1922c,
p. 245, 249-251 ; 1924a, p. 348, 349, 353-354. — Schellenberg, 1922, p. 287-288. — Chatton &
Harant, 1924b, p. 361, 362, 363. — Brehm, 1927, p. 490. — Wilson, 1932, p. 599, 600. — Neave,
1939, p. 241. — Barnard, 1955, p. 237, 241. — Gotto, 1960, p. 216 ; 1962, p. 541 ; 1966a, p. 193.
— Dudley, 1966, p. 155, 157, 159, 160.
Enlorocola Norman, 1868, p. 300 (part).
Intercola Hesse, 1862, p. 349, 354 ; 1865, p. 355.
non Enterocola Norman, 1868, p. 300.
non Enterocola Brady, 1878, p. 147. — T. Scott, 1892a, p. 301 ; 1892b, p. 203-205. — T. Scott and A.,
1895, p. 359-360.
Biocryptus g. n. Hesse, 1865, p. 242-247, 256 (type species not designated). — Canu, 1892, 216 (uncer-
tain species ; actually much resemblance to Enterocola). — Chatton & Harant, 1924b, p. 363
{Enlerocolinae, but unidentiBable as to genus or species). — Neave, 1939, p. 430.
When Chatton & Harant (1924, p. 361-362) formulated their subfamily Enlerocolinae, they
already had so many species of Enterocola to account for that their generic définition is a very general
one, with few particulars. Since that time still more species hâve been added. In order to obtain
first-hand information about the male in the genus, we hâve studied sorae collections made from com-
pound ascidians at Plymouth, England. We hâve found an amount of variation that leaves a definite
question in our minds as to just where species limits lie in the sériés of Enterocola forms coming from
closely related compound ascidians. To avoid designating our material as host forms, which we would
not feel able to do in the absence of substantial suites of specimens for comparative study, we are, with
definite misgiving, designating new species from our material. This expérience of ours points up to
us the close-knit nature of the relationship of the bulk of species so far described in the genus. We
find the major discontinuity in the group to be that séparating out E. büamellata Sars, E. setifera Hansen,
and E. laticeps, described below, and we must point out that the State of knowledge concerning
Sars’ and Hansen’s species is unsatisfactory.
As a resuit of Canu’s excellent présentation on development (1892), the description of additional
species of Enterocola, by Chatton and his associâtes hâve employed an appropriate terminology for
the head appendages, which procedure has been followed by other authors right to the présent (c.f.
Quille, 1964).
The body is eruciform, with 3 distinct régions. The cephalosome forms a head, and the maxil-
lules and prominent maxillae tend to delimit the région ventrally. In the metasome the component
segments are somewhat variably delimited and frequently feature dorsal duplicatures, termed pteroste-
gites by Chatton and his associâtes. These may not occur at ail or may be borne only on some of the
segments. The articulation of the legs with their body segments tends to considérable suppression.
In most species there are lamelliform or mammiform processes between the protopodites of some or
all of the pairs of legs. These are absent in some species. The sixth thoracic segment is characterised
by enlarged pediform projections which envelop much of the dorsal and latéral portions of the urosome.
Source : MttHN, Paris
46
PAUL L. ILLG AND PATRICIA L. DUDI.IiY
These usually are unarmed, but in some cases are setiferous, with at most 2 minute sctae. The caudal
rami arc soniewhat variable in proportions and extent of articulation with the urosome ; they are not
setiferous. The antennule varies froin unsegmented to vaguely bimerous ; Sars States that in E. bila-
meliala there are 4 segments. Hansen shows a multiarticulate appendage. The setation is reduced.
The antcnna is vaguely to clearly bimerous, with the second segment equa! to or much longer than the
basai. The second segment is fialtened, wide and always bears at least 6 setae, these of varying pro¬
portions. The lahrum is always prominent, its surface variously ornamented with spinules, and there
is characteristically at the postérolatéral corners a pair of spinulose setae. These are absent in E. mega-
/oi-a, according to the original description, and the situation is not known for E. seûfera or E. bilamdlata.
Some early statements that mandibular vestiges were présent in species of Enterocola were possibly
based on misidentification of these labral structures. Ciiatton and Harant did point eut (1924a,
p. 351) that if mandibular vestiges occur this would hâve to be verified by study of development. We
feel, from our own study of ascidicolids and reexamination of Canu’s findings, that there are no such
mandibular vestiges after the first or second copepodid and that the conspicuous structures are involved
with the labrum.
The maxillule is characteristic, essentially on a subfamilial pattern, remaining bilobed but
with much suppression of segmentation. The lamellate palp portion tends to be lobed, with 6 setae
distributed on the lobes and on the external margin. Enterocola bilamellata is reported to hâve only
5 such setae.
The maxilla is bimerous and heavily sclerotized. The basal segment supports a medially
extended lobe which may appear even to form an articulation. The distal segment is siibacute at the
apex, with a hook-or claw-like aspect. There may be 1 or more small éléments of ornamentation
set basal to the distalmost hook-process.
There is no maxilliped.
The first to fourth legs are reduced, biramous. There are no setae on the protopodites. The
exopodites tend to hâve a characteristic termination, this usually with a demarcated apex, often of
very distinctive outline, consisting of a dôme with an emerging tip, particularly on legs 1, 2 and 4.
Very consistently the third leg will hâve the exopodite with the margin uninterrupted and forming
in outline a stylet-like aspect. The endopodites may be clearly unsegmented ranging to obscurely
bimerous and always bear at least 2 terminal setae. For ail species for which the host relation is record-
ed, the copepod occurs in the stomach or nearby in the alimentary canal, and hosts previously recorded
were compound ascidians. One new species, described below, p. 91, is froin a solitary ascidian and
furnishes a hint that such problematical species as E. setifera and E. bilamellat-a possibly also corne from
solitary ascidian hosts. Most species so far known corne from hosts of the family Polyclinidae ; 2 are
known from the Didcmnidae.
Tue Male of Enterocola
Canu (1892, p. 216, 218, pl. 18, figs. 3-8), described the male of Enterocola fulg ens and we hâve
been able to make out some features of the male fifth copepodid in our material of E. pterophora. With
this basis we can présent a short sketch of the diagnostic features.
The body is cyclopoid, the division into forebody and urosome well demarcated. The cepha-
losome of the adult male, as described by Canu, is notable in the formation of well-developed pleural
folds. The ventral portions of these folds are covered with fine hairs, disposed in regular sériés. The
cephalosome bears a rostrum and 4 pairs of appendages. The mandibies and maxillipeds are absent.
There is also a distinctive modification of ail the appendages. The metasome consists of 4 segments,
each bearing a pair of biramous swimming legs. The urosome is 6-segmented, with the first segment
bearing rudimentary fifth legs and the second with indications of sixlh legs. Paired spermatophoral
sacs lie in the second urosomal segment. The caudal rami bear elongate terminal setae.
The antennules are 7- or 8-segmented. The setation is well developed and the terminal article
bears an aesthete as well as several setae.
Source : MNHN, Paris
ASCIDICOLIDAE AND SDBFAMILIES
47
The antemia is trimerous and bears an articulated terminal prehensile hook, accompanied by
several setae. The basal article bears a single distal outer sela.
We hâve no positive information about the labrum.
The maxillule in the adult has some resemblance to that of the female, but the basal portion
is reduced, while the palp, which is clearly articulated, bears 6 elongatc plumose setae, rather thaii the
short, hooked setae of the female.
The maxilla consists of 2 articles, the basal miich exceeding the distal in length and somewhat
in width. The basal article has no endite and does not develop the massive expansion and sclerotiza-
tion characteristic of the female. The distal article is directly and siniply articulated on the basal
and does not at ail form a hook-process. The armature of the terminal article is a long, plumose seta,
laterally directed. This seta is very possibly the homologue of the very small, but consistently présent,
seta of the external face of the distal article of the females of ail the species. Between the bases of the
maxillae, but not directly joined with them, is a small ventral protubérance. These 2 mouthparts and
the relatively elongate prehensile antenna and the general habitus of this male appear to us to form
an interesting parallel in modification, following a pattern of réduction which, when it occurs in notodel-
phyids, leads to the formation of what might be called the “ agnathaner ” male type. The modi¬
fications consist of réduction of ail gnathal parts of head appendages, with accompanying general
lengthening of such appendages and of their setae.
By and large ail the swimming legs conform very well to a general cyclopoid pattern, with per-
haps a distinctive feature in the segmentation of the first exopodites. .411 the protopodites are bimerous,
the coxopodites united by laniellae but lacking armature. The basipodites hâve latéral setae and in
the first leg there is the medial spine which occurs so widely through species of gnathostomes. The
endopodites are ail trimerons and they tend somewhat to lack of setae on some of the basal articles.
The exopodites of the first legs are bimerous, ail the others are trimerous. There are some setae lost
on basal articles. There are strong indications of correspondence with males of the Enteropsinae and
equally definite trends to dilTer from males of Ascidicola, which might be said to conform even more
to a general cyclopoid pattern, and of Botryllophilus, which has distinctive supplementary modifications.
The fiflh leg is definitely much reduced, with indications of 2 articles, but the basal one is com-
pletcly coalesced with the body. There is retained a small lobe, bearing a seta. The distal article
is small, somewhat hilobed and bears a single short seta. The position of the fifth leg at the posterior
margin of the first urosomal segment is a very generally occurring cyclopoid trait for males.
The so-called sixth leg is at the posterior margin of the second urosomal segment. There is
no articulated appendage, Init a hilobed prominence at either side ol the segment is set with 2 setae
at the latéral distal margin.
The caudal rami hâve a very cyclopoid aspect, both in general shapc and arrangement and in
the armature of 4 terminal plumose setae and of single dorsal and single latéral reduced setae.
KEY TO SPECIES OF ENTEROCOLA BASED ON FEMALES
1. With mammiform processes at the bases of some or ail of the thoracic legs. 5
1'. Thoracic legs without mammiform processes. 2
2. Pediform projection-fifth leg with very long setae. setifera Hansen, 1923, p. 49
2'. Pediform projection-fifth leg with very short or barely visible setae or without setae.... 3
3. Dorsal surface of metasoinc produced into pairs of conspicuous overlapping folds.
plerophora Chatton & Brément, 1909, p. 50
3'. Dorsal surface without conspicuous overlapping dorsal metasomal folds. 4
4. Urosome lacking visible segmentation. megalova Gotto, 1962, p. 49
Source ; MNHN, Pans
PAUL L. ILLG AND PATRICIA L. DUDLEY
48
4'. Urosome clearly segmeiited. laticeps, n. sp., p. 91
5. Setae of antenna short, hooked, clawlike, length of longest seta less than half greatest width
of appendage. fulgens van Beneden, 1860, p. 63
5'. Setae of antenna normal, flexible, length of longest seta considerably more than half greatest
width of appendage. 6
6. With distinct mammiform processes between all leg pairs. 10
6'. Mammiform processes missîng or very indistinct between one or more pairs of legs. 7
7. Setae of antenna arranged so that an apical pair îs set off on a distinctive lobe.
cUii’elinae, n. sp., p. 66
T.
8 .
8 '.
10 .
10 '.
11 .
11 '.
12 .
12 '.
13.
13'.
14.
Setae of antenna include 4 or more arranged in a compact apical row. 8
Setae of endopods 1-4 set close together apically on ramus. 9
Lateralmost of 2 apical setae of each endopod set more on the latéral surface than apical, and
diverging from the axis of the ramus. hessei Chatton & Harant, 1924, p. 49
Third endopod modified, outline more lobate than styliform ; setae of endopods 1-4 about as
long or shorter than ramus. precaria, n. sp., p. 72
Third endopod with usual styliform outline ; setae of endopods 1-4 distinctly longer than the rami
ianlhina, n. sp., p. 74
Antenna with apex long, narrow, bearing 7 setae. bilamellala Sars, 1921, p. 49
Antenna with apex broad, bearing 6 setae. 11
.Antennule lacking setae. mammifera Chatton & Harant, 1922, p. 49
Antennule with setae. 12
With caudal rami fairly developed, set off to some degree from the anal somite. 13
Caudal rami simply indicated by terminal bilobing of body, not definitely demarcated.
sydnii Chatton & Harant, 1924, p. 76
Antenna clearly unimerous. petili Guille, 1964, p. 80
Antenna completely or partially bimerous. 14
Endopods 1-4 subrectangular, with more or less parallel margins, elongate ; apical setae well
separated at bases. brementi, n. sp., p. 82
14'. Endopods 1-4 with médial margin curved more or less strongly, short ; apical setae set close
together. fertliis, n. sp., p. 85
non Enterocola spp. :
EnUrocola beaumo7\ti Scott & Scott, 1895
Enterocola beaumonli, T. & A. Scott, 1895, p. 359-360, pl. 16, 6g. 9, pl. 17, 6gs. 9-12 (from Valentia,
Ireland, from an ascidian).
Placed in Haplosloma — [Aplosloma) by Chatton & Brément, 1909c, p. 228.
Placed in Haplostomides by Chatton & Harant, 1924d, p. 406-407.
Haplostominae, sp. incertae sedis, Ooishi & lUg, 1977, p. 79.
Enterocola (sic) eruca Norman, 1868
Entorocola eruca Norman, 1868, p. 300 (from Shetland Islands, from Ascidia inteslinalis).
Removed from EnUrocola, suggested as belonging to Aplosloma by Canu, 1886, p. 372. — Placed in
Aplostoma by Brément, 1909, p. 83-85, 87.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Haplostoma eruca, Chatton & Harant, 1922e, p. 419. — Ooishi Si Illg, 1977, p. 12 (see for additional
citations).
Enterocola kibernica, Scott & Scott, 1895
Enlerocola kibernica, T. & A. Scott, 1895, p. 360, pl. XVII, figs. 3*8 (from Valentia, Ireland, from an
ascidian).
Placed in Haplostomides by Chatton & Harant, 1924d, p. 405*406, 407.
Haplostomides hibernicue Ooishi & Illg, 1977, p. 78 (see for additional citations).
SPECIES OF ENTEROCOLA
Enterocola bilamellata Sars, 1921
Enterocola bilamellata Sars, 1921, p. pl. 36 (type locality, Norway Coast, host unknown). — Sewell,
1949, p. 191. — Gotto, 1960, p. 225 ; 1962a, p. 544. — Guille, 1964, p. 289.
Distribution : Coast of Norway.
Host : unknown.
Enterocola hessei Chatton & Harant, 1924
Enterocola heesei Chatton & Harant, 1924b, p. 358-360, 363, fig. 3 (type locality, Baie de Morlaix, France,
from compound ascidians). — Harant, 1931, p. 371. — Gotto, 1960, p. 226 ; 1962a, p. 544. —
GuiUe, 1964, p. 289.
Distribution : France, Channel Coast.
Host : Unknown.
Enterocola mammifera Chatton & Harant, 1922
Enlerocola mammifera Chatton & Harant, 1922a, p. 153-154, 156, fig. 3 (type locality. Golfe du Lion,
France, from Aplidium asperum Drasche). — Chatton & Harant, 1922c, p. 245, 248, 249 ; 1924b,
p. 363. — Harant, 1931, p. 371. — Gotto, 1960, p. 226 ; 1962a, p. 544. — GuUie, 1964, p. 289.
Distribution : Golfe du Lion, Mediterranean.
Host : Aplidium asperum Drasche.
Enterocola megalova Gotto, 1962
Enterocola megalova Gotto, 1962, p. 541-543, fig. 1 (type locality Strangford Lough, Ireland, from Poly-
clinum aurantium Milne Edwards). — Guille, 1964, p. 290. — Gotto, 1966a, p. 193.
Distribution : Ireland.
Host : Polyclinum aurantium Milne Edwards.
Enterocola selifera Hansen, 1923
Enterocola setifera Hansen, 1923, p. 27-28, pl. III, fig. 3a-3i (type locality, S. W. of Iceland, from bottom
sample). — Sewell, 1949, p. 194. Guille, 1964, p. 289, did not accept the species. In a foot-
note he stated “ ... Je ne pense pas que cette espèce appartienne au genre Enterocola... ’ Very
Source : MIIHN, Paris
50
PAUL L. ILLG AND PATRICIA L. DUDLEY
rightly he pointed out the antenna was unlike anything to be expected in Enlerocola and seemed
more related to structures found in the Notodelphyidae. He also noted discrepancies between
other appendages as described by Hansen and the standard characters for the genus. Having
accepted his opinion -we were inclined to exclude the species from Enterocola. However, our
study of the whole family Ascidicolidae and our noting of the range of variation in some appen¬
dages hâve led us to alter our opinion. There are clearly some ambiguities and the situation
demanda that the species be reeollected and redescribed. Hansen himself stated he was dissa-
tisfied with his depiction of some of the important appendages. The animal is without doubt
an ascidicolid and conforms, very roughly, but sufQciently, to be placed as a species of Enterocola,
with the description unsatisfactory. The strange antenna depicted by Hansen now is more
acceptable to us in the light of our study of species of Slyelicola. In these there is a prehensile
apical claw on the antenna. Further, on the basal segment of the antenna of S. bahusia there
is a process, doubtless a much modified spine or seta, of appearance remarkably similar to the
element depicted by Hansen.
Distribution : Iceland.
Host : Unknown.
Enterocola sp. Claus, 1875
EiUerocola sp. Claus, 1875, p. 351-352, 360, pl. xxiv, fig. 31 (locality not given, from an ascidian). —
Délia Valle, 1883, p. 252. — Chatton & Harant, 1922a, p. 149.
Enterocola betencourti, Canu, 1891a, p. 474, 475 ; 1892a, p. 218. — Chatton & Brément, 1909c, p. 228.
Enterocola sp. A Chatton & Brément, 1909
Enterocola fuîgens, Délia Valle, 1883, p. 245-247, pl. 1, fig. 3-10 (from Naples, from Aplidium crystaUi'
num, A. gibbulosum and didemnids).
Enterocola fulgens, Canu, 1892, p. 216.
Enterocola sp. A, Chatton & Brément, 1909c, p. 228. — Chatton & Harant, 1922a, p. 156.
Enterocola sp. Chatton & Harant, 1924b, p. sès.
Enterocola sp. B Chatton & Brément, 1909
Enterocola ? fulgens, T. Scott, 1900, p. 386-387, pl. XIII, fig. 21-27, (from Scotland, from an uniden-
tified ascidian).
Enterocola sp. B, Chatton & Brément, 1909c, p. 228. — Chatton & Harant, 1922a, p. 156.
Enterocola sp., Chatton & Harant, 1924b, p. 363.
Enterocola sp. Schellenberg, 1922
Enlerocola sp., Schellenberg, 1922, p. 294 (from Plattenberg Bucht, South Africa, from Polycitor renieri
Hartmeyer).
Enterocola pterophora Chatton & Brément, 1909
(Figures 12, 13, 14, 15, 16, 17, 18, 19)
Enterocola pterophora Chatton & Brément, 1909b, p. 225-227, fig. 1-5, (type locality, Banyuls-sur-Mer,
Golfe du Lion, France, from “ LeptocUnum commurie ” ; 1911, p. 70 ; 1915, p. 145, 149, 153,
Source ; MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES 51
Fig. 12. — Enierocola pterophora Chatton & Harant, temale : a, habitus, latéral view ; b, antennule ; c, antenna ; d, basal
portion of maxillule ; e, maxillule ; 1, maxilla ; g, apex of maxilla, anterior view ; h, first leg ; i, second leg ; j, tbird
1^ ; k, fourth leg ; 1, pediform projection. Ail scales = 0.1 mm.
Source ; MNHN, Paris
52
PAUL L. ILLG AND PATRICIA L. DUDLEY
F.O. 13. - EM,™* Ch.,.on 4 H,r.n,, Trididmnun, lo.m, h.hi.u., vie» ■ b h.bitu.
n, Z ' n \ i ». Inbrum ; 1, m.Mllule , j, basai portion ol m.sdiiulo , h m.xiil.' ■ i apex
i ô «• “"l l»S i ». enJopodite ol seoond lop , m, Ihird iag ■ i, ’lourth
leg; 0, pediform projection; p. caudal ramus. Scales = 0.1 min. ® . wuru leg , n, lounn
Source : MNHN, Paris
Fig. 14. — Enierocola pterophora Chatton & Horant, Trididemnuin form, female : a, habitua, latéral view ; b. urosome,
latéral ; c, antennulc ; d, antenna ; e, maxillule, basal portion ; f, maxiUule, palp ; g, ûrst leg ; h, second leg ; i, third
leg and medial ventral protrusion ; j, fourth leg, ventral protrusion ; k, margin, pediform projection. Scales
= 0.1 mm.
Source ; MNHN, Paris
54
PAUL L. ILLG AND PATRICIA L. DUDLEY
' v.nTraf'STÏ ^ ^ ‘■. habitua, latéral view ; b, uroaome,
i S k fou^rr ^ J; ' ; «’ : f- maxillule ; g. maxüla ; h. firat leg ; i, second leg
J. thirdleg.k, fourthleg;!, pediform projection ;in, caudal ramua. Scales = 0.1 mm. «naieg,
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIËS
55
16. — Enterocola pterophora Chatton & Harant, Diplosoma form. female : a, habitus, latéral view ; b, antennule ;
c, antenna ; d, basal portion ot maxillule ; e, palp of maxillule ; f, maxilla ; g, first leg, nght ; h, first leg, left ;
i. second leg, left ; j. third leg, right ; k, fourth leg. right ; 1, caudal ramus. Scales = 0.1 mm.
Source : MNHN, Paris
56
PAUL L. ILLG AND PATRICIA L. DUDLEY
Source ; MNHN, Paris
ASCIDICOLIDAE AND SÜBFAMILIES
57
Fig. 18. — Enlerocola pUrophora Chmon & Harant,£>tpIosomÆ form, Banyuls, female : a, habitus, latéral view ; b, anten-
nule : c, antenna ; d, labrum ; e, maxillule ; f, maxillule ; g, maxilla ; h, firat leg ; i, first leg ; j, second leg ; k, third
leg ; 1, fourth leg ; m, caudal ramus. Scales = 0.1 mm.
Source ; MNHN, Paris
n, fouTth leg ; o, caudal ramus, ventral view. Scales = 0.1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
59
fig. 1. — Chalton & Harant, 1922a, p. 151-152, 154, 6g. 2 ; 1922c, p. 245. — Schellenberg,
1922, p. 294. — Chatton & Harant, 19241), p. 363. — Harant, 1931, p. 371. — Sewell, 1949,
p. 183. — Gotto, 1952, p. 674 ; 1954, p. 665 ; 1960, p. 221 ; 1962a, p. 544. — Guille, 1964, p. 289.
— Gotto, 1966a, p. 193.
Distribution : Mediterranean, especially Golfe du Lion.
Hosls : Didemnum commune (Délia Valle), D. asperum maculatum (Milne Edwards), D. fulgens (Milne
Edwards), Diplosoma Usterianurn (Milne Edwards), Parascidia {lava Milne Edwards, Cysto~
dite^ sp.
Spécimens examined :
From Didemnum fulgens Milne Edwards :
Northeast of Port Vendres, and ncar Cap Bear, ncar Banyuls-sur-Mer, France. 90 m, 60 m,
mi.xed trawl, May 14, 1958, 1 female.
Near Banyuls-sur-Mer, 60-100 m, April 25, 1958, 1 female.
Northeast of Port Vendres, mlxed trawl, 70, m, 30-60 m, May 6, 1958, 6 females, 1 male fifth
copepodid.
Cap Doune, near Banyuls-sur-Mer, France, 50 m, May 2, 1958, 1 female.
Near Port Vendres, 50 25 m, May 13, 1958, 6 females.
“ Leptoclinum fulgidum {L. commune Délia Valle ) ” :
Banyuls, September 26, 1910, Chatton number 44, 1 female, 1 male.
“ Leploc. fulgidum ” :
Aquarium, Banyuls, September 27, 1910, Chatton number 67, 1 female, 1 male, 2 fourth cope-
podids.
From Didemnum maculosum Milne Edwards :
Northeast of Port Vendres, and near Cap Bear, 90 m, 60 m, mixed trawl, May 14,1958, 6 females.
North of Port Vendres and off Cap Bear, in Anse de Paulilles, 70-30 m, 20-50 m, mixed trawl,
May 28, 1958, 3 females.
Gaiola, Bay of Naples, 25 m, March 13, 1958, 1 male.
“ Leptoclinum asperum — maculosum" :
Banyuls, October 17, 1910, Chatton number 117, 3 females.
“ Leptoclinum mascul. coriac ” :
Port Vendres, 100 m, January 10, 1912, Chatton number 157, 15 females, 1 male, 1 develop-
mental stage.
From PolysyncToton lacazei Giard :
Cap Doune, 50 m, May 2, 1958, 1 female.
From “ Leptoclinum sp. ” :
« Chalut au large de Cerbère, » September 25, 1912, Chatton number 162, 2 females.
From Trididemnum tenerum (Verrill) :
North and northeast of Port Vendres, 70 m, 30-60 m, mixed trawl, May 6, 1958, 1 female.
Duke Rock, near Plymouth, England 6 fm, September 8, 1958, 1 female.
Duke Rock, near Plymouth, England, 6 fm, September 9, 1958,1 female. 1 male 6fth copepodid.
From Trididemnum niveum Giard :
Troc l’Abeille, Banyuls, October 24, 1910, Chatton number 140, 1 female.
From Diplosoma gelatinosum var. lislerianum Milne Edwards :
Near Port Vendres, 50-35 m, May 13, 1958, 1 female, teratological.
From “ D. spongiforme ” :
Chalut, nord Argelès — Ganet, January 10, 1912, Chatton number 166, 10 females.
From “Diplosoma listeri, var KoUerianum Int-d’oscanius ” :
September, 1912, Chatton number 155, 1 developmental stage.
Our suite of specimens includes material collected by Chatton, at least one with a tentative
Source ; MNHN, Paris
PAUL L. ILLÜ AND PATRICIA L. DUDLEY
identification by him, and extends the rester of host species. Material from the typical host, as best
as we can identify it from the old accounts, establishes a sufiieient range of variation that we hâve felt
confident in assigning material from additional ascidian species to this form. Our discussion below
attempts to account for the range of variation we encountered. We would point out it has only been
very rarely that such an extensive sampling of any species of ascidicolid has been similarly available
for study. We ofîer what we consider to be pertinent supplementary information to the original
description and figure a specimen from the probably typical host, Didemnum fulgens, coUected by
us from the région of the type locality, 2 females and a male fifth copepoidid from Trididemnum tenerum
from Plymouth, and a female from Trididemnum tenerum from the type area, 2 females from Diplosoma
gelalinosum from the type locality, collected by us.
Specimen from Didemnum fulgens (fig. 12) : Overall measurement 0.82 inm, as compared with
the lengths of 0.8 to 1.0 mm given by the authors, The proportions, cephalosome ; metasome ; uro-
some, are 1: 4: 2, corresponding to the authors ; ratio of 1: 5.5: 1 plus, but their figure, when measured,
présents approximately a ratio of 1: 4; 2. The habitus (fig. 12, a) is in agreement with the original
description with the slight exception that the animal has a somewhat greater measurement dorsal
to ventral. The antennule (fig. 12, b) is somewhat more narrowed apically than shown by the authors.
We see 4 setules rather than the 2 they discerned. The antenna (fig. 12, c) is very characteristic in
this species. It appears to be bimerous but the articulation is somewhat incomplète. The distal
article is rather fiat and plate*like with a definite curve of the axis toward the midline. There are
2 small setae on the distal latéral margina and a group of 3 curved setae on the distal médial margin.
The authors saw 4 small setae in the latter position. The maxillule (figs. 12, d, e) is bilobed. We
agréé with the author’s description but the anterior margin of the basal portion bears a seta and a
setule not mentioned by them. The maxilla (figs. 12, f, g) conforms to the original account. The
4 pairs of legs (figs. 12, h, i, j, k) are ail biramous and the protopodites are bimerous. The endopodites
of the first leg (fig. 12, h) and the second (fig. 12, i) are obscurely bimerous, those of the third leg (fig. 12,
j) and the fourth (fig. 12, k) unimerous. Each of the endopodites bears 2 apical setae, both shorter
than the ramus, with the latéral seta about one-third longer than the médial. The exopodite of the
first leg is obscurely bimerous, with at least an interruption in the culicle on one surface. There is a
characteristic terminal element with the shape of a pointed dôme. The exopodites of the second leg
and the fourth leg are unimerous and each has a similar pointed dome-like endpiece. The exopodite
of the third leg (fig. 12, j) is characteristic of the genus and consista of a curved stylet. The pediform
projection (fig. 12,1) is a subcircular plate which bears no éléments of armature. The urosome and cau¬
dal rami are similar to those shown by the authors but in this specimen the strongly difierentiated
segmentation shown by them does not appear.
Female specimens from Trididemnum tenerum (figs. 13, 14, 15) : The overall measurements
are 0.81, 0.75, 0.99 mm, measured aloiig the curve of the body to the tip of the caudal ramus. The
habitus (figs. 13, a, b ; 14, a ; 15, a) in each corresponds well to those of Chatton & Brément and the
specimen discussed above, from Didemnum. We présent additional details for the urosomes (figs. 14,
b ; 15, b, c). The antennules vary from a simple tapering lobe with 4 terminal setules (fig. 13, c) to
a somewhat longer appendage with 3 small terminal setules and a marginal setule (fig. 14, d), and to
a lobe with 3 terminal setules (fig. 15, d). The antenna has a characteristic form for the species, but
the proportions and the setation vary slightly (figs. 13, d ; 14, d ; 15, e). Two of the specimens hâve
exactly the pattern of setation shown by the authors. The labrum (fig. 13, e), not previously illustrated,
consists of a flattened lobe with rows of spinules on the surface and with 2 digitiform spinulose palps
emerging from the distal latéral corners. The maxillules (figs. 13, f, g ; 14, e, f : 15, f) are as described
for the specimen from Didemnum above. The maxilla (fig. 13, h ; 15, g) corresponds to the original
description. The legs show a great degree of minor variation within a consistent basic pattern. They
conform throughout in biramous construction with bimerous protopodites. In ail cases the endopo-
dite bears 2 terminal setae, always shorter than the ramus, but the relative lengths of these vary some¬
what. The endopodite of the first leg varies from unimerous (fig. 13, j) to obscurely bimerous (figs. 14, g :
15, h). The exopodite also varies from unimerous (figs. 13, j, k ; 14, g) to bimerous (figs. 15, h). In ail
specimens there is a subterminal cuticular point, but in one specimen (fig. 15, h) there is an additional
Source : MNHN, Paris
ASCIDICOLIDAI-: AND SUBI'AMILIliS
61
spiiiule at the distal ihird of the latéral margin. In the second legs the endopodite may vary from
obscurely bimerous (fig. 13, 1) to distinctly bimerous (figs. 14, h ; 15, i). The exopodites are ail unime-
rous. In 2 specimens there is no diflerentiated élément except a minute subterminal cuticular point.
In the remaining specimen (fig. 14, h) there is a difîentiated dome-like apex bearing a small subterminal
spine. In the third legs the endopodite varies from obscurely bimerous (figs. 13, m ; 14, i) to unimerous
(fig. 15, j). The exopodite in ail specimens is a curved pointed unimerous stylet. In ail specimens
the fourth leg has unimerous rami. The armature of the exopodite varies from a subterminal cuticular
point (fig. 13, n) to a terminal cuticular point (fig. 15, k) to a diflerentiated dome-like élément with
a terminal point (fig. 14, j). There are never mammiform processes between the legs, but there may
be a conspicuous single médial sclerotized lobe (figs. 14, i, j). The pediform projection is always a large
suhcircular plate. As minor variations there may appear slight marginal indentations (fig. 13, o)
and there may be présent a single marginal or submarginal seta, facing internally (figs. 14, k ; 15, l).
The caudal ramus (figs. 13, p ; 15, m) varies somewhal in length relative to the urosome and indeed
may become extremely elongate (fig. 14, b).
In one collection color features were noted ; the ova in the oviducts were bright pink and the
embryos in the ovisacs had cherry-red yolk.
Three female specimens from Diplosoma gelalinosum (figs. 16, 17, 18) that we dissected indicated
a strong différentiation from the material discussed above, but the sériés presented so much internai
variation that we cannot on the basis of the présent suite ofîer a clear-cut différentiation of these as a
separate taxonomie entity. Further, more extensive study may demonstrate a séparation. The
specimens were relatively large, a représentative specimen measuring 1.23 mm overall. The proportions
cephalosome ; metasome : urosome were 1: 4: 2. The dorsal plates are not as well differentiated as
in ourother specimens (fig. 16, a) but they areclearly indicated and the pediform processis similar insizeand
form to the typical specimens. The body shows a greater curvature than we saw in other material.
The antennule (figs. 16, b ; 17, a) is long, conical, with only a slight taper and with 2 or 3 apicalsetules.
There may be additional setules on the surface. The antenna (figs. 16, b ; 17, c) is indistinctly bimerous,
and although the appendage is long and narrow, the axial curvature is slight. The pattern of setation is
the same as in typical specimens. The labrum (fig. 17, c) conforms. The maxillule (figs. 16, d, e ;
17, d, e) and the maxilla (figs. 16, f ; 17, f) also conform. The first legis rather distinctive (figs. 16, g, h).
There is a setule on the protopodite near the articulation of the exopodite. The endopodite is bimerous
and unusually long. The right leg of the specimen dissected had the usual 2 apical setae, but the left had
an additional seta, probably an abnormality. The exopodite is unimerous, with the usual apical
structure. The second leg (fige. 16, i ; 17, g) also has the seta on the.protopodite and has a bimerous
endopodite, exceptionally long. The third leg (fig. 16, j) as in the other specimens studied, has the
endopodite unimerous, but there is a differentiated area in the cuticle perhaps indicating a suppressed
segmentation. The exopodite is particularly short and départs somewhat from the usual styliform
outline found throughout the genus. The fourth leg (figs. 16, k ; 17, h) may hâve the endopodite unime¬
rous or bimerous ; the exopodite is unimerous with the usual differentiated apical éléments. There
is a setule on the protopodite which we bave not seen on fourth legs of other specimens. The caudal
ramus (fig. 16,1) is elongate, conical.
The only specimen of E. plerophora from Diplosoma gelatinosum from our own collecting, and the
only one recently taken from this host, therefore, is a female which we can only interpret as a teratolo-
gical example (fig. 18). The body shapeisdistorted, and the length measurement probably would not be
significant. It does hâve the usual indications of dorsal plates and the pediform projections conform
to the other specimens from the same host. There is good correspondence with other specimens in the
labrum (fig. 18, d), maxillule (figs. 18, e, f), maxilla (fig. 18, g) and the caudal ramus (fig. 18, m). The
antennule (fig. 18, b) is swollen although it has the usual setation. The antenna (fig. 18, c), although
it has the usual outline, has aberrant setae. The first legs (figs. 18, h, i) are recognizable as those of
the species and would fall within the range of variation of the other specimens. The second leg
(fig. 18, j), however, has an exopodite which has a form more reminiscent of the typical third exopodite.
The third leg (fig. 18, k), although it has the typical exopodite, has a very reduced endopodite with
no terminal setae. The fourth leg (fig. 18, 1) is reduced to a uniramous unimerous, lobe, with 2 setules.
Source : MNHN, Paris
62
PAUL L. ILLG AND PATRICIA L. DUDLEY
Such aberrant specimens are seldom observed among the ascidicolids, doul)tless bccausc of the
relative rarity of almost ail the species. It is interesting to observe tbat although tbis specimen possesses
obviously monstrous features, it does conform to the basic generic and spécifie diagnoses. It empha-
sizes the variability of the leg structure in this particular species.
\Ve did not dissect the specimens from Polijstjncraton lacazei or from Didemnum maculosum,
but feel a reasonably secure identification could be made by superficial examination in connection
with observations made on the whole suite of material. For some of his specimens from D. maculosum
(Chatton No. 157) Chatton recorded that the body was pale rose, with “ ovaries ” and ovisacs violet.
The male fifth copepodid (from Trididemnum tenerum, Plymouth) (figs. 19, a-o) : the body
form (fig. 19, a) is cyclopoid, with the qualification that the habitus is distinctly copepodid. The
overall length, measured from the apex of the head to the end of a cuadal ramus, is 0.70 mm. The
segmentation is well defined. The individual was almost ready to molt into the adult and some of
the features of that stage can be made out through the cuticle. The body is demarcated into 3 régions.
The cephalosome doubtless incorporâtes the body régions corresponding to appendages through the
maxillipeds, but the latter, and the mandibles, are lacking. There are 4 free thoracic segments bearing
the swimming legs. The urosome {fig. 19, b) is 5-segmented. The first segment bears the minute
fifth legs and developing spermatophores can be discerned within this segment. The second urosomal
segment bas some ventral cuticular folds and indications of the spermatophoral sacs which will be pré¬
sent in the adult stage. The remaining 3 segments of the urosome are somewhat narrower than the
first 2. The last segment is approximately equal in length to the combined lengths of the preceding
2 segments, and internally there are indications that this segment will divide into 2 upon molting,
producing a urosome of 6 segment in the adult male.
The rostrum (fig. 19, c) is elongate, tapering and unornamented.
The antennule (fig. 19, d) is 6-segmented. The basal article is about equal in length to the
combined lengths of articles 2, 3, 4 and approximately 2 times wider than ail the remaining articles.
There is little taper in articles 2 through 6. The setation is as follows : I — 10 setae ; II — 2 ; III —
2; IV— 1;V — 1;VI — 7 setae, 1 aesthete. Itcan be observed that article IV will divide into 2 articles
in the adult, producing in the adult male an antennule of presuraably 7 articles.
The antenna (figs. 19, e, f) is trimerous, the articles having the proportional lengths of appro¬
ximately 2 ; 1: 1. The basal article bas a long setiform projection at the distal inner corner. The
second article is unarmed. The terminal article bears an articulated pointed element which contains
the developing articulated hook of the adult male (the outlines of the latter are dotted in Fig. 19, e).
In addition there are 4 subequal setae on the terminal margin and 1 seta, which is at least one-third
longer than the others, in an emargination slightly proximal to the apex on the outer margin.
The labrum was not made out in our préparation.
The maxillule is bilobed and has some resemblance to that of the female. The basal element,
perhaps derived by fusion of the protopodite and parts of the palp, forms a lamella (fig. 19, g) produced
medially and terminating in 2 spinulose processes, one of these articulated and bearing heavier spinules
than the other. The seta and setule of the anterior margin in the female are absent in this male cope¬
podid, and the sclerotization of the apex of the lamella is much weaker in the male copepodid. The
free palp (fig. 19, h) is lamellate, weakly sclerotized, and distinctly articulated. Across the truncate
apex there is a row of 5 subequal long setae bearing fine hairs. The 2 internai setae are clearly articu¬
lated, the other 3 are not articulated. On the external margin at the distal fourth there is an unorna¬
mented articulated seta as long as the terminal setae. Proximal to this the margin of the ramus is
sclerotized. On the internai margin at the distal third there is a small setule, unornamented. This
setule is not represented in the female. Judging from our specimen, in E. Canu’s présentation for
E. fulgens (1892, pl. 18) we would expect that the apical and external éléments will persist in the adult
male and will show increased length and plumosity.
"Hie maxilla (fig. 19, i) is obscurely bimerous. The endite of the basal article présent in the
female is absent here, although there is some lobing of the distal médial margin. The terminal article
is produced apically as 2 setiform processes ornamented with fine hairs. An unornamented seta arti¬
culâtes ou the outer margin. From an adult male and Canu’s figure, it is probable that the latter seta
Source : AlfJHW, Paris
ASCIDICOLIDAE AND SUBFAHILIES 63
maybeanelongate plumosesetainthe adult, and thatthe médial processes willdisappear. Theappendage
is not so heavily sclerotized as in the female and possibly difîers functionally as well.
The first leg (figs. 19, j, k) is biramous, with bimerous protopodite, trimerous endopodite and
bimerous exopodite. There is a bilobcd subtriangular intercoxal lamella. The coxopodite lacks clé¬
ments of armature and ornamentation. The basipodite bears a short seta on the latéral margin and
a distal médial spine which reaches almost to the distal margin of the second article of the endopodite.
Rows of spinules are found on the surface of the basipodite and at the base of the médial spine. The
2 basal articles of the endopodite (fig. 19, k) are unarmed and unornamented. The distal article bears
1 latéral seta, 3 terminal setae and a médial seta. The basal article of the exopodite bas a distal latéral
spine. The second article has a distal latéral spine, a terminal spine, a terminal seta and 2 médial
setae.
The second leg (fig. 19,1) is biramous, with bimerous protopodite, trimerous rami. The coxopo-
dites are joined by a bilobed subtriangular lamella. The basipodite bears a seta on the latéral margin.
The basal article of the endopodite bears a distal médial seta. The second article bears 2 médial setae.
The terminal article bears a latéral seta, 3 terminal setae and 2 médial setae. The 2 basal articles of
the exopodite each bears a latéral spine and a médial seta. The terminal article bears 2 latéral spines,
a terminal spine, a terminal seta, and 3 médial setae.
The third leg is just like the second except that the exopodite (fig. 19, m) has an additional media!
seta on the terminal article.
The fourth leg (fig. 19, n) is similar to the 2 preceding, except that there are only 5 setae on the
terminal article of the endopodite rather than 6, and the exopodite is as in the second leg.
The fifth leg (fig. 19, b) consists of a simple, unornamented lobe well articulated at the distal
margin of the first urosomal segment, accompanied by a seta inserted dorsally at the base of the lobe.
The sixth legs are indicated by bilobed unornamented protrusions at the distal ventrolateral margin
of the second urosomal segment.
The caudal ramus (fig. 19, o) is broad and fiat, about two-thirds as long as the anal somite.
There are 4 setae on the truncate terminal margin, these at least twice as long as the ramus. There
is a short dorsal seta just beyond the middle and a short seta set in an emargination at the middle of
the latéral margin.
We find that Canu’s description of the male and the account of development in Enterocola
copepodids are strongly confirmed by the details of anatomy in this specimen. We hâve remarked
upon the remarkable correspondences with notodelphyids above.
Enterocola fulcens van Beneden, 1860
(Figures 20, 21, 22)
Enterocola fulgens van Beneden, 1860, p. 155-160, 1 pl., 7 figs., (type locabty, coast of Belgium, from
Aplidium sp.). — Gerstaecker, 1870-1871, p. 774, 801. — Aurivillius, 1882b, p. 92-93 ; 1883,
p. 88-89. — Canu, 1890, p. 758 ; 1891a, p. 468-469 ; 1892, p. 29, 37, 48, 49, 59, 66, 67, 85-88,105,
143, 214, 215, 216-218, pl. 18, figs. 1-12, pi. 19, figs. 12, 13. — Norman & Scott, 1906, p. 203.
— T. Scott, 1907, p. 368-369. — Chatton & Brément, 1909c, p. 227. — Hartmeyer, 1911, p. 1736.
— Chatton & Brément, 1915, p. 146, fig. 1, c. — Schellenberg, 1922, p. 287-288, 294. — Chatton &
Harant, 1924b, p. 363. — Harant, 1931, p. 370. — Leigh-Sharpe, 1935, p. 48. — Lang, 1948,
p. 17. — Seweli, 1949, p. 183,188. — Gotto, 1952, p. 674 ; 1954, p. 666. — Barnard, 1955, p. 241.
— Gotto, 1957, p. 259 ; 1960, p. 216, 221 ; 1962a, p. 544. — Guille, 1964, p. 290. — Gotto, 1966a,
p. 193. — Hamond, 1973, p. 350.
non Enterocola fulgens, Délia Valle, 1883, p. 245-248, 251, figs. 3-10 (Naples, from Aplidium cristallinum
délia Valle, A. gibbulosum délia Valle, didemniens).
non Enterocola fulgens, T. Scott, 1900, p. 386 (Scotland, from unidentified ascidian) ; 1901a, p. 352.
Interocola fulgens, Hesse, 1862, p. 349, 354 ; 1864, p. 335.
Source : MNHN, Paris
64
PAUL L. ILLG AND PATRICIA L. DUDLEY
Fig. 20. — Enlerocola fulgent van Benenden, femaie I : a, habitua, ventral view ; b, habitua, dorsal view ; c, habitus,
latéral view ; d, antennulea (Al) and antenna (A2) ; e, antennule ; f, antenna ; g, apex of antenna ; h, labrum,
maxillule (Mxl) and post-oral labial sclerotization (labial acier.) ; i, basai portion ol maxillule ; j, palp of maxillule ;
k, maxilla ; 1, apex of maxilla, poaterior view. Scalea for a, b, c, = 0.2 mm ; other acales = O.I min.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Fig. 21. — Enterocola fulgens van Beneden, female I : a, first leg ; b, second leg ; c, third leg ; d, fourth leg ; e, pediform
projection ; f, Enterocola fulgens, female II, habitus, latéral view ; g, antennule (Al) and antenna (A2) ; h, basal
portion of maxillule. Scales = 0.1 mm.
Source : MNHN, Paris
Fig. 22. — Enterocda julgens van Beneden, female from PoJycJtnum sp. : a, habitus, latéral view; b, antennule (Al)
and antenna (A2) ; c, maxillule ; d, palp of maxillule ; e, maxilla ; f, first leg ; g, second leg ; h, third leg ; i, fourth
leg; j, pediform projection ; k, caudal rami. Scales ~ 0.1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
67
Enterocola belencourli Canu, 1891a, p. 474-475 (type localities, near Boulogne, from Polyclinum suoci-
neum, and Brittany, from Aplidium zostericola) ; 1892, p. 49,67,143, 218, pis. 18, figs. 13,14, pl. 20,
figs. 1-4. Chatton & Brément, 1909c, p. 228. — Hartmeyer, 1911, p. 1736. — Schellenberg,
1922, p. 294. — Chatton & Harant, 1922a, p. 149-151, 154, fig. 1. — Sewell, 1949, p. 183, 184,
188.
Distribution : Mediterranean, West coast of Europe, British Isles, Tafelbucht, S. Africa.
Hosts : Amaroucium erythraeum Michaelsen, A. gibbulosum Savigny, A. mediterraneum, Aplidium
ficoides van Beneden, A. ficus Savigny, A. paUidus Verrill, A. zostericola Giard, A. sp., Botryllus
schlosseri (Pallas), B. sp., Glossoforum luteum Giard, Glossophorum(?sabulosum), Macroclinum pul-
monaria (Ellis & Solander), Polyclinum aurantium Milne Edwards, P. luteum (Giard), P. succi-
neum Milne Edwards, Sidnyum turbinalum Savigny, Synoicum pulmonaria (Ellis & Solander).
Specimens examined :
From Polyclinum aurantium Milne Edwards var. luteum (Giard) :
“ Glossophorum luteum ”, Tatihou, April, 1911, Chatton number 8, 1 female.
” Glossophorum ”, La Hougue, June 5, 1911, Chatton number 50, 2 females.
From Polyclinum sp. :
La Vieille, near Roscoff, France, 20 m, 1958, coll. E, Binder, 1 female.
As indicated above, the bulk of our material is from the Chatton collection, and indeed the
specimen in Chatton number 8 bore the manuscript identification to this species. It very probably
is one of the three adult females collected by Bré.ment at Tatihou and referred to as E. belencourti
in connection with a description and illustrations by Chatton & Harant (1922a, p. 149-151, fig. 1).
AH these Chatton specimens conform rather well to this characterization of the species.
The lengths of 2 specimens measured were 1.78 and 0.95 mm, the latter a specimen which had
obviously been subjected to drying at one time. The authors give the lengths as ranging from 1.3 to
1.7 mm. The proportions of our specimens, cephalosome : metasome : urosome, were 1 : 4 : 1 or 1.5.
The corresponding formula of the authors was 1 : 3.5 : 1.
The habitus (figs. 20, a, b, c ; 21, c) corresponds well with figures by the authors and those of
Canu. Indentations at least indicate the segmentai composition of the body and this was not shown
by Chatton & Harant.
The antennules (figs. 20, d : Al, e ; 21, g : Al) do not correspond well with the illustration of
Chatton & Harant, but we believe that this can bc explained by the présentation of different views.
They probably did not observe the appendages in place on the animal. It is difficult to déterminé
whether there is an actual terminal segment on the appendage. On 1 specimen (fig. 21, g) there would
appear to be a minute conical terminal segment set at the distal latéral corner of the large subquadran-
gular basal segment. Spinules ornament this protubérance and at least 1 spinule is found on the
basal segment. A row of spinules illustrated by Chatton & Harant on the basal segment in reality
consists of a corrugation of the cuticle. In the other specimen (figs. 20, d, e) the apical element is so
small and spinulose that it resembles a modified seta but we believe it is probably homologous to the
terminal article.
The antenna (figs. 20, f, g) or one specimen is very broad, obscurely bimerous, and bears 5 to
6 short, clawlike setae around the distal margin. The longest of these is as long as about one fourth
of the greatest width of the appendage. Rows of spinules ornament the surfaces of the articles. In
the second specimen (fig. 21, g) the antenna is narrower, but this may be an artefact of fixation. There
are 4 apical clawed setae and a médial clawed seta. The longest of these is about one-third the greatest
width of the appendage. These figures correspond well with that of the apex of the antenna shown
by Chatton & Harant, although they apparently found 6 setae consistently.
The labrum (fig. 20, h) has not previously been illustrated.
The maxillule (figs. 20, h, i, j ; 21, h) is set above a complex labial sclerotization (fig. 21, h). It is
bilobed ; the basal portion is lamclliform and heavily sclerotizcd. On a lobe of the anterior margin
Source : Mt-JHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
there is born a seta and accompanying setule. The palp portion carries 5 uncinate, spinulose setae
on its distal margin and an unornamented normal seta on the outer margin at the distal third. Our
illustrations of the palp correspond to that of the “ exopodite of Chatton & Hafant.
The maxilla (figs. 20, k, I) is bimerous and heavily sclerotized. At the distal médial corner of
the basal segment there is a digitiform spinulose endite with a tnmcate distal margin. The terminal
segment terminâtes in 2 unarticulated hooks, one somewhat shorter than the other. On the surface
in an interruption in the cuticle there is set a small conical seta.
The legs (figs. 21, a, b, c, d) are biramous, with bimerous protopodites and unîmerous rami.
The coxopodites are unarmed ; each basipodite bears a small seta at the distal latéral corner. The
exopodites of the first, second and fourth legs hâve differentiated pointed apical dome-like éléments.
The expodite of the third leg has the styliform outline characteristic of most species of the genus. The
endopodites are longer than the exopodites and each bears 2 short setae, these much shorter than
the ramus in each case. The médial of the 2 is slightly longer. Between the legs of the third and
fourth pairs there are pairs of pronouncedly developed mammiform processes. Although there are
sinuous lines in the cuticle between the legs of the first and second pairs there are no differentiated
mammiform processes. Canu and Chatton & Hafant never reterred to the presence of these pro¬
cesses, although they are consistently présent in our specimens. This oversight has led to confusion
in subséquent treatments of the species and the genus.
The pediform projection (fig. 21, e) is a curved plate of circular outline. There are 2 widely
spaced marginal setae. The more dorsal of these is more conspicuous than in any other species of
ErUercola. Although Chatton & Harant refer to this seta they did not see the more ventral seta.
The caudal rami are long, conical and apparently articulate with the body. In our specimens
they were approximately 3 times as long as wide and about half the length of the urosome.
Our specimen from Polyclinum sp. (collected by Binder, 1958) has presented difflculties (fig. 22).
It confornis generally in habitus (fig. 22, a), it is 1.72 mm long and has the proportions cephalosome ;
metasome : urosome, 1 : 3 : 1.5. It should be noted that this specimen was somewhat contracted and
distorted when extracted from the preserved tunicate host. The antennule (fig. 22, b : Al) corresponds
to the second specimen discussed above. The terminal élément here appears even more like a typical
article. It bears more setules on the basal segment also. The obscurely bimerous antenna (fig. 22, b :
A2) is flatlened and very broad and bears 6 setae. The longest is half the greatest width of the appen-
dage. Others are very short and claw-like. The overall configuration conforme reasonably well to
the diagnostic condition, although there is a strong trend toward a more generalized aspect. The
maxillule (figs. 22, c, d) and the maxilla (fig. 22, e) are as described above. The legs (figs. 22, f, g, h, i)
do not conform very well to the previously described specimens, since the setae of each coxopodite are
longer, although they are still shorter than the ramus. The exopodites are proportionately longer
and the first 2 legs hâve slightly differentiated mammiform processes. The exopodite of the third
leg (fig. 22, h) is narrow, elongate and triangular and does not hâve the characteristic styliform aspect.
The pediform projection has 2 tiny setules, both of these much shorter than in the other specimens
(fig. 22, j). The caudal rami (fig. 22, k) are only twice as long as wide and about a third as long as
the urosome. They still articulate clearly. Thus, in the habitus, size, head appendages this speci¬
men conforms well with typical E. fulgens. The legs appear to départ to a considérable degree.
Enterocola clavelinae Chatton & Harant, 1924
(Figures 23, 24, 25)
EnUrocola clxivelinae Chatton & Harant, 1924b, p. 354-356, 363, fig. 1 (type locality. Baie de Morlaix,
France, from Clavelina ruina Lahille). — Harant, 1931, p. 360. — Gotto, 1960, p. 226 • 1962,
p. 544. — Guille, 1964, p. 289.
Distribution ; Channel coasts of France ; Mediterranean.
Hosts : Polycüor [Clavelina) ruina (Lahille).
Source ; MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Fig. !3. - R.mG.nl ol holotyp. ol ÜMercœl,. cMinae Ch.tlon & HaiaM (.-I) and Eb^gI» Ckntlon 4
Harant (g} : a, E. claveliTiae, metasome and urosome, dorsal view ; b, ürst leg ; c, second leg , d, third eg , ,
l«e I I, padilom projection, g, E. ej/dnii, t.mn.nt oi m.ta.om., latéral view ehowing ..eond thmngh ionith leg.
and mairmailorm processes and dorsal processes. Scales — 0.1 mm.
Source : MNHN, Paris
70
PAUL L. ILLf. AND PATRICIA L. DUDLEY
Fig. 24. — EnUrocola clavelinae Chatton & Harant, female : a, habitua, ventral view ; b, habitua, dorsal view ; c, uro-
some ; d, antennule ; e, antenna ; f, labrum ; g, maxillule ; h, maxillule ; i, palp of maxillule ; j, maxilla. Scales =
0.1 mm.
Source ; MNHN, Paris
Specimens examined :
“ Clavelina nana ”, Cmatton number 239, one partially dissected specimen, female, (possibly bolotype).
From Polycitor [Clavelina) nana :
Cap l’Abeille, near BanyuIs-sur-Mer, France, 25 m, June 3, 1958, 1 female.
Remarks ;
From the Chatton specimen, which is very possibly the bolotype of the species, although it
is only a fragment, we can offer some further characters. We présent a dorsal view of the entire rem-
nant (fig. 23, a) to show the paired dorsal duplicatures mentioned by the authors. We were able to
dissect ofï ail 4 swimming legs. Although the authors mentioned that paired mammiform processes
were présent only between the third and fourth legs, we saw partially differentiated processes between
the second legs (fig. 23, c). AU the legs hâve bimerous protopodites and unimerous rami. The exo-
podites of the first (fig. 23, b), second (fig. 23, c) and fourth (fig. 23, e) legs hâve typical differentiated
pointed dome-like apical éléments. The exopodite of the third leg (fig. 23, d) has the characteristic
styliform outline of most Enterocola species. Ail of the endopodites hâve 2 apical setae, these longer
than the ramus itself. Rather characteristic is the placement of the latéral seta somewhat more proxi-
mally than the medial seta. The apical margin of the endopodite is thus somewhat inclined. The
pediform projection (fig. 23, f) is a circular, curved plate, with a single small setule on the margin but
Source : MNHN, Paris
72
PAUL L. ILLG AND PATRICIA L. DUDLEY
facing internally. The authors said that this process, which they termed the oostegite, was aseti-
gerous. The caudal rami, as indicated by the authors, are not delimited from the urosome. The
urosome is shown in latéral view in figure 23, a, because of the distortion of the specimen.
Our specimen from Banyuls (figs. 24, 25) was strongly contracted so the habitus (figs. 24, a, b)
is not really représentative of the normal individual. The length is 0.93 mm, in this contracted State.
The urosome (fig. 24, c) appears segmented, but it is possible that the contraction has produced the
strongly indicated folds. There is a midventral insémination pore and séminal tubes diverge toward
the oviducal apertures far laterally. The antennule (fig. 24, d) is a conical, unsegmented lobe, with 2
anterior setae and 3 smaller terminal setae. Spinules ornament the surface near the apex. The
antenna (fig. 24, e) is strongly bimerous, with the terminal segment longer than the basal. The basal
article is unarmed. There is a short seta at the middle of the médial margin of the terminal segment.
Set apically there are 2 setae which are about 1.3 tiraes as long as the greatest width of the appendage.
Both of these setae are set on low lobes. Separated from the apex, on the latéral margin there are
3 graduated setae which are shorter than the apical setae. Cbatton & Harant also noticed that
there are emarginations between ail the setae and a very marked emargination on the latéral margin
beween the group of 3 latéral setae and the 2 apical setae. The labrum (fig. 24, f) has not previously
been figured. The maxillule (figs. 24, g, h, i) is bilobed ; the basal portion is lamellate and bears a seta
and a setule on a lobe on the anterior margin. The palp bears 5 distal setae and a seta on the external
margin. The maxilla (fig. 24, j) is bimerous and is heavily sclerotized. At the distal médial corner
of the massive basal segment there is an unornamented digitiform endite. The distal segment is bifid
apically, has some spinulose ornamentation and there is a seta set in an anterior cuticular interruption.
The legs (figs. 25, a, b,c, d) agréé very closely with those we illustrate from the CnATTONspecimen with
the exception that we saw a seta on the latéral margin of each basipodite and the mammiform pro¬
cesses were not apparent in this highly contracted specimen. The peditorm projection (fig. 25, e)
bears a salient minute dorsal marginal setule and a more ventral marginal exceedingly minute setule.
The ovisac (fig. 25, f) is distinctly curved anterioly.
Types :
Entbrocola PRECARIA new species
(Figure 26)
Holotypic female (type locality Gaiola, Bay of Naples, Italy, from Polycitor {Eudistoma) mucosus
(Drasche) and paratype listed below.
Specimens examined :
From Polycitor {Eudistoma) mucosus (Drasche) :
Gaiola, Bay of Naples, Italy, 35-40 m, January 2, 1958, holotypic female and female paratype.
Description :
Female (fig. 26) :
The body (figs. 26, a, b) measures 0.85 mm ; the specimen is rather contracted. The propor¬
tions, cephalosome : metasome : urosome, are 1:3:1. Paired dorsal duplicatures are only indicated
on the surface and are not salient. The urosome (fig. 26, c) appears segmented, but because of the
contraction of the specimen, the appearance may be due merely to folding of the integument. The
antennule (fig. 26, d) is unimerous with a strong distal taper from about the basal third. There are
about 8 small setules concentrated at the distal third on the anterior and apical margins. The antenna
(fig. 26, 3) is unimerous, subrectangular. There is a short medial seta at the distal fourth, 4 equispaced
setae on the rather truncate distal margin and a shorter seta at the distal of the latéral margin. The
longest seta is 4/5 the greatest width of the appendage. The labrum (fig. 26, f) is an unornamented
semicircular plate with a spinulose palp extending posteriorly from each posterior latéral corner. The
Source ; MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILllîS
73
; b, habitus, latéral v
Fie. 26. — ËTiUrocola precaria, new speciee, paratypic female : a, habitus, ventral ^ -ii t • . ii.
Bome and pediform projection ; d, antennule ; e, antenna ; f, labrum ; g, maxillule ; h, paJp of maxillule , i, maxilla
i first leg ; k, second leg ; 1, third leg ; m, fourth leg ; n, pediform projection. Scales = 0.1 mm.
Source ; MNHN, Paris
74
PAUL L. ILLG AND PATRICIA L. DUDLEY
inaxillule (figs. 26, g, h) is bilobed. The basal portion is lamellate and bifid distally. On the anterior
margin there is a lobe set with a seta and accompanying setule. The palp bears 5 spinulose setac on
the distal truncate margin and 1 unornamented seta at the middlc of the external margin. The maxilla
(fig. 26, i) is bimerous, with the basal segment massive, heavily sclerotized and bearing at the distal
médial corner a curved articulated spinulose digitiform endite. The distal segment has a terminal
massive hook and a second long narrow digitiform process articulated on a lobe on its surface. A small
seta is set in a cuticular interruption on the anterior margin. AU the legs (figs. 26, j, k, 1) hâve bimerous
protopodites and unimerous rami. The protopodites are unarmed. The expodites of the first and
second legs end in a pointed cuticular specialization and each has a pointed cuticular deflection at
the middle of the latéral margin. The expodite of the third leg has a distinctive outline, lobate distally
rather than styliform (fig. 26, 1). The exopodite of the fourth leg (fig. 26, m) is like that of the first
and second except that it lacks the latéral cuticular point. The endopodites on the whole are very
short, just slightly exceeding the lengths of the exopodites. The 2 setae of each are set apically. In
the first 3 legs these setae are approximately as long as the endopodite and they slightly exceed
it in the fourth leg. The pediform projection (fig. 26, n) is a curved plate with circuler margin. It
bears a single setule in the middle. The caudal rami (fig. 26, c) are simple conical lobate extensions of
the urosome without apparent articulations.
This is very possibly a host form of EnUrocola fulgens, but it does possess distinctive morpho-
logical characters, particularly of the maxilla and the third leg. The name is derived from the Latin
term for uncertain.
Enterocola ianthina new species
(Figure 27)
Types :
Holotypic female USNM (type locality vicinity of Jolo, Philippine Islands, from Polycitor
{Eudistoma) iantkinus Sluiter), and paratype.
Specimens examined :
From Polycitor {Eudistoma) ianthinus Sluiter :
Vincinity of Jolo, Philippine Islands, USNM 6029, Accession No. 53256, Station 5139, Alba-
tross Philippine Expédition, 20 fathoms, February 14, 1908, holotypic female, 1 paratypic
female.
Description :
Female (fig. 27) :
Body (figs. 27, a, b) measures 1.08 mm. Proportions, cephalosome : metasome : urosome :
are 1 : 3 : 1. Dorsal duplicatures are only indicated and are not salient. The antennule (fig. 27, c)
is obscurely bimerous, with the portion corresponding to a distal segment about 1/3 as wide as the
widest part of the basal segment. There are about 7 setules concentrated on the distal half of the ante¬
rior margin and on the apex. The antenna (fig. 27, d) is bimerous, with the apical segment longer.
The basal segment is unarmed. On the apex of the second segment there are 4 subequal cquispaced
setae. At the distal fifth of the médial margin there is a shorter seta. At the distal fifth of the latéral
margin there is inserted the longest seta of the appendage. This seta is longer than the greatest width
of the distal segment of the appendage. The labrum (fig. 27, e) is a semicircular plate with a spinose
lobe extending posteriorly from each posterior latéral corner. A row of spinules ornaments the distal
margin. The maxillule (figs. 27, f, g) is bilobed. The base is lamellate and bifid distally. Set on the
surface near the anterior margin is a seta and an accompanying setule. The palp bears 5 spinulose
setae on the curved distal margin and a spinulose seta at the distal third of the external margin. A
patch of spinules ornaments the surface. The maxilla (fig. 27, h) is bimerous. At the distal médial
corner of the massive basal segment there is an articulated endite which is somewhat bifid apically
Source ; MNHN, Paris
ASCIOICOLIDAE AND SUBFAMILIES
75
Fig. 27. — Enltrocola iarUhina, new gpecies, holotypic temale ; a, habitus, veutral view ; b, habitus, latéral view ;
c, antennule ; d, autenua ; e, labrum ; f, maxillule ; g, palp ot maxillule ; h, maxilla ; i, first leg ; j, exopodite ot
first leg ; k, second leg ; I, exopodite of second leg ; œ, third leg ; n, exopodite of third leg ; o. exopodite of fourth
leg ; p. pediform projection. Scales = 0.1 mm.
and bears a patch of spinules near the apex. The distal segment has a massive terminal hook and a
second articulated shorter hook on the surface. The latter hook is ornamented with spinules. A small
seta is inserted on the anterior margin. Ail the legs (figs. 27, i, j, k, 1, m, n, o) hâve bimerous proto¬
podites and unimerous rami. There are pronounced mammiform processes between the legs of the
second and third pairs, none in the first pair, and a single slightly bilobed plate in the fourth pair.
Although the endopodites hâve a marginal interruption in the cuticle there is no well-developed arti¬
culation on either major surface. In ail legs the basipodites bear a seta at the distal latéral corner.
Source : MNHN, Paris
76
PAUL L. ILLG AND PATRICIA L. DUDLEY
The exopodites of the first legs (figs. 27, i, f) and the second legs (figs. 27, k, 1) are shorter than the endo-
podites and each has a differentiated pointed dome-like élément. The exopodite of the third leg
(figs. 27, in, n) has the characteristic styliform outline of most species of Enterocola and is longer than
the endopodite. The exopodite of the fourth leg (fig. 27, o) ends in a simple pointed element and
does not hâve the articulated dome-like structure. The endopodites taper somewhat distally and
the 2 setae on each endopodite are subequal and longer than the ramus. The pediforra projection
(fig. 27, p) is a semicircular curved plate which has no element of armature although there are 4 very
shallow emarginations on each, these delimiting 2 very shallow lobes, 1 dorsal and 1 ventral. The
caudal ramus (figs. 27, a, b) is subrectangular in form and has no éléments of armature. It articulâtes
distinctly with the urosome.
The spécifie name dérivés directly from that of the host. The species is not a strongly differen-
tiated one ; it is very similar to E. fulgens. The key above indicates differentiating characters.
Enterocola sydnii Chatton & Harant, 1924
(Figures 28, 29)
Enterocola sydnii Chatton & Harant, 1924b, p. 354, 356-358, 363, (type locality Locquirec [Côtes-du-
Nord] France, from Sydnium concrescens Giard). — Harant, 1931, p. 371. — Gotto, 1960, p. 226 ;
1961, p. 153 ; 1962a, p. 544. — GuiUe, 1964, p. 289. — Gotto, 1966a, p. 193.
Distribution : Channel coasts of France and England.
Hosts : Sidnyum turbinatum (Savigny) and S. elegans.
Specimens examined :
From Sidnyum turbinalum (Savigny) :
“ Circinalium concrescens ” : from Locquirec, France, Chatton number 246, 1 partially dissected
specimen, female. Possibly the holotype.
From Sidnyum elegans (Giard) :
“ Fragarium elegans ”, Roscoff, France, 1922, Chatton number 26, 1 female.
From Morchellium argus (Milne Edwards) :
From Duke Rock, near Plymouth, England, 6 m, September 9, 1958, 1 female.
From the Chatton specimen, number 246 which is very possibly the holotype of the species,
although it is only a fragment of the thorax, we can illustrate some further characters beyond those
originally described. The authors pointed out there were dorsal duplicatures on segments 3 and 4
(corresponding to the third and fourth legs). Our fragment shows similar processes also on the seg¬
ment of the second legs. The original description stated there were mammiform projections between
the legs of aU pairs, those of the fourth legs mucronate. We illustrate the projections of the second
through fourth legs (fig. 28, a ; 29, a). We cannot verify the authors’ statement that the endopodites
are twice as long as the exopodites ; the specimen shows the endopodites slightly longer than the exopo-
dites, but we feel sure that the true articulations of these rami were not properly taken into considéra¬
tion in the original description. The illustration presented by Chatton & Harant of the second leg
shows the outlines almost exactly as we see them, but the articulation of the exopodite was not shown
at ail and that of the endopodite was probably taken from the wrong surface. We cannot account for
the apparent exaggeration of the lengths of the setae in the authors’ illustration. We illustrate the
third and fourth legs of the species for the first time.
Female from Sidnyum elegans (fig. 28) : Overall measurement 1.5 mm. This is somewhat
longer than the authors gave for the type material, namely 1.2 mm. The proportions, cephalosome :
metasome : urosome, are 1: 4 : 1.5 ; the corresponding formula from the original description of Chatton
Source : MNHN, Paris
F,O. 28. EnUrocola sydnii Chatton & Harant, female from Siiinyum : a, habitua, ventral ; b. habitua, dorsal ; c
tua latéral ; d, uroaome ; e, antennuies ; f, antenna ; labrum and poat-oral aclerotization ; h, maxillule ;
of maxillule ; j, maxilla ; k, firat leg ; 1, second leg ; m, third leg ; n, fourth leg ; o, pediform projection,
for a, b, c = 0.2 mm, other scales = 0.1 mm.
i, palp
Scales
Source : MNHN, Paris
29. _ Enlerocola sydnii Chatton & Harant, female from Morckellium : a, habitus, ventral ; b, habitus, latéral ;
c, urosome ; d, antennule ; e, antenna ; f, maxillulc, basal portion ; g, maxillule, palp ; h, maxillule, ventral view ;
i, maxilla ; j, first leg ; k, second leg ; 1, third leg ; m, fourth leg ; n, margin of pediform projection of sixth tho-
racic segment ; o, caudal rami. Scales = 0,1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
79
& Habant was 1 : 3.5 : 1.5. The habitus (figs. 28, a, b, c) agréés fairly well with the original descrip¬
tion, although our specimen is somewhat bulkier and does not show the flexure shown. We illustrate
the urosome (fig. 28, d) and show the midventral insémination pore and diverging séminal tubes.
The antennule (fig. 28, e) is unimerous with a slightly difîerentiated apex, which suggests a short
segment. There are 3 short setae and several setules on the anlerior margin. This appendage does
not conform closcly to the appendage illustrated by Chatton & Habant but we feel that their illus¬
tration is from an anterior view and thus could actually be interpreted to conform to the structure
we T-iewed in place on the specimen. The antenna (fig. 28, f) is obscurely bimerous and has a short
seta at the distal third of the médial margin of the portion of the appendage corresponding to the ter¬
minal article. On the curved apex there are arranged 5 short setae. The longest seta is slightly
shorter than the greatest width of the appendage, so the setae are somewhat shorter than those illustra¬
ted by the authors. The labrum (fig. 28, g) has not been previously illustrated. There is a complex
labial sclerotization in the integument of the ventral body surface. The maxillule (figs. 28, h, i) is
bilobed and conforms generally to the original description ; it has not been previously illustrated.
The maxilla (fig. 28, j) is bimerous. At the distal médial corner of the massive heavily sclerotized
basal segment there is an articulated, subrectangular endite which bears rows of spinules both on the
distal margin and on the surface. The distal segment is prolonged medially as two unarmed unarti-
culated hooks, one somewhat shorter. On the anterior margin there is a short stifî seta inserted in an
interruption in the cuticle.
The fîrst legs (fig. 28, k) hâve not previously been illustrated. The protopodites are bimerous
and the rami are unimerous. There is a short latéral seta at the distal latéral corner of the basipodite.
The endopodite is only slightly longer than the endopodite, and the 2 apical setae are slightly shorter
than the ramus. The exopodite bears a terminal differentiated small hook. The remaining legs
(figs. 28, 1, ro, n) correspond well with our observation on the partial specimen from Sidnyum turbina-
luni. Paired mammiform processes occur between the legs of each pair. These between the third
and fourth legs are more prominent than the others ; however, the processes of the fourth legs hâve
a truncate apex rather than the mucronate condition. The pediform projection (fig. 28, o) is as des-
cribed except that there are 2 tiny setules on the margin. The caudal rami (fig. 28, d) are not demar-
cated and appear to be simple prolongations of the urosome.
Femalc from Morchellium argus : The body of this specimen was rather contracted ; it measured
overall 0.92 mm. The habitus (figs. 29, a, b) corresponds fairly well to the specimen from Sidnyum
elegans. Possibly it is the contracted State which is responsible for the more salient dorsal pro¬
cesses. Because of the contracted state, the urosome (fig. 29, c) appears to be segmented. There
are no real articulations. The génital apparatus on the fîrst segment of the urosome consists of a
midventral protrusion supporling an insémination pore ; diverging internai séminal canals procced
toward the oviducal apertures which hâve accompanying complicated sclcrotizations, each with a
setule, at the base of the pediform projections.
The antennule (fig. 29, d) compares well with our other specimens, except that it appears to
be somewhat more heavily sclerotized. The antenna (fig. 29, e) is more clearly bimerous than in
our other material, and the médial seta is found at the middle of the margin rather than at the distal
third. The apical armature is similar, except that the central seta is shorter. The appendage has
heavy sclcrotizations and some spinules on the surface. The maxillule (figs 29, f, g, h) is very similar
to the other specimens. The maxiUa (fig, 29, i) has the same general form but the endite is a curved
digitform process and bears circular rows of spinules. The fîrst two pairs of legs (figs. 29, j, k) havea
somewhat different appearance from those of the other specimens. Although the protopodites are
bimerous, there is no latéral seta on the basipodite. Tht endopodites are similar in outline, but the
endopodite of the first leg has the 2 terminal setae considerably shorter than the ramus. The exopo¬
dite of the first leg (fig. 29, j) terminâtes in a point and there is an emargination at the distal latéral
fourth. There is no differentiated apical élément. The exopodite of the second leg (fig. 29, k) although
constricted at the distal third, lacks the cmarginations of the first leg, and also lacks the usual apical
élément. The third leg (fig. 29,1) is similar to the other specimens in the protopodite and the exopo¬
dite, but the endopodite is much shorter. The fourth leg (fig. 29, m) is very similar to that of the spe-
Source : MNHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
cimen from Sidnyum turbinatum. Mammiform processes are developed between the legs of each
pair but we did not détermine whether those of the fourth pair were mucronate. The pediform pro¬
jection (fig. 29, n) and the caudal rami (fig. 29, o) are very similar to those of our other specimen.
The specimen last described, from Morcheüium argus has some slight différences from the other
material, as pointed out, and so may actually represent a host form, or alternatively, may be a distinct
species. Since we had only a single specimen to study we were unable to explore these possibilities
further. We found the taxonomie référencés to the tunicate hosts involved completely chaotic and
hâve used a terminology which we hope will keep our records of collection clear. It seems from what
we can best make out that our specimens ail came from closely related hosts.
Entebocola petiti Guille, 1964
(Figure 30)
Enterocola pelili Guille, 1964, p. 283-288, 289, fig. 1-3 {type locality, Banyuls-sur-Mer, Golfe du Lion,
France, from Amaroucium densum Giard).
Specimens examined :
From Polycitor {Paradisloma) cristallinus Renier : identification uncertain.
Cap l’Abeille, near Banyuls-sur-Mer, France, 25 m, May 7, 1958, 1 female 1 developmental
stage.
We hâve found one adult female (figs. 30, a, b, c) which appears to us to correspond reasonably
well to the description of this species. The overall length is 1.09 mm, compared with the range of
1.0 to 1.2 mm of the original material of Guille. The proportions, cephalosome : metasome : urosome,
are 1 : 4 : 1, as compared with 1 : 2.7 : 1 of the original description. The author does not depict dorsal
plates ; on our specimen there are slight indications of duplicatures (figs. 30, b, c). The antennule
(fig. 30, d) is unimerous, and does not show the number of setae and setules found by the author but
it is very possible our specimen is somewhat deformed in this regard. The antenna (fig. 30, e) labrum
(fig. 30, f), and maxillule (figs. 30, g, h, i) correspond well to the original description. The maxilla
(fig. 30, j) is a reasonably typical appendage for the genus but does not quite correspond with the original
description and figure. It consists of a massive basal segment with a digitiform spinulose endite at the
distal médial corner. The clearly articulated distal segment terminâtes in 2 hook-Iike extensions, one
unornamented and considerably longer than the other, which is spinulose. There is a short seta set
in an interruption of the cuticle near the anterior margin. The legs (figs. 30, k, 1, m, n) hâve bimerous
protopodites, rather than unimerous as indicated in the original descriptions and the basipodite bears
a short seta at the distal latéral corner. The exopodites of the first legs (fig. 30, k), the second legs
(fig. 30, 1), and the fourth legs (fig. 30, n) are very similar in outüne to those illustrated in the original
description and each bears a pointed dome-hke articulated élément as was shown in the figure 3B'.
The only exception seems to be the presence of an interruption in the cuticle of the exopodite of the
second leg, possibly indicating an incipient bimerous condition. The exopodite of the third leg
(fig. 30, m) although similar in outline to that illustrated by the author, shows an apparent articulation
of the distal styliform process. The endopodites of ail the legs are also similar in outline to the illus¬
trations, with a characteristic bulging curvature of the latéral margins. The apex of each is thus some¬
what narrower than the base and the 2 setae of each, which are longer than the ramus, are set with
their bases close together. The author illustrated many patches of spinules on the legs, which we
saw but did not depict in our figures. The pedifgom projection (fig. 30, o) which the author said was
achaetous we found to carry 2 minute marginal setae. The urosome in our specimen (fig. 30, p) does
not show the distinct segments depicted by the author. The caudal rami are very short lobes, each
set on the end of the urosome with a pronounced articulation. At the center of the urosome near
the anterior end there is located the insémination pore with diverging internai séminal tubes.
Source ; MNHN, Paris
Fie. 30. — EnUrocola petiti GuUle, female ; a, habitus, ventral ; b, habitus, dorsal ; o, habitus, latéral ; d, antennule -,
e, anteona ; t, labrum ; g, maxillule ; h, basal portion ol maxillule ; i, palp of maxillule ; j, maxilla ; k, first leg ;
1, second leg ; m, third leg ; n, fourth leg ; o, pediform projection ; p, urosome. Scales = 0.1 mm.
Source : Paris
82
PAUL L. ILLG AND PATRICIA L. DUDLEY
Entebocola bbementi new species
(Figures 31, 32)
Types :
Hololypic female (type locality, Callot, Channel coast of France, from Aplidium pallUiim
(Verrill) ; and paratypes from the same collection and from Tatihou, listed below.
Specimens examined :
From Aplidium palliduni (Verrill) :
“ Aplidium zostericola ”, Callot, France, Chatton iiumber 254, holotypic female and 2 female
paratypes.
"Aplidium zostericola", Tatihou, France, May, 1911, Chatton number 46, female paratype.
Description :
Female (fig. 31, 32) :
The body (fig. 31, a, b ; 32, a, b) measures 1.4 mm overall length and .98 mm in a contracted
specimen. The proportions of the uncontracted specimen, cephalosome : metasome : urosome, are
1: 6 :1.3. In the uncontracted specimen there is no evidence of dorsal duplicatures. In the contracted
individual folds of the dorsal surface simulated dorsal duplicatures. In the contracted specimen the
urosome (fig. 32, a, b) is folded and drawn into the segment of the pediform projections. In the uncon¬
tracted specimen, the urosome (fig. 31, c) does not show complété segments. Near the anterior end of
the urosome there is a midventral insémination pore and internai séminal tubes diverge from this
toward the oviducal apertures. The antennule (fig. 31, d ; 32, c) shows almost no taper. It is a single
lobe with a few setules on the anterior and apical margins. A few spinules may ornament the surface.
The antenna (fig. 31, e ; 32, d) is obscurely bimerous, with the portion corresponding to an apical seg¬
ment longer than the remainder. There is a short seta at the middle of the médial margin of the por¬
tion corresponding to the apical segment and 3 setae of intermediate length on the apex. Two addi-
tional somewhat longer setae insert on the latéral margin at the distal third and fourth. The longest
seta of the appendage is just slightly shorter than the greatest width of the appendage. The maxil-
Iule (fig. 31, e, f, g; 32, e, f) is bilobed. The lamellate basal portion has a projecting lobe (fig. 31, f ;
32, e) ornamented with spinules at about the middle of the anterior margin. Near the insertion of
the palp there is a prominence set with a seta and a setule. The apex of the medially directed lobe
of the basal portion, seen on an en face préparation (fig. 32, e) has a slightly bifid character, because
of the development of fairly strongly developed anterior and posterior tooth-like processes. There is
a row of spinules at the base of each tooth. The palp bears 5 spinulose setae on its distal margin,
separated into an internai group of 2 and a more external group of 3. At the distal third of the exter-
nal margin there is a long, narrower, unornamented seta, approximately twice as long as the longest
apical seta. The maxilla (figs. 31, h ; 32, g) is bimerous, with the basal segment massive, heavily
sclerotized and bearing at the distal médial corner a curved articulated spinulose digitiform endite. The
distal segment is bifid apicajly, ending in 2 hook-like processes. These are unornamented, with 1 of
the hooks somewhat shorter than the other. A small seta is set in an interruption of the cuticle near
the anterior margin. The legs (figs. 31, i, j, k, 1 ; 32, h, i, j, k) hâve bimerous protopodites and uni-
merous rami. Each basipodite has a small seta at the distal latéral corner. The exopodite of the first
leg (figs. 31, i ; 32, h) the second leg (figs. 31, j ; 32, i) and the fourth leg (figs. 31, 1 ; 32, k) ends in a
differentiated pointed dome-like élément. The exopodite of the third leg (figs. 31, k ; 32, j) has the
characteristic styliform outline of most species of the genus. In 1 specimen (fig. 32, j) the styliform
apex is set off by a line in the cuticle. The endopodites of the legs are subrectangular and the 2 apical
setae of each are rather widely separated from each other. The setae are consistently shorter than
the rami. The first and second legs of the endopodites are considerably longer than the exopodites.
However, in the third and fourth legs the rami are approximately equal. Paired mammiform processes
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
83
leg ; 1, fourth leg ; m, podjform projection. Scaîes = 0.1 mm.
Source : MNHN, Paris
84
PATRICIA L. DUDLEY
\
PAUL L. ILLG AND
Fio. 32. — EnUrocoh brementi, oew species, paratypic femaJe : a, habitus, ventral ; b, habitus, dorsal ; c, antcnnule :
d, antenna ; e, basal portion ol maxillule ; f, palp of maxillule ; g, maxilla ; h, first leg ; i, second leg ; j, third
leg ; k, fourth leg ; 1, pediform projection. Scales = 0.1 mm,
are found on the legs of each pair in both specimens. The pedifornt projection (figs, 31, m ; 32, 1)
is a curved plate with a circular margin. It bears a minute dorsal setule on the margin. The caudal
rami (fig. 31, c) are simple conical lobate extensions of the urosome without évident articulations.
It should be noted that ail specimens studied were from the Chatton collections and we hâve
not collected this species ourselves. The species bears a great resemblance to E. petiti, also typically
Source ; MNHI\I, Paris
ASCIDICOLIDAE AJMD SUBFAMILIES
from a species of Amaroucium but it diflers considerably in the form of the rami of the legs and in
the length o( the external seta of the maxiliular palp and the form of the antenna.
Enterocola fertilis new species
(Figures 33, 34, 35, 36)
Types ;
Holotypic female (type locality, near Port Vendres, from Amaroucium (Parascidia) areolala
Chiaje.
Near Banyuls-sur-Mer, France, holotypic female, allotypie male, paratypic females and female
paratypes listed below.
Specimens examined :
From Amaroucium (Parascidia) areolata Chiaje :
Near Port Vendres, near Banyuls-sur-Mer, France, 50-25 m, May 13, 1958, 5 females, 1 male,
including holotype, allotype, and paratypes.
“ Parasc. flavum ”, Argelès, near Banyuls, July 10, 1910, Chatton number 100, 7 females
paratypes.
Description :
Female (figs. 33, 34) :
The body (figs. 33, a, b, c ; 34, a, m) measures 1.76, 1.37 and 0.8 in a very contracted specimen.
Paired dorsal duplicatures are only indicated on the surface and are not salient. The proportions,
cephalosome ; metasome : urosome, are 1 : 5 : 2 or 1.7, in the uncontracted specimens. The urosome
(figs. 33, d ; 34, b) has folds simulating segments. Near the anterior end there is a midventral insémi¬
nation pore and diverging internai séminal tubes. The antennule (figs. 33, e ; 34, c) is a lobe termi-
nating subacutely, set with small setules. The antennule of the contracted specimen (fig. 34, n) is
slightly more developed. The antenna (figs. 33, f ; 34, d) is obscurely bimerous There is a short
médial seta at the middle of the médial margin of the portion corresponding to an apical segment.
Apically there are 3 closely-set setae of intermediate length ; 2 somewhat longer setae articulate on the
curving latéral margin at the distal fourth and fifth ; the longest seta is only slightly shorter than the
greatest width of the appendage. The labrum (fig. 33, g) is an unornamented plate with 2 spinose
lobes extending posteriorly from dorsal insertions just anterior to the posterior margin. The maxil-
lule (figs. 33, h, i) (34, e, f) is bilobed with the base extending directly medially. The palp is lamellate,
somewhat bifid distally. At the distal third of the anterior margin of the portion of the appendage
directed medially there is an unornamented lobe. Near the articulation of the palp there is a promi-
nence bearing a setae and a setule. The palp bears 5 spinulose setae on its distal truncate margin,
grouped as 2 external and 3 internai setae. At the distal third of the external margin there is an unor¬
namented seta which is about one and a half times as long as the longest apical seta. The maxiila
(figs. 33, j ; 34, g) has a massive basal segment which bears at its distal médial corner an articulated
digitiform, spinulose endite. The apical segment, which is narrower than the basal segment but also
heavily sclerotized is bifid distally, with one process somewhat shorter than the other. Set in an inter¬
ruption of the cuticle near the anterior margin there is a very short seta.
AU of the legs (figs. 33, k, 1, m, n ; 34, h, i, j, k, o, p) hâve bimerous protopodites and the basi-
podite of each bears a small seta at the distal latéral corner. The exopodite of the first legs (figs. 33, k ;
34, h, o), the second legs (figs. 33, m ; 34, j) has the characteristic styliform outline of most species of
the genus, but there is a subapical line in the cuticle setting ofT the apex of the process. The endopo-
dites are consistently shorter than the exopodites. The ramus has a more curved latéral margin than
the médial margin. The 2 apical setae of each endopodite are set closely together and are longer than
the ramus. Belween the legs of each pair there is a pair of well-developed mammiform processes.
Source : MNHN, Paris
1. second leg ; m. third Ug ; n. founh îeg o Vedii «”* '«f
0.2 mm ; other scales = 0.1 mm. ^ ’ ’ ^ projection : p. ovisac with embryos. Scales for a. b, c. p L
Source : MNHN, Paris
Fig. 34. — Etüerocola jertilU, iiew specios, paratypio female 2 (a-l), paratypic female 3 (m-p) : a, habitus, ventral;
b, urosumc ; c, antenuulc ; d, antenna ; e, maxillule ; t, palp of maxillule ; g, maxilla ; h, fîrst leg ; i, second
leg : j, tliird ieg ; k, fourth leg ; 1, pediform projection ; m, habitus, ventral ; n, antennule ; o, flrst leg ; exopodite ;
p, second leg. Scalcs = 0.1 inm,
Source ; MhSHH, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
Fio. 35. — Enteroeola fertilU, new apeoies, allotypie male : a, habitua, latéral ; b, cepbaloaome, ventral view : At =
antennule ; A2 = antenna, Mxl = maxillule, Mx2 = maxilla ; c, uroaome, ventral ; d, antennule and rostrum ;
e, antenna ; {, remuant of labrum ; g, maxillule ; h, maxilla. Scale for a = 0.5 mm ; acale for f b 0.2 mm ; other
scales B 0.1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
n Bpecies, allotypie male : a, first leg ; b, second leg ; c, third leg ; d, fourth leg ;
The pediform projection (figs. 33, o ; 34, 1) is a curved plate with subcircular margin. Set distally
on the margin are 2 separated minute setuies. The caudal rami (figs. 33, d ; 34, b) are conical and
apparently form a definite articulation with the urosome. The ovisac (fig. 33, p) has a rather strong
curvature.
The female in this species is very close to E. petüi and E. brementi, but differs from the former
by the form of the antenna, the external seta of the maxillule and outlines of the endopodites of the
legs. From the latter it dillers by the form of the outlines of the endopodites of the legs.
Male, from Amaroucium {Parascidia) areolata (figs. 35, 36) : The overall len^h measures 0.93 mm
from the tip of the cephalosome to the end of the caudal rami, excluding the caudal setae. The body
(fig. 35, a) is divided into 3 régions. The cephalosome doubtiess incorporâtes the body régions corres-
Source : MtJHN, Paris
90
PAUL L. ILLQ AND PATRICIA L. DUDLEY
ponding to the appendages through the maxillipeds, but the latter and the mandibles are lacking.
There are 4 free thoracic segments bearing the swimming legs. The urosome (fig. 35, b) is 6-segmented.
The first 2 segments bears the fifth and sixth legs. The second segment contains the bilaterally arran-
ged spermatophoral sacs. There is an unaccountable single mid-ventral slit-like aperture. The
remaining segments of the urosome are somewhat narrower than the first 2.
The rostrum (figs. 35, c, d) is elongate, tapering, and bifid terminally. The head appendages
are shown in an en face view {fig. 35, c). The antennule (fig. 35, c, d) is elongate, 7-segmented. The
basal segment is approximately a third wider than the remaining segments which show little taper.
The basal segment has a line on one surface but it is not subdivided by a real articulation. The setation
of the segments is as follows : I — 13 setae ; II — 2 setae ; III — 2 setac ; IV — 1 seta ; V — 1 seta ;
VI — 1 seta ; VII — 8 setae, 1 aesthete.
The antenna (fig. 35, e) is trimerous, the articles having the proportional lengths of 1.5 : 1 : 1.
The basal article has a long seta at the distal inner corner. The second article is unarmed. The ter¬
minal article bears an articulated hook and 4 short setae.
The labrum (fig. 35, f) is obviously degenerated and consists of a crumpled lobe, without setae
or palps.
The maxillulc (fig. 35, g) is uniramous, and obscurely bimerous. The base has lost tlie médial
projections of earlier copepodids and the terminal portion is fan-shaped, with 6 long, plumose setae
uniformly spaced on the distal margin. The setae are not clearly articulated with the appendage liul
extend directly.
The niaxilla (fig. 35, h) is uniramous and unimerous. It terminâtes in a long, unarticulalcd,
plumose seta, which lacks a clear articulation.
The first leg (fig. 36, a) is biramous, with bimerous protopodite, trimerons endopoditc and
bimerous exopodite. There is a trapezoidal intercoxal lamella. The coxopodite lacks armature or
ornamentation. The basipodite bears a short seta on the latéral margin and a distal médial spinc,
which reaches slightly beyond the distal margin of the first article of the endopodite. A cluster of
hairs is found just medial to the spine. The 2 basal articles of the endopodite are unarmed. The
terminal article bears 2 latéral setae, 2 terminal setae, and a medial seta. The basal article of the
exopodite bears a distal latéral spine. The second article has a distal latéral spine, a terminal spine,
a terminal seta and 2 medial setae.
The second leg (fig. 36, b) is biramous with bimerous protopodite, trimerous rami. The coxo-
podites are joined by a subrectangular intercoxal lamella. The basipodite bears a seta on the latéral
margin. The basal article of the endopodite bears a short seta at the distal medial third. The second
articles has 2 medial setae. The terminal article bears a latéral seta, 3 terminal setae, and 2 medial
setae. The basal article of the exopodite bears a distal latéral spine. The second article bears a distal
latéral spinc, a medial seta. The terminal article bears 2 latéral spine, a terminal spine, a terminal
seta, and 3 medial setae.
The third leg (fig. 36, c) is much like the second except that the basal segment of the exopodite
bears a short medial seta and the terminal segment bears 5 setae rather than 4.
The fourth leg (fig. 36, d) is similar to the 2 preceding except that there are only 5 seta on the
terminal article of the endopodite rather then 6, and the exopodite is as in the third leg.
The fifth leg (fig. 36, e) consists of a bimerous setiferous plate-like lobe on the latéral ventral
surface of the first urosomal segment. It is not readily possible to détermine if the plate is proxirrially
articulated because it is so strongly outlined. There is an additional setiferous projection far laterally
on the segment which perhaps représenta a basal portion of the usual fifth leg.
The caudal ramus (fig. 35, b) is broad and fiat, slightly longer than the anal somite. There are
4 setae on the truncate terminal margin, a short dorsal seta at the distal fifth, and a short seta set
in an emargination at the middle of the latéral margin.
We find a very good correspondence of our specimen with the account of Canu for the male
of E. fulgens, which is described in general terms. Since we hâve not had access to a specimen of the
latter for detailed anatomical comparison we are not able to provide a spécifie discrimination at this
point.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
9i
Enterocola laticeps, new species
(Figure 37)
Types :
Holotypic female, (type locality, San Juan Islands, Washington, in Styela gibbsii (Stimpson)) ;
paratypes, ail specimens listed below.
Specimens examined :
From Washington :
From Styela gibbsii (Stimpson) (ail specimens less than 7 mm in diameter) :
Mixed collection, Harney Channel 48o34.7' N, 122®53.1' W, 30-40-15 fathoms, P. L. Dudley,
June 20, 1964, 2 females.
Blakely Island West, 122^50.1 W, 48®33.3' N, 16-18 fathoms, P. L. Dudley, June 20, 1964,
4 females.
Blakely Island West, 16-18 fathoms, J. Nolfi, August 15, 1966, 1 female.
From British Columbia, Canada :
From Styela gibbsii (Stimpson) :
Nanoose Bay, shallow dredge, R. Campbell, August 9, 1964, 1 female third copepodid.
Description :
Female (fig. 37) :
Body (fig. 37) 3.1mm, 2.8 mm, 2.5 mm, 2.0 mm and 1.5 mm, total length from 5 available
specimens measured from the anterior margin to the end of the caudal rami. The body is very heavily
sclerotized overaU, with strong démarcation into cephalosome and metasomal and urosomal segments.
The cephalosome is broad and rather fiat and there is no rostrum. The major body articulation lies bet-
ween the fourth and fifth leg-bearing segments, so the metasome is composed of 4 well delimited seg¬
ments. The urosome (fig. 37, b) is very distinctive in configuration, 5-segmented. The first urosomal
segment expands laterally as conspicuous, larged curved lamellae, which are unarticulated but which
probably include the fifth legs. On the same segment at the ventral midbne is a conspicuous lobed
sclerotization associated with insémination pore. From this diverging tubes lead to latéral séminal
réceptacles at the oviducal apertures. The remaining urosomal segrfients bear no appendages ; the
caudal rami are well developed and bear a characteristic armature. The urosome is generally covered
ventrally with broken rows of spinules.
The head appendages are described from an en face préparation (fig. 37, c) so the terms of orien¬
tation refer to this and not to the basic anatomical relations on the body.
The anteimule (fig. 33, d) is unimerous, but the pattern of sclerotization indicates 4 ingrédient
segments. The portion representing a basal article bears 1 seta and 2 setules distally on the anterior
margin. The short second component has 2 anterior setae ; third 2 setae ; the terminal component
bears 9 setae around the apical and posterior margins.
The antenna (fig. 37, e) is clearly bimerous, heavily sclerotized and with distinctive sclerotized
pataches on the basal article. The basal article is longer than the terminal article and lacks armature.
The terminal article bears apically 5 setae, 1 much longer than the others.
The labrum (fig. 37, f) is heavily sclerotized and bilobed. It bears 2 spinulose, extremely heavy
setae laterally and dorsally inserted. The general ventral surface has rows of denticles. There is
a complicated postoral apparatus on the body surface (fig. 37, c).
The maxillule (fig. 37, g) is bilobed with the articulation on the head very complicated. The
basal portion is heavily sclerotized and considerably modified. It is produced medially to terminale
in heavy lobes and there is a latéral subterminal thumb-like process, which has rows of heavy spinules.
At the base of this process there is inserted a stout seta ornamented with long spinules. The palp is
flattened and bilobed, 1 lobe with 2 spinulose seta, the other with 3 terminal setae, 2 of these stout
Source ; MNHN, Paris
v.nt,.| ; À2““ri™."L‘ ‘ ‘l P"*!»»"” projection.,
Mx2 = maxiUa, shaded ; d, antennule ; e antenna • f lâhn.m antenna, L _ labrum. Mxl = maxillule unshaded,
or .eoond lep k, third l.g, 1. exopodit. ot ihird lej'- txToJiU S Wh i ! j' «"l '•» i i, exopodit.
1.0 mm ; other Bcales = 0.1 mm. ® ’ “P caudal ramue. Scale for a s:
Source : MNHN, Paris
ASCIDICOLIDAE
SUBFAMILIES
93
and covered with heavy spinules, lhe third unornamented. There is an additional short seta at the
proximal fourth of the ventral margin. The latéral face is heavily sclerotized and is ornamented with
rows of heavy spinules.
The maxilla (fig. 37, h) is bimerous and heavily sclerotized. The basal segment bears an endite
with a spinulose thumb-like process. The articulation of the component articles is very complicated.
The distal article ends medially in 2 pointed spinulose, heavily sclerotized processes. There is a single
seta, with a much expanded base on the anterior margin.
The first leg (fig. 37, i) is biramous and its articulation with the body is rather weakly delimited.
The entire appendage is very heavily sclerotized. The coxopodite has a pointed protrusion at the distal
latéral corner. Rows of spinules form the ornamentation of the anterior surface. The basipodite
tapers somewhat and bears a seta at the middle of the latéral margin. The anterior surface bears
rows of spinules. The endopodite is bimerous. The basal article lacks armature but has 2 rows
of spinules on the anterior surface. The distal article is slightly tapered to an apex bearing 2 setae,
one about one-fourth longer than the other. Short lines of spinules ornament ; the anterior surface.
The exopodite is obscurely bimerous and very small in proportion to the endopodite, equalling about
half its length. The ramus tapers and bears on the truncate apex a broad, flattened spine. The basal
article has no armature or ornamentation.
In most specimens examined the second leg is just like the first. In one case the exopodite
is definitely bimerous and there is a small spinule at the distal latéral corner of the basal article {fig. 37, j).
The third leg (fig. 37, k) is much like the first and second with the following exceptions. The
basipodite is subrectangular rather than trapezoidal and the rami are both borne on the terminal margin.
The exopodite is strongly bimerous and there is a heavy spine-like element at the distal latéral corner
of the basal article. The features of this leg are in strong contrast to the structure in most species of
Enterocola, where the third leg differs in a characteristic way from the others.
The fourth leg is in most cases like the third. In 1 case 1 exopodite (fig. 37, 1) has 2 flattened
terminal spines, contrasting with its partner (fig. 37, m).
The pediform projection of the sixth thoracic segment (fig. 37, b) is a very large curved flap
which reaches almost to the end of the urosome. There are 3 minute setae spaced along the posterior
margin, possibly indicating involvement of the fifth leg.
The caudal ramus (fig. 37, n) is flattened, almost rectangular and slightly longer than the anal
somite. It has 3 terminal lobes and an apical flattened, short seta, 1 subterminal dorsomedial setule
and a minute knob-like element on the latéral margin at the proximal third. Rows of fine spinules
ornament the dorsal surface.
The animal is eyeless. It is usually a glistening white, but may acquire a brownish tinge from
gastric sécrétions of the host. There may also be orange streaks on the dorsal surface. AU specimens
were taken from the stomachs or proximal intestine of specimens of Styela gibbsii less than one centi-
meter in diameter.
No males hâve been found so far.
This species has been dealt with in an extended description because of its many différences from
the species of Enterocola associated with compound ascidians. In its habitus it definitely approaches
the other species, with the body eruciform ; the head well demarcated, flat and broad ; in the presence
of expansive pediform projections on the sixth thoracic somite ; and with the urosome much shorter
than the anterior body. The elongate legs and the well-developed caudal rami are perhaps fairly
obvioua différences. In more detailed characters, the distinction becomes considérable. The anten-
nule has indications of 4 segments and it bears more setae than does any previously known species
of Enterocola. The antenna, although bimerous as in most species, has the basal article much longer
than the terminal and there are only 5 setae on the latter, as opposed to the consistent appearance in
other species of 6 or 7 setae. The most distinctive différences are in the characters of the legs, and
lhese also differ from those of Lequerrea and Enlerocolides. Although the 4 thoracic legs are extremely
large in proportion to the body, the ventral lamelliform or mammiform processes characteristically
associated with the legs of many other species do not occur here. Further, the notably differentiated
third leg is not encountered in the présent form.
Source ; Mt-JHN, Paris
94
PAUL L. ILLG AND PATRICIA L. DUDLEY
The latéral seta on the basipodite on each leg is not found at ail in other species nor does any
other hâve the endopodites clearly bimerous throughout. The exopodites are clearly related to the
basic plan of legs 1, 2, and 4 of Enterocola species, but differ by élaboration of an auxiliary process of
the basal article on the third and fourth exopodites. The exopodites of the third and fourth legs of
Lequerrea show great similarity to these, but in Lequerrea the processes of the basal articles hâve no
articulations. The endopodites of Lequerrea are quile dissimilar, being unimerous and without arma¬
ture, as again arc those of ErUerocolides. The pediforin projections of the sixth thoracic segment bear
3 setae, and none other of the enterocolins has more than 2 here ; in many species of Enterocola there
are none. None of the other enterocolins hâve comparably developed caudal rami and ail lack armature
on the ramus ; in this species there are at least 2 setae represented, although these are modified. It is
probably considerably significant that our species is symbiotic in a solitary ascidian. AU species of
Enterocola for which the host is known corne from aplousobranch compound ascidians, mostly from
the tamily Polyclinidae, with 2 species from Didemnidae and 1 from Clavelinidae. Lequerrea, which
also reaches large size, is known only from species of the simple ascidian Polycarpa. We are somewhat
inclined to suspect that the so far poorly understood species, E. bilamellata Sars and E. selifera Hansen,
although they were found free in bottom trawls, may turn out to hâve solitary ascidians as hosts.
INDETERMINABLE SPECIES
Biûcryptus flavuH Hesse, 1865, p. 244-247, pl. 6, figs. lc-6c (no locality, in compound tunicate attached
to Cystoseira fibrosa). — Enterocola flavus, Gerstaccker, 1870-71, p. 801. — Hartmeyer, 1911.
Biocryplus roseus Hesse, 1865, p. 242-244, pl. 6, fig. IB-lOB (no locality, in social tunicate attached
to frond of Zoslera). — Enterocola roseue, Gerstaecker, 1870-71, p. 774, 801. — Hartmeyer,
1911, p. 1736.
Biocryplus calthaeus sp. n. Hesse, 1872, p. 29-30 (unidentiHed tunicate, no localitv). — Hartmeyer,
1911, p. 1736.
SUBFAMILY ENTEROPSINAE AURIVILLIUS 1885
Enteropsidae Aiirivillius, 1885a, p. 236-237. — 1885b, p. 282. — Shimkevich’, 1889, p. 76.
Enteropsidés Canu, 1892, p. 30, 107.
Ascîdicolidae Gerstaecker, 1870-1871, p. 719 (part), — Canu, 1891, p. 472, 474, 475 (part) ; 1892, p. 30,
107, 186 (part). — T. Scott, 1901b, p. 241-245 (part). — Caïman, 1908, p. 177, 182 (part). —
ChattOD & Brément, 1909b, p. 196 (part).
AscidicoUnae Schellenberg, 1922, p. 219-220 (part).
Aplostomiens Canu, 1886a, p. 373-374 (part).
Enterocolidae Sars, 1921, p. 73-74 (part). — Gray, 1933, p. 523 (part). — Gotto, 1953, p. 659 (part) ;
1954, p. 659. — Bresciaiii & Lützen, 1962, p. 376 (part). — Monniot, 1965, p. 160 (part). —
Dudley, 1966, p. 155, 160 (part). — Gotto, 1966a, p. 193 (part). — Stock, 1967a, p. 9 (part).
When Aurivillius (1885a) described the genus Enteropsis, he proposed a family for it also ;
he was struck by the distinctive features of the copepod. In the relatively few publications concerniug
members of the genus there has been very little considération for the familial position. The very broad
treatment of Canu (1892) and Schellenberg (1922) included the genus in the Ascidicolidae and Asci-
dicolinae, respectively. Sars (1921) placed it in the Enterocolidae, with Mycophilus (his spelling),
Chatton & Harant, (1922, p. 163) recognized the family but termed it the Enteropsidés and assigned
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
95
the much better known Mychophilus also to the family. The latter often has been confused with
EiUeropsis and indeed Chatton & Brément (1909c, p. 239-240) point out that the généra probably
should bc combined, We are following their precedent in conserving the genus Mychophilus (see p. 103)
but it is obvious that the diflerences belwecn Mychophilus and the sériés of Enteropis species now
known are not profound ones. Accordingly, the subfamilial diagnosis is a relatively cohesive one.
We bave stated (p. 15) that we consider it necessary to place Enleropsis pilosus Canu as a species incerta
sedis in the family Ascidicolidae, so do not account for its characters in our diagnosis of the subfamily
Entcropsinac. Canu has many garbles in the description of E. pilosus. We strongly suspect he had
a species of Enleropsis betore him but that he mixed in some details deriving from a haplostomin.
In the female the body is eruciform to fusiform, inflated, with segmentation usually not obvious,
lendiiig to indication only by indentations and the positions of the legs ; without pterostegites and
without pediform projections of the sixth thoracic segment (that is, without any indication of fifth legs).
The démarcations of the 2 posterior régions of the body tend to be obscure. The habitus and posture
may be highly distinctive, The size ranges from minute in Mychophilus to very large dimensions in
species of Enleropsis (length up to 8 mm).
The cephalosome does not form a distinct articulation with the body but is typically rallier well
set off, hy a constriction or by a sharp inflection, and may be strongly indicated laterally by pleural
folds. The appendages are the antennules, antennae, maxillules, maxillae. Practically ail the litera-
lure is erroneous in référencé to these ; the maxillules are usually reterred to as mandililes, and in
inany cases the maxillae are termed maxillipeds.
The metasome effectively consists of the segments of the 4 major thoracic legs, and the démar¬
cation from the urosomc is obscure. In the absence of fifth legs or of the corresponding pediform pro¬
jections, it is not possible to déterminé whether the corresponding segment is metasomal or iirosomal.
The metasome is the widest and longest body région. The legs protrude from it conspicuously, without
any distinct articulation.
Although the urosome may appear annulated, the segmentation is usually obsolescent. The
anterior portion is marked by the présence of the génital apparatus. Therc is usually a midventral
insémination pore which may be seen te connect with diverging internai tubes which loop anteriorly
then laterally toward the oviducal apertures. The latter are usually marked by various sclerotiza-
tions and sometimes by setules. The caudal rami are always reduced and vaguely articulated on the
body and vary in the presence or absence of a fceble setation.
The antennule varies from unsegmented to obscurely Irimerous. The setation is always somc-
what reduced, or may be absent. There are at most 10 setules and setae, mostly borne on the terminal
article.
The antennae are modified appendage, always much tapered and varying in segmentation.
Up to 4 articles may be indicated ; most forms hâve the appendage bimerous. There may be actual
setae on the appendage, but usually it tends to prolongation in 1 or 2 setiform processes. It is very
characteristic that circular patches of spinules occur, sometimes on actual lobes on the surface of the
appendage.
The labrum is rather central on the head and typically conspicuous. The subterminal margin
may be unornamented, but in many species is set with a row of spinulose setiform éléments, ranging
in number from 2 to 9 in various species of Enleropsis. There is no mandible in the adult.
The maxillule is a complex structure of obscure anatomical dérivation in which articulations may
be suppressed. There are always 2 prominent lobes, one usually articulated. The armature is various,
ranging from fully artictilated setae to setiform processes, or to obsolescence with vestiges as lobe-
like projections. There is a tendency for the basal médial process rather consistently to hâve 2 élé¬
ments of armature. The palp-like lobes are more various and may hâve 2 or 3 éléments of armature.
The maxiUa is a massive appendage, the greatest bulk representing a basal article. In Mycho-
pkilus there is a small subapical spine ; in Enleropsis there is a terminal article forming a complicated
articulation on the basal article.
There is no maxilliped, but there are many published référencés to such an appendage due to
misconceptions as to the head appendages.
Source ; MNHN, Pans
96
PAUL L. ILLG AND PATRICIA L. DUDLEY
The 4 pairs of thoracic legs are ail uniramous and bimerous. The basal article is vaguely arti-
cuiated with the body and strongly articulated terminally. There is no armature on this article. The
terminal article has a characteristic apical hooded hook with an opposed process.
ScHELLENBERC înterpretcd in Enteropsis onychopkorus 2 feeble latéral prominences at the pos-
terior margin of the metasome as fifth legs. No other forra in the subfamily has fîfth legs and we
consider Schellenberg’s interprétation as dubious.
There are complicated structures associated with the oviducal aperatures on the sides of the
urosome, which may include sclerotizations and, in a few cases, setules. The adult male is not known
in the subfamily, but see below (p. 105) our discussions of male fifth copepodids in Enteropsis and
Mychopküus. It is expected that males for this subfamily will be readily diagnosable on the basis
of the mouthparts, which probably will correspond well in basic structure with those of the females.
The subfamily shows close affînity with the Enterocolinae, sharing with the latter many of the
tnost distinctive anatomical adaptations. Many astute observera (e. g. Sabs, Gotto, Monniot, Stock)
hâve referred one or both of the enteropsin généra to the “ Enterocolidae
Many published misconception as to the head appendages in Enteropsis stem frora misapprehensions
of Canu with regard to his Enteropsis pilosus. Ail of his conclusions as to structures in this animal
seem to hâve derived from purely anatomical observations on the adult Futhermore, as stated above,
his original description would seem to présent features derived from an undoubted Enteropsis plus
some others from some other ascidicolid. Curious, Canu presented several times subséquent to the
original description exactly the same figured représentation and seems never to hâve made fresh obser¬
vations. The ambiguities in his présentation hâve been obvions to ail subséquent students and it is
very strange that they escaped his keenly observant eye.
Using purely anatomical criteria he termed the mouthparts of Enteropsis the mandibles and
the second maxillae. As to the mandible he was definitely mistaken, a most curious circumstance
because his démonstration, on the basis of observations on a sériés of developmental stages that the
exactly équivalent appendages in Enterocola are the maxillules was an outstanding tour de force. These
appendages in the two généra are by no means identical but they are much alike and both differ so
siibstantially from any mandible known among ascidicoles that it is most surprising that he was not
struck by the discrepancy.
As to the “ second maxillae ” he was right, but for wrong reasons. His terminology would
hâve as équivalent to this appendage in Enteropsis a similar prehensile appendage in Enterocola, and
lhe strongly prehensile mouthpart of Haplostoma, calling them ail “ simple second maxillae ”. This
terminology stemmed from his misconception, derived curiously enough, from his very perspicacious
observations on development, that the second maxillae of copepods develop in bipartite fashion, the
endopodite and exopodite respectively evolving as separate lobes from the general body surface. He
never used the term “ maxilliped ”, nor considered that such an appendage belonged to the complé¬
ment of copepod appendages. In modem terminology his “ external second maxilla ” is the true
maxiUa, his internai second maxilla ” is the maxilliped, and they indeed develop as the separate
lobes he observed.
In the case of Enterocola he saw a single lobe develop behind the maxillule so termed it a “ simple
second maxilla This was a fortunate term because the appendage thus designated conforms to
current terminology. His choice was less fortunate in the case of Haplostoma where again he found
a single lobe and termed it a “ simple second maxilla ” ; In this instance current information identifies
this appendage as the maxilliped.
In his description of adult anatomy of Enteropsis pilosus he compared the structures to those
of Aplostoma He was correct in terming the first mouthpart of Haplostoma the mandible, but
wrong in the case of Enteropsis. He then compared the second mouthparts in the two and considered
them as counterparts. They are not, but here it was his erroneous terminology in calling them both
second maxillae which had the resuit of applying the right term to Enteropsis for the wrong reason.
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBPAHILIES
97
KEY TO FEMALES OF GENERA OF ENTEROPSINAE
1. Anus displaced dorsally to just posterior to thorax at asterior end of urosome ; maxilla a mas¬
sive unarticulated lobe with articulated subapical spine. MychophÜus
1'. Anus terminal on urosome ; maxilla distinctly bimerous, with massive base and well developed
distal article directed medially. EnteropaU
Mycuophilus Hesse, 1865
Mychophilus Hesse, 1865, p. 233-236, 255 (no type species designated ; type species, Mychophilua
roseus, by reference to its synonym M. curvatus Ghatton & Brément, improperly designated as
type by Ghatton & Brément, 1909). — Gerstaecker, 1870-1871, p. 719. — Ghatton & Brément,
1909d, p. 235-236, 238 ; 1910, p. 91-92. — Hartmeyer, 1911, p. 1736. — Ghatton & Harant,
1922b, p. 163 ; 1922c, p. 249. — Schellenberg, 1922, p. 292, 294. — Harant, 1931, p. 372. —
Wilson, 1932, p. 602. — Neave, 1939, p. 234. — Sewell, 1949, p. 183, 189. — Stock, 1967a, p. 9.
Myehephilus, Canu, 1892, p. 216.
Mycophilua Sars, 1921, p. 78-79. — Gray, 1933, p. 525-527. — Lang, 1948, p. 16-17, 22. — Gotto, 1952,
p. 674 ; 1954, p. 659 ; 1960, p. 216 ; 1961, p. 151. — Bresciani & Lützen, 1962, p. 376. — Gotto,
1966a, p. 193.
Michophilus Ghatton & Harant, 1922b, p. 163.
linieropais, T. Scott, 1901b, p. 241-242 (part) ; 1907, p. 369 (part).
Adranesiua, Ghatton & Brément, 1909a, p. 202 (part).
Gbatton & Bbéhent, 1909d, pronounced the genus of Hesse recognizable, but claimed the
2 species he described were indéterminable. They then designated as type of the genus the species
tiiey described from their own material, Mychopilus curvatus Ghatton & Brément, 1909. This dési¬
gnation is contrary to the International Gode of Zoological Nomenclature (1961). In any case
the spécifie name is a synonym of Hesse’s species M. roaeua.
This genus has been studied by us through bibliographie sources and from some specimens
coUected at Plymouth, England and Banyuls-sur-Mer, France. Because all of the appendages of
M. roseus hâve never been completely described, we hâve had to obtain anatomical details to compare
with the species of Enteropsis in order to détermine the taxonomy of the subfamily Enteropsinae. There
remain many probleras involving the biology, the life history, the morphological variation, and perhaps
genetic différentiation within this genus, as pointed out especially by Lanc (1948) and Gotto (1954).
Although several speciBc names hâve been proposed in the genus, all referred to M. roaeua
lintil 1967, when Stock described a second species.
The characteristics of the genus are essentially those of the subfamily Enteropsinae, discussed
above, with the salient additional distinctive features utilized above in the key to the généra.
The body form of Mychophilus is highly distinctive, with the displacement of the anus and the
élongation of the urosome proportional to the metasome. In M. roseus there is a tendency to extreme
latéral displacement of the first legs. In most of the characters of the appendages the différences from
forms of Enteropsis seems to be only at about the spécifie level. In the maxillae, however, there is
a deflnite distinction. The males of Enteropsis and Mychophilus would seem to approach each other
even more than the females but the séparation can undoubtedly be made at the spécifie level, even
on the fifth copepodid males now avaüable.
Source : MNHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
KEY TO SPECIES OF MYCHOPHJLUS, BASED ON FEMALES
1. Antenna distinctly not prehensile ; urosome and cephalosome-metasome roughiy subequal....
rosens Hesse, 1865
1'. Antenna prehensile ; urosome much exceeding the cephalosome-metasome.. fallax Stock, 1967
Mychophilus roseus Hesse, 1865
(Figures 38, 39)
Mychophilus roseus Hesse, 1865, p. 232-235, pl. VI, fîgs. 1-8 (type locality, coast of Finistère, France,
from compound tunicate attached to Zostera murina). — Gerstaecker, 1870-1871, p. 774, 801.
— Hartmeyer, 1911, p. 1736. — Chatton & Brément, 1909d, p. 234, 238 (indéterminable species).
— Stock, 1967, p. 9.
Mycophilus roseus Sars, 1921, p. 79-80, pl. 36, fig. 2. — Lang, 1948, p. 3, 17-22, fig. 7-14, 15-17. —
Sewell, 1949, p. 189. — Gotto, 1952, p. 674 ; 1954, p. 659-665, 6g. 105 ; 1960, p. 216 ; 1961,
p. 151. — Bresciani & Lützen, 1962, p. 376. — Gotto ,1966a, p. 193.
Mychophilus pachygasler Hesse, 1865, p. 235-236 (type locality, coast of Finistère, France, from com¬
pound ascidian). — Gerstaecker, 1870-1871, p. 774, 801. — Chatton & Brément, 1909d, p. 238
(indéterminable species). — Hartmeyer, 1911, p. 1736. — Stock, 1967a, p. 9.
Mycophilus pachy gaster, Sewell, 1949, p. 189.
Mychophilus curoatus Chatton & Brément, 1909d, p. 236-238, 6g. 1 (type locality, Banyuls-sur-Mcr.
Golfe du Lion, France, from Polycyclus renieri], — Schellenberg, 1922, p. 292, 295. — Harant,
1931, p. 371. — Stock, 1967a, p. 9.
Mycophilus curvatus, Sewell, 1949, p. 183, 189.
Mycophilus rosovula Gray, 1933, p. 523-528, 6gs. 1-2 (type locality, Millport, Scotland, from Botryl-
loides leachii Savigny). — Stock, 1967a, p. 9.
Enteropsis oararensis, T. Scott, 1901b, p. 241-242, pl. XVII, figs. 28-34 (type locality, Moray Firth,
Scotland, from Botryllus sp.) ; 1907, p. 369.
Adranesius vararensis, Chatton & Brément, 1909b, p. 202. — Mychophilus vararensis, Chatton &
Brément, 1909d, p. 239. — Stock, 1967a, p. 9.
Distribution :
European west coast ; British Isles ; Mediterranean.
Hosts :
Botryllus scklosseri (Pallas) B. smaragdi, B. violaceus ; Polycyclus renieri Lamarck ; Botrylloides leachi
(Savigny).
Specimens examined :
From Botryllus scklosseri (Pallas) :
Off Cap Béar, Anse de Paulilles, near Banyuls-sur-Mer, France, 50-20 m, May 28, 1958, imma¬
ture male.
Nord et nord-est Port-Vendres, near Banyuls-sur-Mer, France, mixed trawl, 70 m and 30-60 m,
May 6, 1958, 2 females.
From Botrylloides leachi (Savigny) :
Gaiola, Bay of Naples, Italy, 35 m, February 20, 1958, 1 female.
Source : MNHN, Paris
F,O. 3g. Mychopküus roseus Hes»e. femaie : a, habitus, latéral ; b, en face view of cephalosome : Al = antennule,
A2 = anteoua, Mxl = maxillule, Mx2 = maxilla ; c, sclerotized area on dorsal cephalosome and antennule, d,
antenna, ventral ; e, antenna, dorsolateral ; f, maxillule -, g, maxillule ; h, palp of maxillule ; i, maxilla ; j, maxilla ;
k, apex of maxilla ; I, first leg. médial ; m. distal article of second leg ; n, distal article of third leg ; o, fourth leg,
anterior ; p. génital aperture and ornament ; q, caudal ramus. Scale for a = 0.2 ram ; for b = 0.1 mm ; other
scales = 0.05 mm.
Source : AlfJHM, Paris
100
PAUL L. ILLG AND PATRICIA L. DUDLEY
Fig. 39. — Mychophilus roseua Hesse, male fifth copepodid ; a, habitus, latéral ; b, urosome, ventral ; c, antennule ;
d, antenna ; e, maxillule, basal portion ; f, maxillule, palp ; g, first leg ; h, second leg ; i, fourtb leg ; j, anal somite
and caudal ramus. Scales ss 0.1 mm.
Duke Rock, near Plymouth, England, 6 m, September 8, 1958, many females.
From Polycyclus renieri Lamarck :
Near Banyuls-sur-Mer, France, September 26, 1910, Chatton number 212, 60 females.
From Polycyclus sp. :
Near Banyuls-sur-Mer, France, October 24, 1910, Chatton number 136, 1 female.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
101
Description ;
Adult Female :
The overall lengths of 2 specimens from anterior ends to ends to ends of urosome were 1.49 mm
and 1.40 mm.
The body (fig. 38, a) has been described and fîgured many times. It is notable for the curved
posture, the elongate cylindrical form and the unique recurvature of the intestine, with conséquent
displacemcnl of the anus dorsally to the anterior end of the urosome. The latéral displacement of
the first legs, rcffered to particularly by Lang (1948) and by Gotto (1954) is notable.
The head appendages are described below from the orientation on an en /ace'préparation
(fig. 38, b) so the terms of orientation used are pertinent only to this préparation and not necessarily
to the general anatomy of the body.
There is no rostrum, but on the dorsal anterior part of the cephalosome there is a strongly
sclerotized area (fig. 38, c) and 4 setiiles emerge from pits in the surrounding unsclerotized cuticle, at
the corners,
The antennulc (fig. 38, c) is obscurely articulated with the cephalosome, it is unimerous, flalte-
ned and has a slighl taper. Heavy scierotizations form a distinctive pattern on the surface. Termi-
nally there are 4 large, flattened, spiniform projections. Two setules, 1 on the dorsal face, 1 on the
ventral face, emerge from pits. There is a general covering of small protubérances.
The antenna (figs. 38, d, e) is bimerous, with a strongly sclerotized fold separaling the articles
both dorsally and ventrally. The basal article has 2 setules emerging from pits on the ventral proxi¬
mal surface. Several patches of spinules also ornament this surface. The distal article is essentially
conical, but the pointed tip actually probably represents a fused apical seta. There is a short articu¬
lated spiniform process on the posterior margin just proximal to the apical projection.
The labrum is an unornamented hemispherical lobe.
The roaxillule (figs. 38, f-h) is bilobed, with a médial basal prolongation and fused setiferous
lobe representing the palp. The entire appendage is relatively heavily sclerotized. There are 2 rows
of spinules on the ventral face of the base. The palp portion is stout and somewhat flattened. On
the ventral surface there is an apical dépréssion, supported to either side by scierotizations, so that in
views from some aspects the tip appears bilobed {fig. 38, g] in others appears entire (fig. 38, f). The
entire ventral surface is covered with spinulose stout pointed projections. In some specimens (fig. 38, h)
this apex appears as 1 such projection and rounded spinulose lobe. On our specimens there is no
large seta basal to the palp, as was described and figured by Chatton & Brément (1909d).
The maxilla (figs. 38, i, j, k) is unimerous, indistinctly articulated on the body and consists of
a lobe with a single apical medially directed, articulated spine. There is no other ornamentation or
armature.
The 4 pairs of legs are much alike (figs. 38, l-o). Each leg is bimerous and uniramous. The
whole appendage is covered with generally distributed fine protubérances. Scierotizations form a
distinct ve pattern on the articles. The basal article has 2 setules emerging from pits in the sclero-
tization on the ventral survace. The distal article has a complicated apical structure. There is a
médial lobe, opposing an articulated spine, which is invested, when retracted, by a cuticular covering.
On the second leg, there is a setule at the base of the spine, which is lacking in the other legs (fig. 38, m).
There seem to be no fifth legs.
At approximately the level of the anus, on the sides of the urosome, complicated structures
perhaps involving the sixth legs are situated at the oviducal apertures (fig. 38, p). Each consists of
2 sclerotized pièces. The more anterior of these is bilobed and it is somewhat exceeded anteriorly
and dorsally by the more posterior piece. A membrane between the pièces is pierced by the oviducal
aperture and anterior to this on the membrane, a small setule is inserted.
The caudal rami, exceedingly small, are at the apex of the urosome. Each (fig. 38, q) is a cône
with no ornamentation or element of armature. It is a reraarkable feature of the genus that the anus
is displaced completely away from the bases of the rami.
Source : MNHN, Paris
102
PAUL L. ILLG AND PATRICIA L. DUDLEY
The many females, taken near Plymouth, in Bolrylloides leachi, were ail found in the branchial
baskets of the hosts.
The literature provides a iiumber of statements as to localization within the hosts of the cope-
pod. Chatton & Brément reported they found specimens at Banyuls in the matrix of colonies of
the ascidian hosts. Scott found his form in branchial cavities of a species of Bolryllus. Schellen-
BERG added a finding in the cloacal cavity of a compound ascidian. Gray stated his specimens occu-
pied canal Systems in the matrix of Bolrylloides leachi. Gotto, 1954, p. 665, provided the summary
statemciit that “ The précisé relationship of Mycophilus {sic) to its host remains problcmatical
Male fifth copepodid (figs. 39, a-j) :
The body form (fig. 39, a) is cyclopoid, with the qualification that the habitus is distinctly
copepodid (immature) and the exact configuration of the adult cannot directly be inferred. The ove-
rall length, measured from the apex of the head to the end of a caudal ramus is 0.92 mm. The seg¬
mentation is well defined ; the cephalosome iiicludes the appendages through the maxilla. The man-
dible and maxilliped are absent, but the corresponding body régions are incorporated into the cepha¬
losome. There are 4 frce thoracic segments. The urosome (fig. 39, b) is 5-segmented. The first
segment bears the minute fifth legs, and developing spermatophoral sacs are visible in the second
segment. The third and fourth segments bear rows of spinules on the ventral surface.
We did not observe whether there was a rostrum, before dissecting the specimen.
The antennule (fig. 39, c) is 6-segmented ; the basal article is about 2 times as long and 2 times
as wide as any other article. The terminal 5 articles are ail of approximately the same width, but
vary in length. The setationis as follows ; Article 1 —11 setae ;II — 2;III — 2; IV — 2;V — 2;VI —
8 setae, 1 setule, 1 aesthete. Two rows of spinules ornament the anterior surface of the basal article.
The antenna (fig. 39, d) is obscurely trimerous. The basal article bears an elongate seta which
reaches to the distal fourth of the appendage. The second article is unarmed and unornamented.
The terminal article tapers to a point and is longer than the other 2 articles combined. It is orna-
mented with rows of spinules and a row of spinules lies on the apex.
We did not inake observation on the labrum.
The maxillule is bilobed. The palp (fig. 39, f) is a lamella bearing a short seta on the margin
and terminâtes in 2 apical setiform processes, these approximately one and a half times longer than
the palp proper. The surface of the appendage are ornamented with rows of spinules. The basal
portion of the appendage extends medially to form a tapered lamella, terminating in 2 setiform pro¬
cesses, these much shorter than the lamella (fig. 39, e).
The maxilla was not observed, because of difficulties in dissection.
The first leg (fig. 39, g) is biramous, with bimerous protopodite, trimerous endopodite, bimerous
exopodite. There is a bilobed triangular intercoxal lamella. The coxopodite is elongate, approxi¬
mately equalling in length the remainder of the appendage. The basipodite lacks the distal médial
spine characteristic of most notodelphyids and occurring in some ascidicolids. The usual latéral
seta is présent. The endopodite is about one-fourth longer than the exopodite. The basal segment
of the endopodite has no setae, the second segment bears a distal médial seta, the terminal segment
bears a latéral, 2 terminal, 2 distal medial setae. The basal segment of the exopodite bears a distal
latéral spine ; the terminal segment, which is about 2 times longer than the basal segment, bears a
distal latéral spine, a terminal spine, a terminal seta, 2 medial setae. The setae of the appendage are
not much longer than their respective rami. Spinules ornament the distal margins of all the leg seg¬
ments except the coxopodite and the terminal endopodite article.
The second (fig. 39, h) and the third legs hâve bimerous protopodites and obscurely trimerous
rami. There is a subquadrangular intercoxal lamella. The coxopodite has no armature ; the basi¬
podite bears a short seta on the latéral margin. The endopodites are slightly shorter than the exopo-
dites. The basal and second segments of the endopodites each bear a seta at the distal medial corner.
The terminal segment bears a latéral, 2 terminal, 3 medial setae. The basal and second segments of
the exopodite each bear a latéral spine, 1 medial seta. The terminal segment bears 2 latéral spines,
Source : MNHN, Paris
ASCIDIGOLIDAE
SUBFAMILIES
103
1 terminal spiiie, 1 terminal seta, 3 médial setae. Spinales ornament the distal margins of the basal
and second segments of the rami.
The fourth leg (fig. 39, i) lias the protopodite and exopodite the same as the second and third
legs ; the endopodite difîers by bearing on the second segment 2 (rather than 1) setae and on the ter¬
minal segment 5 (rather than 6) setae.
The fifth leg (fig. 39, b) consiste of a simple lobe at the distal margin of the flrst urosomal seg¬
ment, bearing a terminal seta. The is no indication of sixth legs on the second urosomal segment.
The caudal ramus (fig. 39, j) is articulated on the urosome so as to diverge from the other ;
the shape is rather characteristic and may reflect the immature .State. There is a stout latéral seta
at ahout the middle of the latéral inargin, in the emargination. There is a seta at the distal third of
the dorsal surface and there are 4 subequal stout terminal setae.
A single specimen was available to us for examination, and the details we made out are only
partial. Lang (1948) and Gotto (1954) hâve documented the great rarity of occurrence of the males
of this copepod. From their publications, we judge that the male described by Lang must be the
fifth copepodid, because of the correspondence to our specimen. Our impression as to the stage of
our specimen dérivés from a certain juvenility of the habitus and some of the appendages, as compared
with other ascidicoles, and the 5-segmented urosome. We concur with Gotto that the male descri¬
bed by him was probably subadult ; we would estimate it as the fourth copepodid. Some possible
discrepancics in our description and the tabulations of the other authors, as to the setation of the legs,
cannot be dealt with noM' because of the scarcity of material and the general juvénile faciès of ail the
specimens so far studied.
From these studies we feel that we can infer for this species a male which will conform very
satisfactorily with the general features of an ascidicolid male, as discussed above in our characteriza-
tion of the family.
The many correspondences of Mychophilus to Enteropsis hâve been pointed out before (i. e,.
Chatton & Brément, 1909c) but we feel that there is considérable desirability in retaining the generic
séparation at présent, particularly with the recent reënforcing contribution of the addition of a second
species in Mychophilus (Stock, 1967a).
Mychophilus fallax Stock, 1967
Mychophilus fallax Stock, 1967a, p. 9-11, fig. la-li, (type locality Dahlak Archipelago, Red Sea, from
Botryüoidss nigrum Herdman).
Distribution :
Red Sea.
Host :
BolryUoides nigrum Herdman.
Enteropsis Aurivillius, 1885
Enteropsis AurivilUus, 1885a, p. 237-239, (types species, by monotypy, Enteropsis sphinx Aurivilius,
1885) ; 1885b, p. 282. — Canu, 1886a, p. 370-372 ; 1890, p. 759 ; 1891, p. 469, 475. — Shimke-
vich’, 1889, p. 76. — Schimkewitsch, 1896, p. 345, 350, 352. — Canu, 1892, p. 25, 29, 30, 52,
58, 66, 108, 117, 132, 133, 218, 220, 221, 222 (part). — T. Scott, 1901b, p. 241-242 (part) ; 1907,
p. 369 (part). — Caïman, 1908, p. 182. — Chatton & Brément, 1909b, p. 196-198 ; 1909 d, p. 239 ;
1910, p. 91-92. — Hartmeyer, 1911, p. 1734, 1735. — Chatton & Harant, 1922b, p. 157-162 ;
1922c, p. 249, 250. — Schellenberg, 1922, p. 290-292. — Harant, 1931, p. 371. — Wilson, 1932,
p. 599, 600, 602. — Neave, 1939, p. 241. — Sewell, 1949, p. 192. — Gotto, 1960, p. 226. — Mon-
Source : MNHN, Paris
104
PAUL L. ILLG AND PATRICIA L. DUDLEY
niot, 1961, p. 98. — Gotto, 1961, p. 151. — Monniot, 1965, p. 160. — Dudley, 1966, p. 155, 157.
158, 159, 160.
Haligryps Aurivillius, 1885a, p. 242-243 (type species, first listed, Haligryps acideatus .\urivillius,
1885) ; 1885b, p. 282. — Neave, 1939, p. 552.
Synthetys Canu, 1886a, p. 371 (lapsus).
This genus is very cohesive, once Enleropsis pilosus Canu 1886, is removed ; we hâve assigned
the latter form as Ascidicolidae species incerla sedis (p. 132). The body of the female of an Enleropsis
species is characteristic in form, basically eruciform, elongate, with indistinct segmentation. Thcre
is usually a general covering over the cuticle, typically of very fine hairs. The anus is terminal. The
cephalosome is somewhat set off from the metasome, may hâve pleural folds developed and bcars,
4 pairs of appendages. The mandibles and maxillipeds are always lacking.
The metasome is the iongest and widest portion of the body and bears 4 pairs of equispaccd
uniramous legs. The sixth thoracic segment may possibly be involved in the metasome, but if so.
the participation is not expressed in the form of fifth legs or pediform projections.
The urosome may be demarcated by a groove and the anterior portion carries the génital appa-
ratus. There is a midventral insémination pore ; internai diverging canals ; and at the latéral ovi-
ducal apertures there may be sclerotizations and setules. The caudal rami may be présent or ab.sent,
but are always reduced.
The antennule, antenna, labrum, maxillule and legs conform to the subfamilial diagnosis.
The maxilla is generically diagnostic and usually not distinctive in the species. It is a bimerous
appendage, with a massive basal article, forming a complicated articulation with a medially directed
terminal article. There is a prominence on the médial distal corner of the basal article opposing the
terminal article. The latter is drawn out as a slightly falcate, sclerotized process. It bears a sela on
the distal margin near the articulation.
No species in the genus is really well known. The majority are very rare and almost ail are
described from very few specimens. Some of the descriptions are confusing and incomplète. The
original descriptions of Aurivillius and those of varions forms in his collection hâve been variously
interpreted in subséquent révisions. We propose to continue the usage of most authors (Canu, 1892,
ScHELLENBERG, 1922) and recognize the various forms described by Aurivillius from the single host
Molgula ampulloides {Enleropsis sphinx ; Haligryps teres ; Haligryps aculeatus) as forms and stages
of a single species, E. sphinx Aurivillius. We propose to recognize Scbellenberg’s form from the
Antarctic, E. sphinx var. georgianus as a nominate species E. georgianus Schellenberg (1922).
After careful study we cannot accept the arguments of Chatton & Harant, (1922b, p. 158-
162) as to the probable bispecific composition of Aurivillius’ material. We feel that in the absence
of new information from actual specimens that the best interprétation of the 1885 material is to treat
all as the same species. We acknowledge there are discrepancies among various of Aurivillius’
statements, but the correspondences, when allowances are made for the fact that the 2 sexes and indi-
viduals of various sizes and âges were treated, are truly remarkable and, in our opinion, substantiatc
identification.
The specimen from Tethyum papillosum from Port-Vendres, identified as E. sphinx by Chat-
ton & Harant (1924b, p. 158-159, figs. 1-3) and figured in part, below, we consider to represent a
different species, which has been subsequently described as Enleropsis chattoni Monniot, 1961. As a
resuit of this reconsideration we relegate Enleropsis teres (Aurivillius) sensu Chatton & Harant (1922li,
p. 162) to the synonym of Enleropsis sphinx Aurivillius.
In chronological order, the species of Enleropsis we are recognizing here are : E. sphinx Auri¬
villius, 1885 ; E. roscoffensis Chatton & Brément, 1909 ; E. georgianus Schellenberg, 1922 ; E. onyckophorus
Schellenberg, 1922 ; E. chattoni Monniot, 1961 ; and we are proposing the following new species : E. copi-
tulatus ; E. minor ; E. superbus, and E. abboUi.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAHILIES
105
ENTEROPSIS FIFTH MALE COPEPODID
So far there bas never been an adull male of a species of Enteropsis available for study. The
only information which could apply at ail to males dérivés from Aurivillius’ descriptions (1885)
of Haligryps leres and H. aculealus ({885a, p. 243-246). Canu (1891, p. 469 ; 1892, p. 218-22) proposed
lhat the Haligryps species of Aurivillius were thc adult and immature males of Enteropsis sphinx
Aurivillius. This scheme bas received subséquent concurrence (Schellenbebg, 1922) ; and Chat-
ton & Habant (1922b, p. 161), correctly, in our opinion, revised the attribution of II. aculeatus to
the aiilepenultimate stage (fourth copepodid) (note below that we do not agréé with the complété
synonymy of Chatton & Habant in connection with the Haligryps forms). We would make a fur-
ther révision of considering H. leres the male fifth copepodid. Even though the séminal vesicles are
illustrated by Aurivillius, they are not necessarily at the definitive stage of spermatophore formation.
Further, such development of the séminal vesicles, well up in the thoracic région, as depicted by Auri¬
villius, is diagnostic of male late fifth copepodids in the Nolodelphyidae. The urosome is 5-segmented,
a consistent feature of fifth copepodids.
There is a considérable correspondence of the Enteropsis male fifth copepodid, as can be deduccd
from Aurivillius’ présentation, and the male copepodid of Mychophilus, as described and discussed
above (p. 102). We présent our inferences below, but must emphasize that the characters of the defi¬
nitive male could very well be further modified.
The body form is essentially cyclopiform, with the cephalosome including the appendages
though the maxilla, and probably also the segment corresponding to the maxilliped, although this
appendage is absent. The metasome bas 4 free leg-bearing segments. The urosome, including the
segment of the fifth legs, is 5-segraented.
The antennule is 6 or 7-segmented, and présents a rather general ascidicolid habitus.
The antenna is much like that of the female Enteropsis sphinx Aurivillius. In both cases
this author described and depicted 4 segments in the appendage, but we feel that he may have misin-
terpreted some surface details and the appendage is perhaps 2-segmented, as it is throughout most
of the species of the genus. The terminal armature consista of 2 unequal setae.
The labrum is a distinct lobe, and the author did not describe ornamenting setae.
The maxillule is much like that of the female, bilobed. One lobe bears 3 setae ; the other bears
2 setae.
The maxilla corresponds well to lhat of the female. It consista of a massive basal lobe, with
a terminal hooked precess. There seems to be a pad on the basal part, opposable to the distal hook.
The correspondence here of male with female appears to be much greater than is the case in Mycho-
pkilus.
The legs are ail Liramous and some of the details of armature can be inferred. In the first and
fourth legs the coxopodites have no armature, and the basipodite bears a latéral seta. In the first
legs the endopodites are trimerous, the basal segment bears no armature, the second segment bears
a médial seta, the third segment has 1 medial, 2 terminal, 3 latéral setae. The exopodite is bimerous ;
the basal segment bears 1 latéral spine ; the terminal segment bears 1 latéral spine, 1 terminal sea,
2 medial setae.
The endopodite of the fourth leg is trimerous. The first segment bears 1 medial seta, the second
segment bears 2 medial setae ; the terminal segment bears 1 medial, 2 terminal, 2 latéral setae. The
exopodite is trimerous. The basal and second segments each bear 1 latéral spine and 1 medial seta.
The terminal segment bears 2 latéral spines, 1 terminal spine, 1 terminal seta, 3 medial setae.
A single seta is depicted as representing the fifth leg, and there is no indication of sixth legs.
The caudal rami diverge ; each bears 1 latéral seta, 1 dorsal setae, 4 terminal setae.
We are struck with the correspondence of the characters of this male copepodid with those of
the female Enteropsis sphinx, particularly in the mouthparts. Even in the fifth stage, the male of
Source ; A1JJHW, Paris
106
PAin- L. ILLG AND PATRICIA L. DUDLEY
Mychophilus seems to show a somewhat greater dimorpliism in the mouthparts, although the male
retains the Basic pattern of the female appendages. In features of the antennules, the swiinming legs,
the fifth legs, the caudal rami and the general habitus, the male fifth copepodids of Enleropsis, and
Mychophilus show striking similarity. Probably in this subfamily it could be said the males will be
readily diagnosable by the characters of the females, but the mouthparts will hâve longer setae. This
would secm to indicate a different trend of dimorphism from that found in the genus Bolryllophilus
and in the haplostomins, where the mouthparts of the male tend to reduce greatly beyond the condition
found in the female. There is also some contrast with the enterocolins where the trend is also toward
réduction of niasticatory cléments, with other modifications, such as élongation of more distal setae.
KEY TO SPECIES OF ENTEROPSIS, BASED ON FEMALES
1 .
1 '.
2 .
2 '.
3.
3'.
4.
4'.
5.
5’.
6 .
6 '.
7.
7'.
8 .
8 '.
Lahruni without setiform processes ; antenna with 2 apical processes. 2
Labnim with setiform processcs ; antenna with 1 or 2 apical processes. 3
Palp portion of maxillule with 2 setiform processes. chatloni Monniot, p. 106
Palp portion of maxillule with 3 more or less setiform processes.. superbus, new species, p. 108
Antenna with 1 apical process. 4
Antenna w-ith 2 apical processes. 7
Promiiient seta-like structure at each latéral corner of labrum. 5
Row of seta-like structures on margin of labrum. 6
Maxillule complex basally, prominent médial extension terminating in 2 subequal setiform acute
projections . minor, new species, p. 111
Maxillule complex basally, prominent médial extension bifid terminally, one element acute, seti¬
form, 1 forming a narrow lobe with spatulate apex. abhotii, new species, p. 128
Antennule unimerous, short; antenna bimerous.... roscoffensis Chatton & Brément, p. 114
Antennule longer, indistinctly bimerous; antenna unimerous.
onychopkorus Schellenberg, p. 131
With 8 or 9 setiform processes on labrum. sphinx Aurivillius, p. 120
With 5 or 6 minute setiform processes on labrum. 8
One apical process of antenna twice the length of second ; caudal ramus very short, terminating
in tiny point, placed far laterally to anus. capitulatus, new species, p. 123
Apical processes of antenna subequal ; caudal ramus subterminal, terminating in setiform process.
georgianus Schellenberg. p. 126
Entebopsis chattoni Monniot, 1961
(Figure 40)
Enleropsis sphinx (non Aurivillius) Chatton & Harant, 1922b, p. 156-162, Gg. 1 (from Port-Vendres,
Golfe du Lion, France, in Halacynthia papillosa [L.]). — Gotto, 1960, p. 225 ; 1961, p. 151-152.
Enleropsis sp. Monniot, 1961a, p. 98.
Enleropsis chattoni Monniot, 1961b, p. 113-116, fig. 1 (type locality, BanyuIs-sur-Mer, Golfe du Lion,
France, in Microcosmus vulgaris Heller) ; 1965, p. 160.
Source : Mt-JHN, Paris
ASCIDICOLI0AE AND SUBFAMILIES
107
Fio 40 — Enteropsis ekatlom Monniot, female : a, habitus, ventral ; b, habitus, latéral ; c. antennule and anterior
end of cephalosome ; d, antennule ; e, antenna ; f, labr.im ; g. maxillule ; h, maxillule ; i, maxilla ; j, second leg ;
k, caudal ramus. Scales = 0.1 mm.
Distribution :
French and Italien Mediterranean coasts ; British Isles.
Hosts :
Mtcrocosmus viiigaris Heller, Hahcynihia papillota (L.), Diamna violacea Savigny.
Source ; MNHN, Paris
108
PAUL L. ILLG AND PATRICIA L. DUDLEY
Specimens examined ;
From Microcosmus vulgaris Heller = . , ,
Poîzuoli, Bay of Naples, Italy, 40m, January 16, 1958, 1 female.
We ofîer what we consider to be pertinent supplementary information to the descriptions of Mon
moT and of Chatton & Harant, and Bgure our specimen to compare it with other specimens and
other species in the gcnus. The specimen, an adult female, measured 3.48 mm, overaU length (d
Monniot : “ environs 4 mm ”). The proportions of our specimen, cephalosome : metasome : urosoms’
are about 1.5 : 5 : 3.
The habitus {figs. 40, a, b) agréés well with that figured by Monniot. Chatton & Harant
did not présent a figure of the habitus. We figure the oviducal aperture and the insémination pore
(figs. 40, a, b), not previously illustrated. Monniot refers to a movable rostrum, but we hâve notso
far foun’d any such in ail the species of Erüeropsis we hâve studied ; there is a pronounced ventral
bulge of the anterior portion of the cephalosome, and it may carry some ornamentation. The anten-
nule (figs. 40, c, d) agréés essentially with the earlier description in ornamentation and setation,but
the terminal article is shorter than that illustrated by Monniot and by Chatton & Harant. The,
antenna (fig. 40, e) appears as it was illustrated and described by Monniot. The figure and illustra¬
tion of Chatton & Harant would seem to départ strongly, because they consider as a terminal seg¬
ment what we are convinced is the swollen basal portion of the larger distal setiform process of the
appendage. These processes, doubtless homologous with setae, do not actually forra the strong arti¬
culations which previous illustrations would tend to indicate. The labrum (fig. 40, f) is not prorainenl
and there are no labral setiform ornamentations ; this concurs with the findings of the previous authors.
The maxillule, termed mandible by the French authors, agréés in essential characters (fig. 40, g, h),
but one of the terminal setae of the médial extension of the basal lobe is somewhat longer in our mate-
rial than was illustrated by Monniot. There are no actual articulations, on the appendage although
the représentations of strong outlines in the previous publications would seem to indicate joints. The
maxilla (fig. 40, i) agréés with the published accounts and figures. The four thoracic legs (fig, 40, j]
are ail essentially similar in structure. We feel, on the basis of extended studies on several species
in the genus, that there is a misinterpretation of the structure of these legs in the original descriptioe
of E. chattoni. They are stated to bear “ deux stylets égaux, parallèle, l’un dorsal, l’autre ventral
Referring to this character, Monniot pointed out a différence from the description of Chatton & Harant
He accordingly accepted their désignation of their material as E. sphinx. We hâve discussed our
findings on the legs 1 to 4 in Enteropsis and our conviction is that there is a remarkable uniformily
throughout the genus. We would suggest that the appearance of 2 stylets could readily be derived
from some views of the terminal articulated spine in its sheath. Our specimen from Naples, confoN
ming throughout otherwise to the description of E. ckaltoni, has the structure of the legs as we hâve
found them in other species. We illustrate the caudal ramus (fig. 40, k).
Our record extends the known distribution to the Bay of Naples. Gotto provides (1961, p. 152)
interesting data on the form he observed from Diazona violacea. AH the young and non-ovigerous
stages were found in the stomach of the host. Adult females and egg-strings were found in the pha¬
rynx. Gotto suggests that the female migrâtes through the long, slender esophagus just priortothe
extrusion of the egg-strings and he offers a brief discussion of possible factors involved in the adaptive
significance of this behavor. The occurrence in the branchial basket is a relatively rare feature for
a species of Enteropsis. Most of the living ovigerous females we observed were obtained from the
atrial cavity of the host.
Types :
Enteropsis superbus new species
(Figure 41)
1 t female, San Juan Islands, Washington, August 1, 1966, in Pyura kaustor (Stimpeon).
1 female, 7.7 mm. Paratypes listed below. ^ ’ 8 . > »
Source ; Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
Fio. 41. _ Entenpais auperbus, new species, paratypic female : a, habitus, latéral ; b, habitus, ventral ; c, en face view
of eephalosome : Al = antennule, A2 = antenna, L = labrum, Mxl = maxillule, Mx2 = maxilla ; d, antennuk,
ventral ; e, antenna, ventral ; f, labrum ; g, maxillule, ventral ; h, maxilla, ventral ; i, first leg, anterior ; j, génital
aperture ; k, caudal ramus. Scales for a, b, c = 1.0 mm ; other scales = 0.1 mm.
Source : MhlHN, Paris
110
PAUL L. ILLG AND PATRICIA L. DUDLEY
Spccimens examined :
From Pyura hatalor (Stimpson) :
Upright Channel, San Juan Islands, Washington, 1 female, 5.3 mm.
Lopez Pass, San Juan Island, Washington, 15 July, 1950, 1 immature female, 7.2 mm.
Parker Reef, Orcas Island, Washington, 8 July, 1950, 1 immature female, 1.54 mm.
“ Potato Patch ”, San Juan Islands, Washington, July, 1952, 1 female, 8.2 mm.
West Sound, Orcas Island, Washington, 3 fourth copepodids, 3 fifth copepodids, 2 adult females,
1 5.3 mm long — from many hosts examined.
San Juan Islands, Washington, August, 1966, 1 female, 7.5 mm.
Description :
Female :
The overall hody length from the anterior end of the cephalosome to the posterior end of the
caudal rami ranges from 1.54 mm to 8.2 mm in availablc specimens, with an average of 6.9 mm in mature
adults.
The hody (figs. 41, a, b) is almost uniformly cylindrical, rounded at the extremes and with the
well spaced legs markedly salient. The cephalosome is not set olT notably from the body. The meta-
some is only slightly indented as an indication of the component segments. The urosome is not dis-
tinctively articulated or otherwise set ofî from the metasome and exhibits only a slight taper. The
minute caudal rami are terminal on the rather truncate posterior surface of the urosome, set somewhat
ventrally. In the overall impression of the body the urosome is prominent. The proportions of the
lengths of the three major body régions in the specimens seen so far are about 1 : 5 : 2.5, cephalosome :
metasome : urosome. The thickness (that is, the depth at midpoint in latéral view) of the body is
about 1/7 the overall length. The body is generally covered with very fine hoirs. The illustrations
wil depict these variously or omit them, depending on the degree of magnification applied.
The head appendages are described from an en face préparation, so the terms of orientation
refer to this {fig. 41, c), not necessarily to the basic anatomical relations on the body.
The antennule (fig. 41, d) is obscurely bimerous but distinctly articulated on the head. Therc
is a strong articulative line on the ventral surface but no interruptions of the dorsal cuticle. Thcre
are no ornamenting spinules or hoirs. The basal article has 2 setules on the anterior margin at about
its middle. The terminal article is tapering and truncate. Therc are a stout seta and 2 setules apically,
a stout seta on the anterior margin at the proximal third. There are 2 setules near the base, 1 each
on the dorsal and ventral surface, and an additional setule on the ventral surface at the base of the
second anterior seta.
The antenna (fig. 41, e) is clearly bimerous, heavily sclerotized uniformly overall, distinctly
flattened, and the outline is tapering. The basal article has 3 rounded lobes along the posterior margin
and a similar lobe on the distal margin on the ventral surface. These lobes are set with patches of
elongate heavy spinules and there are additional curved rows of shorter spinules proximally and distally
on the ventral surface. The dorsal surface is unornamented. The distal article has one terminal,
articulated seta and a subapical articulated, subequal spine on a prominence at the distal fourth.
There are rows of spinules on the ventral face, 1 row' at the base of the distal spine, 1 row across the
face of the segment just proximal to the subapical spine base and a short row at the proximal fourth.
The dorsal surface has patches of finer spinules.
The labrum (fig. 41, f) is a rather simple lobe with sclerotized distal margin and lacking margi¬
nal “ setae ”. The ventral surface is finely spinulose.
The maxillule (fig. 41, g) is bilobed with the lobes well articulated on the unsegmented base.
Thé .appendage is only weakly sclerotized but the surfaces are roughened by a general covering of small
protubérances. The basal lobe has no armature. The latéral lobe is flattened and subrectangular
in outline, the distal margin formed by 3 unarticulated flattened processes, triangular in outlfne and
Source: Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
111
équivalent to the setae of the palp and base of other species. Two of tlie processes are apical, the other
is just subapical and the 3 are subequal. A curving row of spinules crosses the ventral surface at about
the middle and a patch of hairs lies proximal to these. Some rows of finer spinules lie at the basal
région of the subapical process. The articulated spine is accompanied by a subterminal spiniforin
setule, about two-thirds as long as the spine. Three rows of spinules curve across the ventral surface
beyond the middle.
The maxilla (fig. 41, h) is distinctly bimerous and distinctly articulated on the body. Heavy
sclerotizations form a distinctive pattern. The surface of the sclerotized areas are uniformly pitted
and each pit surrounds a small hair. The unsclerotized surfaces are roughened by a general covering
of small protubérances. The appendage is probably représentative of the basic structure ihroughoul
the genus, and distinctive proportions would only be significant as such information becanie availabie
from the previously described species. The basal article has no armature. The articulation of the 2
component articles is most complicated but the terminal article can be doubtless move only in a res-
tricted fashion. It is directed medially as a resuit of the arrangement. The distal article is tapering
in outline, and curves slightly at the tip. The heavy sclerotization contributes to its development as
a stout, somewhat flattened claw. Set in a subcircular unsclerotized space on the ventral face, there
is a modilied seta, encircled at the base by a sclerotized ring. The seta, about twice as long as thick,
is peg-like with a smoothly curved apex.
The 4 pairs of legs are much alike (first leg, fig. 41, i) ; each leg is bimerous and with the apex
directed somewhat medially. The articulation of the leg with the body is obscure because the cuticle
is essentially uninterrupted. The leg has generally distributed fine hairs over the whole surface.
The proximal article has heavy sclerotizations, probably in a distinctive pattern. On the anterior
surface there are about 6 short setules, emerging from pits in the sclerotization. The distal article
is sclerotized overall and the apex is complicated in structure. There is an apical médial tuberculated
lobe beside a socket which reçoives the opposable articulated terminal spine, which is heavily sclero¬
tized. The terminal spine is probably rétractile into the terminal article.
There is no fifth leg.
A complicated structure, perhaps involving the sixth leg (fig. 41, j) is proximal to the posterior
margin of the sixth thoracic segment near the middle of the latéral surface. This structure is a bilobcd
sclerotized ridge, with a minute setule on each lobe. Underlying this structure there is an ovoid lobe
of cuticle with surface spinulations. Under this is a slit, the oviducal opening, surrounded by a pecii-
liarly striated cuticle.
The caudal rami (fig. 41, k) are minute and very inconspicuous on the rounded posterior end
of the body. Each is a conical process, unarticulated at the junction with the body, but with a clearly
articulated tip.
This rare copepod occurred in the atrium or distal intestine of the host. The animal is very
handsomely colored. There is an overall bright yellowish-orange coloration, with a conspicuous
bright, light-red eye. The ova in the oviducts, which form conspicuous, paired, extensive tubes along
the sides of the body, are scarlet-purple. The movements are as described for E. roscoffensis below,
with the addition that antennae and maxillae show strong striking movements when the animal is
turned on its back.
Enteropsis minor new species
(Figure 42)
Types :
Holotypic female : Peavine Pass between Blakely Island and Obstruction Island, Washington,
July 13, 1956, 1 female, 1.6 mm, from Metandrocarpa taylori Huntsman ; paratypes, listed below.
Specimens examined :
From Metandrocarpa taylori Huntsman ; Peavine Pass between Blakely Island and Obstruction Island,
Washington, August 15, 1966, 3 females, 1.4 mm, 1.6 mm, and 1.6 mm.
Source : MNHN, Paris
112
PAUL L. ILLG AND PATRICIA L. DUDLEY
Fig. 42. — Enteropsis minor, new species, paratypic female ; a, habitus, latéral ; b, habitus, ventral i c, en (ace view
of cephalosome : Al = antennule, A2 = antenna, L = iabrum, Mxl = maxillule, Mx2 = maxilla ; d, antennule ;
e, antenna ; 1, Iabrum ; g, maxillule, attached to head scierotization ; h, maxilla ; i, first leg ; j, oviducal aperlurè
and omament. Scales for a and b = 0.2 mm ; other scales k 0.1 mm.
Description :
Female :
The overall body length from the anterior end of the cephalosome to the posterior end of the
urosome ranges from 1.4 to 1.7 mm in the available specimens of mature females.
The body (figs. 42, a, b) has only a slight anterior taper, is more pronouncedly tapering posteriorly.
Source : MIJHN, Paris
ASCIDICOLIDAE
SUBFAMILIES
113
The cephalosome is not set off markedly from the metasome. The metasome consists of 4 leg-bearing
segments, these clearly delimited by cuticular Unes dorsally and ventrally. Laterally, the segments
are not defined except by indentations. The slightly tapered urosome, with 2 ingrédient segments
marked by indentations only, is rounded posteriorly and no caudal rami are seen. The proportions
of lengths of the 3 major body régions in the specimens seen are about 1 : 4.5 : 2, cephalosome : meta¬
some : urosome. The depth of the body at midpoint in latéral view is about 1/5 the overall length.
No ornamentation could be discerned on the general body surface. This species differs greatly from
other local species by its small size.
The head appendages are described below from the orientation of an en face view {fig. 42, c)
so the terms of orientation refer to this and not neccssarily to the basic anatomical relations on the
body.
The antennule (fig. 42, d) is unimerous and articulated very indistinctly on the head. There is
a small sclerotized patch on the dorsal surface and also a row of spinules just distal to this. The ven¬
tral surface has no ornamentation. The éléments of armature are inserted apically and subapically
on the anterior margin. There are 3 stout setae, 1 filiform seta, and 1 setule.
The antenna (fig. 42, e) is incompletely bimerous, because on the ventral surface the articulation
reaches only part way across the appcndage. The terminal spine is so stout and the jointing of the
articles is so strongly indicated that casual observation migbt well lead to interprétation of this as
a trimerous appendage. The basal “ article ” bears no setae or spines but is ornamented with rows
of spinules near the distal margin on the ventral face. The second “ article ” tapers and the apex
is occupied by the articulation of the very strong terminal spine which is longer than the article. The
whole ventral surface of the article and the entire basal third and both margins of the spine are orna¬
mented with spinules.
The labrum (fig. 42, f) is a simple lobe with strongly convex sclerotized posterior margin. Direc-
ted posteriorly from the latéral bases of the lobe are 2 strong, plumose setae. The ventral surface
of the labrum is covered with fine hairs.
The maxillule (fig. 42, g) appears to be uniramous and the distal lobed portion has no articula¬
tion with the base. The entire appendage is relatively heavily sclerotized and there are ventral scie-
Fotized areas on the basal portion. The sclerotizations bear pits through which fine hairs emerge.
At the base of the appendage, on the head surface, there is a sclerotization of distinctive outline. The
basal portion is expanded as 2 heavily sclerotized lobes thickly covered with rows of spinules. Our
sériés of local species, in the genus Enteropsis offers sufFicient graduation of structure in maxiUules that
we are convinced that these lobes represent a vestige of the palp, indeed of its apical setae. The medial
process of the basal portion is long, tapered, and the distal margin is formed, without articulation, by
2 stout, ciliated setae, one of which is about three-fourths as long as the other.
The maxilla (fig. 42, h) is bimerous and distinctly articulated on the body. Heavy scleroti¬
zations form a distinctive pattern on both articles. The surface of the sclerotized areas is pitted with
a fine hair emerging from each pit. In addition, there are a few heavy spinules on the ventral sclero¬
tized surfaces of the appendage. In general, the appendage conforms to generic characteristics. The
distal article is complexly articulated and tapers in outline. It curves slightly at the tip to form a
long, stout, flattened claw. Set in a subcircular unsclerotized space on the ventral side there is a
relatively long, simple seta.
The 4 pairs of legs (first leg, fig. 42, i) are much alike, ail bimerous and uniramous, but obscu-
rely articulated with the body. The proximal article has heavy sclerotizations distinctly arranged.
On the anterior surface of the sclerotization are 2 pits, from each of which a spinule emerges. Just
proximal to the more medial pit there are 2 rows of spinules, Two rows of spinules also ornament the
unsclerotized portion of the anterior surface. Curved rows of fine, long spinules ornament the distal
margin. The distal article is sclerotized overall and has a complicated apex. There is a medial lobe,
composed of a medial, heavy, sclerotized rod in the cuticle supporting anterior and posterior unsclero¬
tized cuticular extensions with lobùlated latéral margins. A triangular sclerotized base piece subtends
th»,structure. Apposing the lobe is an articulated spine. There is a sclerotization around the poste¬
rior surface of the spine which may provide a hooded covering.
8
Source ; MNHN, Paris
114
PAXJl- L. ILLG AND PATRICIA L. DUDLEY
At the postérolatéral boundaries of the first urosomal segment are sclerotized plates and flaps
associated with the oviducal aperture (fig. 42, j). Part of the complex may represent a sixth leg.
From an insémination pore found mid-ventrally at the posterior margin of the first urosomal segment,
elongate, looping séminal tubes diverge toward the oviducal apertures.
There are no caudal rami. Broken rows of spinules ornament the ventral surface of the uro-
some.
The coloration of the female is an overall dull orange, with a large conspicuous red eye. The
ova in the oviducts are purplish red, embryos in egg sacs are cerise red. In several hundred spéci¬
mens of Metandrocarpa, only 3 females were found. In 4 other specimens of Metandrocarpa, free
in the atrium, egg sacs containing embryos of Enteropsis minor were found, but there were no females.
In 2 of these cases, 2 egg sacs were found, these containing embryos of different âges. Thèse obser¬
vations lead to the spéculation that females may move from individual to individus! of the social
tunicate. This species appears to be much more active than any of the other local species observed.
When the female is placed ventral surface down on a substratum, she can progress slowly in a coordi-
nated manner by moving the legs metachronically. When the female is placed on her back, she is
unable to turn over, but when she is in this position, the antennae are moved with violent striking
movements, the maxillae strike medially and the legs move like those of Enteropsis roscoffensis, with
first a fuU extension and then a posteriorly directed flexion during which the apical spine is retracted
and then the distal article télescopés into the basal. Flexion of the legeonsists of a distinctively ciawind
action. The body can generally elongate and contract, or curve in a bow-like fashion, and the uro-
some can swing actively. Because the host is so minute and extraction of the large copepod is extre-
mely difficult we hâve been unable to déterminé the actual site of occurrence within the body of the
ascidian.
Enteropsis roscoffensis Chatton & Brément
(Figures 43, 44, 45, 46)
EnUropsis roscoffensis Chatton & Brément, 1909b, p. 198-200, fig. 1-5 (type locality, near Roscofî,
France, from Styelopsia grossularia van Beneden). — Schellenberg, 1922, p. 291. — Chatton
& Harant, 1922b, p. 159-160. -- Harant, 1931, p. 371. — Sewell, 1949, p. 189. — Gotto, 1960,
p. 226.
Distribution :
Atlantic and Mediterranean coasts of France.
Hosts :
Dendrodoa grossularia (van Beneden), Pyura microcosmus (Savigny), Tethyum saoignyi.
Specimens examined :
From D. grossularia (van Beneden) :
Plymouth, England, exact locality of collection unrecorded, from laboratory specimens, Sep-
tember 5, 1958, 1 female.
Roscoff, France, August, 1909, Chatton number 27, type lot, 5 females.
Coast of France, June, 1911, Chatton number 51, 3 females.
From Phaüusia mammillata (Cuvier) :
No collection record, Chatton number 249, 1 female.
From Styela gibbsii (Stimpson) :
Off Upright Head, Lopez Island, Washington, 1 ovigerous female, approximately 3 mm .
Friday Harbor Labs, Washington, August 21, 1953, 8 females ; 4 specimens measured were
5.1 mm, 3.9 mm, 3.5 mm, 3.8 mm.
Source : MIJHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
115
If,Q 43 _ EnleropaU roscoffensis Chatton & Brément, female frora Dendrodoa : a, habitus, latéral ; b, antennule ; c,
antenna ; d, labrum ; e. maxiilule ; f, maxilla ; g, first leg ; h, caudal ramus. Scale for a = 0.2 mm ; other scalea =
0.1 mm.
Source ; AlfJHM, Paris
116
PAUL L. ILLG AND PATRICIA L. DUDLEY
nule : c. antenna 1™“ Dtj^rodoa. Plymouth ; a, habitus lat
Scalea
Source : MNHN, Paris
Fig 45 — EnteropsU roscoffentis Chatton & Brément, female from Pkallusia : a, habitus ; b, antennule and anterior
end of cephalosome ; c, antenna ; d, iabrum ; e, maxillule ; f, maxillule ; g, maxilla j h, second leg ; i, caudal ramus.
Scale for a = t.O mm ; other scalea = 0.1 mm.
Source : Mh}HN, Paris
Fiq. 46. — EnteropsU roscoffensis Cbatton & Brément, temale from Styela gibbaii : a, habitus, latéral ; b, habitus, ven¬
tral : c, habitus, ventral, showing egg sacs ; d, detail of general body ornamentation ; e, en face view of cepbalo-
some : Al = antennule, A2 = antenne, L = labrum, Mxl = maxillule ; Mx2 = maxilla ; f, antennule, ventral ;
g, antenna, dorsal ; h, antenna, ventral ; i, labrum ; j, maxillule, ventral ; k, apex of maxillule, ventral ; 1, maxilla’
ventral ; m, maxilla, ventral ; n, first l^, anterior ; o, oviducal aperture ; p, caudal ramus. Scales for a, b, c =
1.0 mm ; other scales = 0.1 mm.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBPAMILIES 119
Of! Lopez Island, Washington, June 26, 1954, 5 females, 5.0 mm, 4.3 mm, 4.1 mm, 3.7 mm,
3.3 mm.
Upright Head, Lopez Island, Washington, August 23, 1954, 1 ovigerous female, 3.1 ram.
Üpright Head, Lopez Island, Washington, August 6, 1954, 1 female, 4.0 mm.
Friday Harbor Laboratory, Washington, August, 1966, 3 females, only 2 measured : 3.6 mm,
and 2.8 mm.
Reef Island, Washington, 10-17 fathoms, August 26, 1966, 2 females : 3.1 mm and 3.7 mm.
We ofîer what we consider to be pertinent supplementary information to the original descrip¬
tion and figure one specimen from the type host, Dendrodoa grossularia, from the type collection, one
collected by us from D. grossularia, from Plymouth, England, one specimen from Pkaüusia mamm.il-
laie, from the Chatton collection, but not identified by him and a représentative specimen from a
considérable sériés we hâve accumulated from Styela gibbsii from Washington. As in many species
of EnUropsis the cuticle over the body has a general covering of fine hairs. This has not been depicted
in some of our figres.
Specimens from D. grossularia (figs. 43, 44) : overall measurements, 3.5 mm and 3.77 ram, lying
about in the middle of the range of size for the species as reported by Chatton & Bbément. The
proportions of our specimens, cephalosome : metasome : urosome, were 1 : 9 : 3 or 4. The corresponding
formula given by Chatton & Brément (p. 198) read 1 : 14 : 13. which we must consider a misprint.
Examination of their figures suggests the actual reading should hâve been 1 : 14 : 5.
The habitus (figs. 43, a ; 44, a) is in agreement with the earlier figure and description, except
the formula for the proportions. The oviducal aperture on the urosome we saw just as was represented
in the original figure and we présent an enlargment (fig. 44, h). The antennules (figs. 43, b ; 44, c)
agréé essentially with the earlier figure and description ; we saw a few more setae. The antenna (figs. 43,
c ; 44, d) appears as it was figured in the original description ; we interpret the appendage as obscurely
trimerous, since the terminal portion has a strong articulative line on one face. The basal articulation
is strong and complété. We agréé with the authors’ interprétation of the labrum and présent figures
(figs. 43, d ; 44, e). The bilobed maxillule, termed mandible by Chatton & Brément, (figs. 43, e;
44, f) is just as figured earlier. In furnishing supplementary information on the species, Chatton
& Harant (1922b, p. 159) evidently confused the positions of the parts, since they stated that the
spinulose seta of the basal portion inserts at the base of the endopodite. This seta is actually at the
base of the latéral lobe, this possibily representing an entîre palp. The maxilla agréés with the authors’
statements and we présent figures (figs. 43, f ; 44, g). In the legs we see the details somewhat diffe-
rently from the authors’ description and figure of the first leg. We agréé that ail the legs are alike
and from our specimens figure the first legs (figs. 43, g ; 44, h). As they State, the leg is uniramous
and the base is massive. They describe the terminal article as bearing a triangular, lamellose crotchet,
not protruding prominently. As in ail other species we find a hooded triangular articulated spine,
opposing an unarticulated, tuberculated lobe. The caudal ramus of the specimen from France is as
described originally (fig. 43, h), but in the specimen from Plymouth (fig. 44, i) we find 2 small setae
at the apex rather than 1. We consider this condition definitely anomalous and not significant in
the taxonomy. The record from Plymouth adds the species to the British faunal list.
Specimen from Phallusia mammillata (fig. 45) : In the Chatton collection there was found
a vial labelled simply “ Enleropsis des Phallusia mammillata ”, with no other data available. Aside
from minor différences, we can identify the specimen as E. roscoffensis, and herewith offer figures.
The overall length measures 4.5 mm. The proportions, cephalosome : metasome : urosome, are 1 :
10 : 3. The habitus (fig. 45, a) corresponds very well to the original. The antennule (figs. 45, e, f),
maxilla (fig. 45, g), second leg (fig. 45, h) correspond. The caudal ramus (fig. 45, i) is somewhat smaller
than in the original form and bears a tiny apical point rather than an elongate setiform process.
Specimen from Styela gibbsii (fig. 46) : The overall body length is 4.82 mm ; additional speci¬
mens measured ranged in length from 2.8 to 5.1 mm, with an average of 3.7 mm. The body (figs. 46,
a, b, c) is somewhat plumper than in the European material, with the proportion, cephalosome : meta¬
some : urosome, about 1:6:2. As in ail the other specimens the cuticle of the general body surface
Source : AItJHW, Paris
120
PAITL L. ILLG AND PATRICIA L. DUDLEY
is covered with moderately fine hairs interspersed with spinules (fig. 46, d). The head appendages
are described from the orientation of an en face préparation (fig. 46, e). The antennule corresponds
well to the original description (fig. 46, f). The segmentation of the antenna (figs. 46, g, h) is appa-
renlly suppressed, with a single strong articulation évident, and only on the dorsal surface, but with
indications of trimery. The outline of the appendage is just like that in the other material. The
labrum (fig. 46, i) bears only 5 setiform ornamenting éléments, rather than 6 as in the original descrip¬
tion and in ail the European specimens we hâve studied. The maxillule (figs. 46, j, k) is bilobed, but
with neither lobe set off by a clear articulation. The basal portion extends without interruption into
the lobes but at the basal divergence of these, on one side there is a small prominence set with nume-
rous spinules, which forms the base for an articulated spinulose seta. ' Just proximal to this on the
ventral surface is a second spinulose, prominence. At this level there is an internai sclerotization
fumishing the insertion for a major muscle. The armature of the lobes is as described previously.
The maxilla (figs. 46, 1, m) and the leg (fig. 46, n), oviducal aperture (fig. 56, o), and the caudal ramus
(fig. 46, p) agréé with the original description.
The coloration of this female is an overall soft light orange, with the red eye conspicuous. The
ova in the oviducts are purplish-light red. The egg string is lively bright red, the transparency increas-
ing as development progresses. Observations on living specimens give some information about move-
ments of the body and some of the appendages. The body can elongate and contract, and it flcxes
in a curved, bow-like fashion. The major flexure is usually between the second and third legs. Secon-
dary flexures are at the junctures of cephalosome and metasome, metasome and urosome, and also
between the other pairs of legs, The urosome beyond its juncture is immobile and the caudal rami
are inert. The legs move powerfully, mainly in the parasagittal planes. At full extension the leg
is directed slightly laterally and the apical spine extends almost fully free of the tip of the leg. The
first legs point anteriorly at full extension, the other legs reach about a right angle with the body axis.
In flexion the leg moves posteriorly and medially. First the spine is retracted, then the distal article
télescopes into the basai. The latter does not intrude into the body. The resuit of these components
of flexion is a distinctly clawing action of the leg.
None of the specimens was collected from the branchial baskets of the hosts. The usual site
of détection was the atrium, but at least some specimens were taken from the digestive tract proper.
Freshly collected examples may be difficult to observe because they are coated with sticky detritus-
like' material, probably the feces of the host.
We adopt an illustrious precedent, set by CsATTON and his colleagues, in assigning a perhaps
Bomewhat improbable identification to our specimens from Washington. The general anatomical
conformity between our specimens from the Pacific and the Atlantic is so thorough that we feel the best
solution is to conform to a purely morphological diagnosis. ,
Entbropsis sphinx Aurivillius, 1887
(Figure 47)
Enleropsis sphinx Aurivillius, 1887, p. 238-242, pl. 8, fig. 12-28 (type localities. Stations 28, 66, 76,
Vega Expédition : Station 28, W. of Taimurlandet, 76o99' N latitude, 92o20' E longitude !
Sta 76 — no data, from Molgula ampulloides van Beneden). — Shimkevich’, 1889, p. 75, 78, 79'
— Canu, 1890, p. 759 ; 1891a, p. 469 ; 1892, p. 30, 53, 54, 220. — Chatton & Brément, 1909b'
p. 200, 201. — Hartmeyer, 1911, p. 1734. — Schellenberg, 1922, p. 290. — Harant, 1931, p. 37l’
non EnUropsis sphinx, Chatton & Harant, 1922b, p. 158-159,160 (Port-Vendres, Golfe du Lion, France,
from Tethyum [= Cynthia] papillosum) (synonym of Enleropsis chaüoni Monniot, 1961).
Haligryps aculealus Aurivillius, 1887, p. 244-246, pl. 9, fig. 11-20 (type localities. Stations 28, 29, Vega
Expédition : W. Taimurlandet, 76o88' N latitude, 92o20' E longitude : Station 28 ; Taimurlandet
76®18' N latitude, 95“30' E longitude : Station 29, from Molgula ampulloides).
Source : MNHN, Pans
Fie. 47. _ EnUroptU tphinx Aurivillius, female : a, babitus, latéral ; b, habitua, ventral ; c, uroaome ; d, cephaloaome,
ventral ; Al = antennule, A2 = antenna, L = labrum, Mxl = maxiliule, Mx2 = maxüla ; e, antennule ; f, antennule ;
g, antenna ; b, apex of antenna ; i, labrum ; J, maxiliule ; k, maxiliule ; 1, maxilla ; m, maxilla ; n, firat leg, ante-
rior surface; o, oviducal aperture ; p, caudal ramus. Scale for a, b ^ 1.0 mm ; other scales — 0.1 mm.
Source : MNHN, Paris
122 PAUL L. a.LG AND PATRICIA L. DUDLEY
Haligryps teres Aurivillius 1887, p. 243 244, pl. 9, fig. 1-10 (type localities Stations 28, 76, Vega expé¬
dition, from Molgula atnpulîoides).
Synlhetys sphinx, Canu, 1886a, p. 371 (lapsus).
Enteropsis teres Chatton & Harant, 1922b, p. 161-162 (for specimens of Aurivillius’ collections includ-
ing the adult and ovigerous females and the male adults and antepenultimates : the Haligryps
teres and aculeatus specimens). — Harant, 1931, p. 371.
Distribution :
Siberian Arctic coast.
Hosts :
.Molgula manhatlensis (DeKay).
Specimen examined :
From D. grossularia (van Beneden) : Off Alaskan Coast, Lat. 67‘’43.3' N, Long. 164®55' W, 90 feet,
Station 57, SS Hugh Smith, taken by diving, 1 female.
Description :
Female (fig. 47) :
Overall body length, from anlerior end of ccphalosome to posterior end of caudal rami mea-
sures 3.95 mm. The body (fîgs. 47, a, b) is eruciform, with tapering anterior and posterior ends. The
segments of the body are not articulated, but are indicated by the legs and by indentations at appro-
priate positions. The body is not particularly contracted and there are no obvious ventral projections
between the thoracic legs. The proportions of cephalosome : metasome ; urosome, are 1:6:2. The
urosome (fig. 47, c) is not dislinctly articulated or otherwise set off from the metasome. The minute
caudal rami are inserted somewhat laterally. The body is generally covered with short rows of very
fine hairs.
The head appendages are described from an en face préparation so the terms of orientation refer
to this (fig. 47, d) and not to basic anatomical relations on the body.
The antennule (figs. 47, e, f) is obscurely bimerous and obscurely articulated on the head. There
is a strong articulative line on the ventral surface but none on the dorsal surface. The portion of the
appendage corresponding to the usual basal segment has only 3 small spinules on the anterior margin.
The tapering terminal portion has 4 setae on the anterior margin and apex, accompanied by 5 setules.
The antenna (figs. 47, g, h) is obscurely bimerous. The distal portion bears 2 articulated mar¬
ginal setae, one twice the length of the other. There is a basal cuticular emargination and the larger
of the 2 apical setae has a swoUen base, thus suggesting a 4-segmented composition upon superficial
observation. Patches of spinules lie on the ventral surface.
The labrum (fig. 47, i) is a hemispherical lobe, with 8 setiform ornamentations on the dors.il
surface, near the posterior margin.
The maxülule (figs. 47, j, k) is bilobed, the unsegmented projections continuing from the base
without definite articulations. The palp portion bears 3 unarticulated setiform processes, 2 borne
apically, 1 on the anterior margin. The médial extension of the base is subrectangular in outlinc,
the distal margin formed of 2 setiform processes, one about twice as long as the other. Patches of
spinules ornament dorsal and ventral surfaces of the appendage.
The maxilla (figs. 47, 1, ra) is distinctly bimerous. Heavy sclerotizations form a distinctive
surface pattern. The surfaces hâve a general covering of relatively heavy hairs and patches of spi¬
nules. The basal article has no armature. The distal article continues into a curved claw. Set in
a subcircular ventral space there are 2 stout, short modified setae.
The 4 pairs of legs are alike (first leg, fig. 47, n). Each leg is bimerous, with the apex directed
Source : MNHN, Paris
ASCIDICOLIDAlî AND SUBFAMILIES
123
somewhat medially. The articulations of the legs on the body are indistinct but cuticular speciali-
zations outline médial areas which seem to unité the legs of each pair. The proximal article has a
heavy sclerotized plate, doubtless serving for the insertion of major muscles. On the anterior surface
of the article there are groups of short setules, emerging froin pits. The distal article has a compli-
cated apex. There is a médial tuberculated lobe, beside a sheath which encloses the opposable arti-
culated terminal spine. This spine is rétractile into the sheath. There is no fifth leg.
A complicated subtriangular structure possibly involving the sixth leg (fig. 47, o) is located
laterally on the area of the urosome corresponding to the corresponding thoracic segment. In this
species this lobe is salient. It consists of heavy sclerotized ridges and among these the oviducal aper-
ture is located centrally. Muscles attach to the sclerotized ridges and presumably fonction in the
control of the aperture.
The caudal ramus (fig. 47, p) is proportionately long, conical, with a terminal articulated spi-
niform seta. The ramus is obscurely articulated with the body.
This specimen was found in the stomach of the host, the mouth of the copepod was at the open-
ing of the esophagus into the stomach. Ovisacs, containing embryos, wholly separated from the
copepod, were found lying in the atrium, apparently anchored at their proximal ends to the outer wall
of the pharynz. The question of the identity of this specimen is a vexed one, for several important
reasons. The original description offers deûnite discrepancies from several anatomical features subse-
quently established for the genus. The segmentation of the cephalic appendages very likely was
described on the basis of superficial observations of in tolo préparations. We can draw on analogous
information from two other copepods described from the same collections by Aurivillius, namely
Doropygus demissus and Schizoproctus inflatus. The author tended to represent the general features
of habitus with great accuracy. In features of the appendages, particularly in segmentation and
numbers of éléments of ornamentation, he tended to ofîer numbers of units which differ from those
typical for the taxa involved. We strongly suspect his estimâtes for this species were excessive for
the numbers of segments in the antennule, the antenna, and the maxülule. In general outline the
appendages of our specimen correspond fairly well with his présentation. His largest specimen is
bigger than ours, but hc shows equal development of the spécifie characteristics on a much smaller
représentative as well. We are sure he represented graphically his observations on the contracted
State of the bodies and the résultant protrusions and groovings of the surfaces However, we are
oqually sure that the basis for much of these features was response to fixation procedures and they
are not a reliable basis for taxonomie treatment. The ventral protrusions of the thoracic segments
are a case in point. They very likely are artifacts, but subséquent authors hâve relied on them very
strongly in the différentiation of other taxa from E. sphinx. EnUropsis duhius Shimkevich’ would
he one example ; E. sphinx georgianus Schellenherg is another. We feel this character must be disre-
garded. Drawing on full published information we hâve concluded that there is a strong possibility
that E. dubius is conspecific ; that E. sphinx georgianus most likely is not ; and that E. sphinx Chat-
ton & Harant definitely is not (see our treatments of E. chattoni, p. 104, and E. georgianus, p. 126).
The question of the host specificity of E. sphinx is a very interesting one. There seems little
doubt of the reliability of the original détermination of the host as a species of Molgula. The finding
of our specimen in Dendrodoa grossularia is somewhat counter to the generaUty of our expérience in
the occurrence of Enteropsis species, where there seems to be some taxonomie conformity of hosts.
However, we hâve good evidence that a considérable degree of polytopy may occur, as in E. chattoni.
There are good geographical grounds for our identification of our specimen. It occurs in the same
marine basin as the original collection, the Chuckchi Sea, and we hâve found host diversity in the
Arctic ascidicole copepods to be almost a general rule. Furthermore we find the truly Arctic assem¬
blage of ascidicoles to be a very sparse one. The total numher of species cannot be very great.
Enteropsis capitulatus new species
(Figure 48)
Enteropsis sp., Dudley, 1966, p. 157, 158, 159, 160.
Source : MNHN, Paris
Fio. 48. — Enteropiii eapitulatus, new species, paratypic female : a, habitus, latéral ; b, habitus, ventral ; c, detail
of general body ornamentation, marginal j d, detail of general body ornamentation, surface view ; e, en /ace view
of oephalosome : Al = enUnnule, A2 = antenna, L = labrum, Mxl = maxillule, Mx2 = maxilla ; f, antennule,
ventral ; g, antennule, dorsal ; h, antenna, ventral ; i, labrum ; j, maxillule, ventral j k, maxilla, ventral j 1, firat
leg, anterior ; m, insémination pore, séminal tube, oviducal aperture ; n, posterior end of body showing position
of caudal rami ; o, caudal ramus. Scales for a, b = 1.0 mm ; other scales = 0.1 mm.
Source : MtJHN, Paris
ASCIDICOLIDAB AND SUBFAMILIES
125
Types :
Holotypic female, West Sound, Orcas Island, Washington, November 26, 1955, 2 females,
3.5 mm and 4.0 ram (type locality, San Juan Islands, Washington, in Boltenia villcsa (Stimpson)) :
paratypes, specimens listed below.
Specimens examined :
From Boltenia oillosa (Stimpson) :
Blake Island, Washington, May 27, 1956, 1 female, 4.0 mm.
West Sound, Orcas Island, Washington, July, 1956, 1 female, 5.5. mm.
West Sound, Orcas Island, Washington, August, 1956, 1 female, 3.3 mm.
Description :
Female (fig. 48) :
The overall body length from the anterior end of the cephalosome to the posterior end of the
caudal rami ranges from 3.3 mm to 5.5. mm, with an average of 4.1 mm in 5 available specimes of
mature females.
The body (fîgs. 48, a, b) is strongly tapered anteriorly with the notably small head set off by
a neck-like constriction or indication. The posterior taper is not strong, and the overall aspect is
somewhat fusiform. The legs are relatively inconspicuous. The metasome is rather elongate, with
the legs widely spaced. There are very slight indications of the component segmentation of the meta¬
some. The urosome is most inconspicuously set ofl from the metasome. The caudal rami are minute
and set far lateraDy. The proportions of lengths of the 3 major body régions are about 1:8:4, cepha¬
losome ; metasome : urosome. The body is generally covered with broken rows of spinules, these
somewhat more closely spaced on the ventral surface of the urosome (figs. 48, c, d).
The head appendages are described below from the orientation of an en face préparation (6g, 48, e),
so the terms of orientation used are pertinent only to this préparation and not necessarily to the general
anatomy of the body.
The antennule (6gs. 48, f, g) might be termed obscurely bimerous, since there is no articulation
of the dorsal surface, but there is a strong, well developed sclerotization, serving for insertion of the
major muscles, extending across the entire ventral face, and clearly demarcating 2 ingrédient articles
of the appendage. The basal article has a stout, curved seta, articulated on a prominence, subter-
minally on the anterior margin. There are 5 short setules, 5 heavy spinules and a general covering
of small spinules on the dorsal surface ; there is no additional ornamentation on the ventral surface.
The distal article is truncate and somewhat tapered. At about the middle of the anterior margin,
and somewhat on the ventral side, there is an unarticulated heavy seta. At its base there is oneach
surface a setule. Terminally there are a hooked seta, a filiform seta and a setule. Additional éléments
of ornamentation include proximal setules on the dorsal surface, several small spinules and some short
hairs on the distal 6fth of the posterior margin. The entire appendage is relatively heavily sclerotized.
The antenna (Bg. 48, b) is derived from 2 basic articles, but the articulation is represented only
i>y a weak fold on the dorsal surface, although ventrally there is a strongly sclerotized fold. The appen¬
dage is generally heavily sclerotized. The basal article has no spines or setae but is ornamented with
numerous spinules arranged in patches over 3 lobes along the posterior margin. Two rows of spinules
are set on the distal margin on the ventral side. The distal article is essentially conical but the pointed
tip actually represents the fused apical seta. A subapical sharply pointed process doubtless represenls
a posterior seta. Two small lobes, set with spinules, protrude on the posterior margin, one just basal
to the posterior process, one more proximal to this. The apical process has rows of heavy spinules,
the posterior process is unornamented.
The labrum (fig. 48, i) is a semicircular lobe with an unornamented entire posterior margin,
Source : MNHN, Paris
126
PAUr. L. ILLG AND PATRICIA L. DUDLEY
well sclerotized. The ventral surface is covered with rows of fine protubérances. Directed posterior
from the anterior margin of the labrura there is a row of 5 to 6 stout, somewhat curved, non-plumose
setae.
The niaxillule (fig. 48, j) is hilobed, with the palp set off by a faint articulative line on the dorsal
surface only. The entire appendage is relatively heavily sclerotized. Distally on a lobe drawn out
as base for the palp there is set a stout pointed process, doubtless representing a fused seta of the base.
There are 2 patches of spinules on the ventral surface of the base. The palp is short, flattened, sub-
rectangular and the distal margin is formed by 2 equal, stout, sharply pointed processes, these thus
triangular in oiitline and doubtless équivalent to setae of other species. The ventral surface is covered
with spinules. The médial lobe of the basal portion is about twice as long as the palp, truncate termi-
naUy, bearing on the distal margin a stout, articulated, curved, spiiiform seta accompanied by a shorter
seta which is unarticulated. The ventral surface is covered with spinules.
The maxilla (fig. 48, k) is bimerous and distinctly articulated on the body. Heavy scleroti-
zations form a distinctive pattern on both articles. The surface of the sclerotized areas is uniformly
pitted and each pit surrounds a small hair. The unsclerotized surfaces are ornamented by rows of
spinules. The appendage in general conforms to the generic characteristics. The basal article has
no setae or spines but is produced at the distal medial corner as a short conical process which probably
opposes the movable terminal hook. The distal article, complexly jointed, is tapering in outline and
curved slightiy at the tip, forming a stout, flattened hook. Set in a subcircular, unsclerotized space
on the ventral side there is a short seta, surrounded at its base by a sclerotized ring. There are several
(about 4) heavy spinules distally, on the ventral face of the hooked tip. There are several rows of
spinules just distal to the seta.
The 4 pairs of legs (first leg, fig. 48, e) are much alike, ail bimerous and uniramous. The whole
appendage is covered with genorally distributed fine hairs. The proximal article has no setae or spines
but has heavy sclerotizations, probably distinctively arranged. The distal article is sclerotized overall,
with a complicated apical structure. There is a medial tuberculated lobe opposing an articulated
spine, which is invested except at the tip, when retracted, by a cuticular covering.
There is no fifth leg.
A complicated structure, perhaps involving the sixth leg (fig. 48, m) is proximal to the posterior
margin of the first urosomal segment near the middle of the latéral surface. This consists of 2 lobes,
containing sclerotized zones, which on each side surround a sclerotized flap, which covers the génital
aperture. There is a mid*ventral insémination pore at the posterior margin of the first urosomal
segment from which diverging séminal tubes lead to the area of the latéral oviducal apertures.
The caudal rami (figs. 48, n, o) are minute and set somewhat ventrolaterally on the rounded
posterior end of the body. Each is a heavily sclerotized flattened lobe covered with heavy spinules.
The coloration is an overall orange, with a bright red eye. The embryos in the egg strings are
bright purplish-pink. Two specimens, one mature but non-ovigerous, one with egg strings, were
taken from the intestines of the host tunicates. The intestinal wall encased the copepod very firmly
m each case, and even in living matériel had to be picked away. The ova in the oviducts of these
individuals were garnet red.
Enteropsis georgianus Schellenberg, 1922
(Figure 49)
Enteropsis sphinx Aurivillius var. georgianus Schellenberg, 1922, p. 291, 295 (type locality, South
Georgia, from Polyzoa pictonis var. georgiana (Michaelsen).
Distribution :
South Georgia and Patagonia.
Source : MNHN, Paris
Source : MNHN, Paris
PAUL L. ILLG AND PATRICIA L. DUDLEY
Host :
Polyzoa opuntia Lesson, subspecies ? (= Polyzoa piclonis).
Specimens examined ;
From Polyzoa opuntia Lesson : San Julian, Patagonia, Argentina :
ÜSNM No. 10565, Accession Number 97902, Station 123, May 6, 1927, collector W. L. Schmitt,
id. W. G. Name, 1 female.
We ofîer below what we consider to be pertinent supplementary information to the description
of ScHELLENBERG and present our Ogures for comparison. Our specimen, an adult female, measures
2.5 mm in overall length, as compared to Schellenbebg’s 4 specimens, which ranged from 2.7 to
3.2 mm. The proportions, cephalosome ; metasome : urosome, are 1 : 4 : 2, as compared to Schel¬
lenbebg’s 2:9:7. We present a figure of the habitus for the first time (fig. 45, a). The urosome
(figs. 45, b, c) has a midventral insémination pore set on a raised area ; diverging séminal tubes pass
laterally toward the oviducal apertures. The antennule indicated by the author as being bimerous
or trimerous, with a seta on the basal segment and several on the outer margin of the tip, is definitely
bimerous in our specimen (fig. 49, d). The terminal segment is approximately as long as the basal
segment but only two-thirds as wide, but the appendages agréé in other details. The antenna (fig. 49, e)
is bimerous and has 2 subequal spinulose terminal setiform processes. Schellenberg stated that
the appendage had 4 segments and that the terminal éléments were unequal. Although this would
seem to be a distinct différence between our specimens, we feel the matter of interprétation of segmen¬
tation of the head appendages has not been resolved satisfactorily. We are sure there are many
instances of accepting superficial grooves and folds and partial articulative lines as expressions of
fuU segmentation. We hâve tried to take a consistent position of describing segment numbers based
only on defînite interruptions in the cuticle. The labrum (fig. 49, f) bears 6 setiform ornamentations,
concurring with Schellenberc’s indications. The maxillule, termed mandible by Schellenberg,
is bilobed (fig. 49, g). We find bis terminology of the “ rami ” more or less the reverse of our own.
In the en face view, the most ventral lobe is the palp, thus latéral (he called it endopodite). In our
specimen it terminâtes in 2 equal unornamented setiform processes. At the base of this lobe, not
the “ endopodite ”, there is a stout hairy setiform process, born on a lobe which is strongly set off.
The médial extension of the base terminâtes in a broad flattened hairy setiform process, which is accom-
panied by an articulated narrower and shorter hairy seta. The appendage has a strongly set off basal
article. Patches of spinules ornament the surfaces of the base and the palp. The maxilla, termed
maxilliped by Schellenberg, concurs essentially with his description (fig. 49, h). The legs (first
leg, fig. 49, i) hâve elongate basal segments, measuring about 3 times as long as the terminal segments,
and about twice as wide. The apex of the terminal segment bears the usual characteristic hooded
spine. The caudal ramus (fig. 49, k) is elongate, conical, ending in an abruptly constricted flexible
setiform tip. The oviducal apertures (fig. 49, j) are found laterally on the urosome. They occur
on subtriangular lobes. Complex sclerotization to which muscles attach surround the apertures.
As we pointed out above we find no great difficulty in reconciling our specimen with Schel-
lenberg’s indications. Our study of several species within the genus furnishes us the background
for the opinion that the différentiation from E. sphinx is essentially at the spécifie level.
Type :
Enteropsis ABBOTTi new species
(Figures 50, 51)
Holotypic female, from Academy Bay, Santa Cruz Island, Galapagos Islands, from stomach
or anlerior intestine of Styela sp., collected under intertidal rock, by D. P. Abbott and J. L. Barnard,
8 February, 1964.
Source : MNHN, Paris
Fio. 50. _ Enleropaia <i66o<(t, new spccies, holotypic female : a, habitus, latéral; b, posterior end of body, latéral;
c, head, ventral ; d, antennule and antenna ; e, antennule ; f, antennule from other side ; g, antenna ;
h. antenna, another view ; i. antenna, apical portion ; j, labrum ; k, maxillule, dorsal ; 1, maxillule, ventral ;
rn, maxillule of other side, ventral ; n, maxilla ; o, maxilla, another view. Scales = 0.1 mm.
Source : MNHN, Paris
130
Fie. 51. — EnUropsiê abboUi, new apecies, holotypic female : a, firsl leg; b, second legj c, third Icg; d, fourth Icg ;
e, inaemination pore, semiaal tube and oviducal aperturle, latéral. Scales = 0.1 mm.
Specimen examined ;
Holotype, unique specimen.
Female (figs. 50, 51) :
The overall body length from the anterior end of the cephalosome to the posterior end of the
urosome is 3.0 mm. The body (fig. 50, a) is almost unîformly cylindrical, with gently curved apices,
and held in a gently curved posture in our specimen. The egg sacs were lacking. There are some
sligbt indentations of the cuticle but they do not seem to indicate directly the component segments
of the body. The cephalosome is very slightly set off dorsally by indentations. The boundary of
the metasome and the urosome is not at al! clearly indicated and the major evidence of the composi¬
tion of the latter is the pair of prominent pair of oviducal apertures (figs, 50, b, e). There are no caudal
rami. By rough estimâtes the body régions measure in the ratio cephalosome : metasome : urosome
about 1:7:2.
The appendages correspond well to the generalizations given in the diagnosis of Enteropsis
with the following spécifie features : Thé antennules (figs. 50, d, e, f) are simplified in outline, unseg-
mented, ornamented with 5 about equal setae arranged around the apex ; there are no other anten¬
nules just like thèse in the species we hâve studied. The antennae (figs. 50, d, g, h) show no apparent
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
131
segmentation, although there is a strong line across one face (fig. 50, g). The sclerotized apex is pro-
duced in a strong pointed process. The labrum {fig. 50, c) shows distinctive characters in consisting
of a rounded lobe, with strong seta-like processes inserted at the proximal corners, and without margi¬
nal setiform teeth, so that the only species to which it bears any resemblance in this regard is E. minor.
The maxillules (figs. 50, k, 1, m) are unique although the basic composition is of the general type for
the genus. There is a basal portion with indications of 2 component articles. These bear marginal
spiniform projections and the more distal article is also much produced about midway on the margin.
There are 2 diflerentiatcd apical processes, not completely articulated on the more distal article, one
of these setiform, broad basally and with plumose margin. The other is very distinctive with a some-
what spoon-like outline of the terminal portion, and with sclerotized margins. One of the two usual
major lobes is very little developed. Probably the process representing the palp in other species here
is reduced to a pointed process on the basal portion of the appendage.
The maxillipeds (Ggs. 50, n, o) are not especially distinctive. The basal portion bears rather
long hair-like ornaments on one margin.
The first to fourth legs (figs. 51, a, b, c, d respcctively) do not show specifically distinctive cha-
racteristics. As is true of much of the general surface of the body these appendages bear lines of fine
hair-like ornaments.
The insémination apparatus and the complications of the cuticle at the apertures of the ovi-
ducts (fig. 51, e) do not show markedly distinctive features. The absence of the caudal rami is unusual
for the genus.
We dedicate this interesting species to Dr. D. P. Abbott, who sent it to us, in acknowledgment
of this and many other favors we owe him.
Enteropsis ONYcnopnoRUS Schellenberg, 1922
Enteropsis onychophorus Schellenberg, 1922, p. 291-292, 295 (type locality Tauranga, New Zealand,
From Alloeocarpa llnlenii Micliaelsen).
Distribution :
New Zealand.
Ilost :
Alloeocarpa thilenii Michaelsen.
There hâve been no subséquent reports of this species. We feel the description of the fifth
legs for the species is misleading. From observations on other species of Enteropsis we are sure that
Schellenberg observed structures connected with the oviducal apertures and termed them fifth legs.
We are convinced that the latter appendages are lacking throughout the genus.
INDETERMINABLE SPECIES
Enteropsis dübius Shimkevich’, 1889
Enteropsis dubius Shimkevich’, 1889, p. 75-92, pl. 3, fig. 1-17, pl. 4, fig. 19-27, 34, pl. 5, fig. 53-58 (type
locality, White Sea, from Molgula groenlandica). — Canu, 1892, p. 53, 91, 92, 215, 220. —
Schimkewitch, 1896, p. 342, 345, 352, pl. XV, fig. 36-38. — Chatton & Brémont, 1909b, p. 201-
202. — Chatton & Harant, i922b, p. 161. — Schellenberg, 1922, p. 290, 295. — Sewell, 1949,
p. 192.
Source : MNHN, Paris
132
PAÜI- JL. ILLG AND PATRICIA L. DUDLEY
EXCLUDED SPECIES
Enteropsis pilosus Canu, 1886
Enteropsis pilosus Canu, 1886a, p. 365-374, pl. III, fig. 1-5 (type locality, vicinity of Iles Glenans,
Concarneau, France, from Diasona kebridica [Forbea]) ; 1892, p. 52-53, 67, 143, 219, 220, fig. 4,
17. — Chatton & Bréinent, 1909b, p. 197, 201. — Hartmeyer, 1911, p. 1735. — Schellenberg,
1922, p. 290, 295. — Chatton & Harant, 1922b, p. 160-161. — Harant, 1931, p. 371. — Sewell,
1949, p. 184, 188. — Gotto, 1960, p. 226 ; 1961, p. 151, 152.
Gotto, 1961, called attention to dilTiculties he encountered in trying to deal with E. pilosus.
Our study of the literature and comparison with specimens of other species leads us to conclude that
this species cannot be classified in the genus Enteropsis. The head appendages correspond excellently
to the generic characters. However, the thoracic legs, particularly the description of fîfth legs, defi-
nitely give grounds for exclusion. It is tempting to speculate that there was some confusion of spe¬
cimens involved in Canu’s original description. It gives every evidence of hasty assembly and there
is some distinctly garbled information presented. In two different parts of the paper he compares
the thoracic appendages to those of Aplostoma (sic) and explicitly States he finds the fifth legs to cor¬
respond with those of “ Aplostoma ”.
Enteropsis vababensis T. Scott, 1901
Enteropsis vararensis, T. Scott, 1901b, p. 241-242, pl. XVII, fig. 28-34 (type locality, Moray Firth,
Scotland, from branchial chambers of Botryüus sp.). This species was correctly assigned to
Mychophilus by Chatton & Brément, 1909c, p. 239, and to the synonymy of M. roseus by
Sars, 1921, p. 79.
SUBFAMILY HAPLOSTOMINAE Chatton & Harant, 1924
This subfamily, proposed by E. Chatton & H. Harant (1922-1924) for a small group of closely
allied généra, was reviewed by Ooisiii & Illg (1977) in connection with description of a sériés of new
species. Essential to the préparation of these descriptions were extensive developrhental studies on
species of the general Haplostoma, Haplostomides, Haplostomella. The results of these, to be published
in a paper now in préparation following a lead established by Dudley (1958, 1966), and the findings
included in the présent paper hâve been crucial to our morphological and taxonomie conclusions in
treating our diverse collection of species as a phyletic assemblage.
We repeat here, for completeness, selected strategie référencés from our synonymies of the sub¬
family and généra and keys to the généra and species. We refer the interested reader to the fuU treat-
ment in the paper cited.
Ascidieolidae (part). — Canu, 1891, p. 475 ; 1892, p. 186. — Gotto, in Anderson & Rossiter, 1969,
p. 464.
Ascidicolidés (part), Brément, 1909, p. 61-62, 86-87.
Ascidicolmae (part), Chatton & Brément, 1915, p. 143, 144. — Schellenberg, 1922, p. 220, 277-281.
EnterocoUdae (part), Sars, 1921, p. 73-74. — Blake, 1929, p. 6 ; 1933, p. 226. — Lang, 1948, p. 25-27.
— Monniot, 1962, p. 570. — Dudley, 1966, p. 155. — Gotto, 1966, p. 193.
Source : MNHN, Paris
ASCIDICOLIDAB AND SUBFAMILIES
133
Notodelphyidae (part), Sewell, 1949, p. 174.
Aplostomiens Canu, 1886, p. 373-374.
Haplostomiens, Chatton & Harant, 1922c, p. 250-252 ; 1924d, p. 406-407.
HaploBtominae, Chatton & Harant, 1924e, p. 415, 416-421. — Gotto, 1959, p. 9-10. — Ooishi & ïllg,
1974, p. 365 ; 1977, p. 12-15.
KEY TO GENERA OF THE SUBFAMILY HAPLOSTOMINAE
1. Exopod of legs 1 to terminating in a single lobe, ornamented with 1 or 2 thorn-like spines
apically. 2
1'. Exopod of legs 1 to 4 terminating in 2 lobules, one of them setiferous apically.
Haplostomella, p. 135
2. AH mouthparts présent. 3
2'. One or more pairs of mouthparts lacking. Haplostoma, p. 133
3. At least some of the cephalic mouthparts partially setiferous. Haplostomides, p. 134
3'. Mandible, maxillule and maxilla reduced lobes without normal setae. Haplosaccus, p. 135
Haplostoma (Canu, 1886)
EnUrocola (part), Norman, 1869, p. 300.
Aplostoma Canu, 1886a, p. 313-320 (types species, by monotypy, A. brevicauda Canu, 1886) (not Aplos-
toma Mouquin-Tandon, 1856) ; Canu, 1886b, p. 1025-1027 ; 1891, p. 471, 474, 475 ; 1892, p. 220-
223. — T. Scott, 1906, p. 363-364 ; 1907, p. 361, 369-370. — Brément, 1909, p. 78-87. — Chat¬
ton & Brément, 1909b, p. 228 ; 1910, p. 80-81, 88-92. — Schellenberg, 1922, p. 288-289. — Sal6,
1926, p. 1-2. — Neave, 1939, p. 256. — Dudley, 1966, p. 155, 157, 158, 160.
Cryplopodus Hesse, 1865, p. 237-241, 255 (unidentifiable genus for 2 unidentiûable species ; no type
designated). — Canu, 1892, p. 222 ; unidentifiable genus. — Sars, 1921, p. 74-75 (part). —
Blake, 1929, p. 6 ; 1933, p. 226. — Lang, 1948, p. 3.
Tranesloma Wilson, 1924, p. 14, for Aplostoma Canu, preoccupied (type species, by monotypy, Aplos¬
toma brevicauda Canu, 1886) ; 1932, p. 601 (in key).
Haplostoma Chatton & Brément, 1915, p. 144, 145, 153. — Chatton & Harant, 1922c, p. 249-252 ;
1924b, p. 363; 1924c, p. 399, 405; 1924d, p. 407; 1924e, p. 413, 418-419. — Harant, 1931, p. 371.
— Wilson, 1932, p. 598, 600 (in key). — Neave, 1939, p. 570. — Gotto, 1952, p. 674 ; 1954,
p. 665 ; 1959, p. 9, 10 ; 1960, p. 216 (in key) ; 1966, p. 193 ; 1970, p. 272. — Monniot, 1962,
p. 573. — Ooishi & Illg, 1974, p. 365 ; 1977, p. 15-21.
KEY TO SPECIES OF HAPLOSTOMA, BASER ON FEMALES
1. One pair of mouthparts (maxillules) lacking. elegans Ooishi & Illg, 1977
1'. Two (maxillules and maxillae) or 3 (mandibles, maxillules and maxillae) pairs of mouthparts
lacking. 2
Source : MNHN, Paris
134
2 .
2 '.
3.
3'.
4.
4'.
5.
5'.
6 .
6 '.
7.
7'.
8 .
9.
9'.
10 .
10 '.
11 .
11 '.
PAUL L. ILLG AND PATRICIA L. DUDLEY
Armature of terminal segment of antenna including 3 spines. 3
Armature of terminal segment of antenna including 4 spines. 4
Mandible with 1 small seta. setiferurn Ooishi & Illg, 1977
Mandible with 2 small setae. gibberum (Schellenberg, 1922)
Posterior margin of labrum with processes. 5
Posterior margin of labrum without processes. 9
Posterior margin of labrum with 8 distinct processes. denialum Ooishi & lUg, 1977
Posterior margin of labrum with 4 to 6 processes. 6
Ëxopod of legs 3 to 4 with 4 spines. 7
Exopod of legs 3 to 4 with 2 spines. 8
Posterior margin of labrum with 4 indistinct processes. ambiguum Ooishi & Illg, 1977
Posterior margin of labrum with 6 distinct processes. banyulensis (Brément, 1909)
Posterior margin of labrum with 6 distinct processes ; body length beyond 2 mm.
eruca (Norman, 1869)
Posterior margin of labrum with 6 indistinct processes; body length less than 1.5mm.
miniUum Ooishi & Illg, 1977
Exopod of legs 3 to 4 with 2 spines. albicalum Ooishi & Illg, 1977
Exopod of legs 3 to 4 with 3 or 4 spines. 10
Legs 1 to 4 without distinct endopod lobe. canui Chatton & Harant, 1924
Legs 1 to 4 with endopod represented by subconical lobe. 11
Fifth leg with 2 apical setules. brevicauda (Canu, 1886)
Fifth leg with 3 apical setules. mizoulei Monniot, 1962
Haplostomides Chatton & Harant, 1924
Enlerocola (part), T. & A. Scott, 1895, p. 359-360.
Aplostoma (part), Brément, 1909, p. 84, 85. — Chatton & Brément, 1909b, p. 228 ; 1910, p. 84-86. —
Schellenberg, 1922, p. 228, 289.
Haplostoma (part), Chatton & Harant, 1924b, p. 363. — Gotto, 1959, p. 10.
Cryplopodus (part), Blake, 1929, p. 6; 1933, p. 226.
Haplostomides, Chatton & Harant, 1924d, p. 406-412 (type species, by original désignation, H. scoUi
Chatton & Harant, 1924) ; 1924e, p. 418. — Harant, 1931, p. 371. — Wilson, 1932, p. 600
(in key). — Neave, 1939, p. 570. — Gotto, 1952, p. 674 ; 1954, p. 665, 666 ; 1960, p. 213, 216 ;
1966, p. 193 ; 1970, p. 271. ~ Ooishi & Illg, 1974, p. 365 ; 1977, p. 77-78.
KEY TO SPECIES OF HAPLOSTOMIDES, BASED ON FEMALES
1. Armature of terminal segment of antenna consisting of 2 spines.
kibernictis (T. & A. Scott, 1895)
1'. Armature of terminal segment of antenna consisting of 4 spines. 2
2. Maxilla with 4 setae. luteolus Ooishi & Illg, 1977
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAHILIES
135
2 '.
3.
3'.
4.
5.
5'.
Maxiila wilh 2 or 3 éléments of armature..
Maxilla with 2 setae .
Maxiila with 3 variously developed éléments of armature
Mandible with 3 apical setae.
Mandible with 2 apical setae.
Mandible with 3 apical setae.
Mandible with 2 apical setae.
3
4
5
scotti Chatton & Harant, 1924
bremenli Chatton & Harant, 1924
. amarouci (Blake, 1929}
. beÜus Ooishi & Illg, 1977
Haplosaccus Chatton & Harant 1924
Aplostoma (part), Chatton & Brément, 1910, p. 86*92. — Schellenberg, 1922, p. 289. — Salfi, 1926,
p. 1.
Haplostoma (part), Chatton & Harant, 1922c, p. 249, 251, 252.
Haplosaccus, Chatton & Harant, 1924e, p. 413, 415, 419 (type species by original désignation, Aplos-
toma sacculus Chatton & Brément, 1910). — Harant, 1931, p. 371. — Wilson, 1932, p. 600
(in key). — Neave, 1939, p. 569. — Gotto, 1960, p. 227 ; 1970, p. 271. — Ooishi & Illg, 1974,
p. 365 ; 1977, p. 88*90.
KEY TO SPECIES OF HAPLOSACCUS, BASED ON FEMALES
1. Antenna with 1 terminal spine ; exopod of legs 1 to 4 with 1 simple claw-Uke element.
sacculus (Chatton & Brément, 1910)
1'. Antenna with 1 terminal and 1 subterminal spine ; exopod of legs 1 to 4 with 1 deeply and equally
bifurcated claw-like element (2 spines). elongatus Ooishi & Illg 1977
Haplostomella Chatton & Harant 1924
Aplostoma (part), Chatton & Brément, 1910, p. 82-84, 89-91. — Schellenberg, 1922, p. 289. — Salfi,
1926, p. 1-4. — Sewell, 1949, p. 174.
Haphstomella, Chatton & Harant, 1924c, p. 398-406 (types species, by original désignation, H. mala-
cocera Chatton & Harant, 1924) ; i924e, p. 413, 417, 419-421. — Wilson, 1932, p. 601 (in key). —
Neave, 1939, p. 570. — Gotto, 1969 (in Anderson & Rossiter), p. 464 ; 1970, p. 267-272. —
Ooishi & Illg, 1974, p. 365-374 ; 1977, p. 96-99.
Bhabdomcrpha, Fukui, 1965, p. 61-63 (type species, by monotypy R. kalocynthiae Fukui, 1965).
KEY TO SPECIES OF HAPLOSTOMELLA, BASED ON FEMALES
1. Postérolatéral protrusion for fifth leg distinct ; mandible developed as lobe. 2
r. Postérolatéral protrusion for fifth leg indistinct j mandible reduced into spiniform projection
or absent. ^
2. Mandible unimerous with 1 terminal seta ; exopod of legs 2 to 4 with 1 terminal seta.
tuberculata Chatton & Harant, 1924
Source ; MNHN, Paris
136
2 '.
3.
3'.
4.
4'.
5.
5'.
6 .
6 '.
7.
7'.
8 '
9.
9'.
PAUL L. ILLG AND PATRICIA L. DUDLEY
Mandible partially biraerous each with 1 seta ; exopod of legs 2 to 4 with 2 terminal setae....
dubia Ooishi & Illg, 1977
Antennule cylindrical with rounded end ; body lacking indentations or folds to suggest body
régions or segments. reducta Ooishi & Illg, 1977
Antennule conical ; body with or without indentations or folds to suggest body régions or segments.
Urosome relatively short, comprising about 1/7-1/10 body length. 5
Urosome relatively long, comprising about 1/3 body length. 8
Caudal ramus lacking setae. sycozoae (SaIG, 1926)
Caudal ramus with setae. 6
Antennule 4-segmented ; caudal ramus with 1 terminal seta.
magellanica (Chatton & Brément, 1910)
Antennule unsegmented ; caudal ramus with 5 setae. 7
Maxilla with 1 seta ; ûfth leg with 2 seta and 1 spine.. malacoeera Chatton & Harant, 1921
Maxilla with 2 setae ; fifth leg with 2 setae. oceanica Ooishi & lUg, 1977
Each metasomal segment with well-developed dorsolateral plates of suboval outline. 9
Each metasomal segment with weakly-developed dorsolateral plates of subtriangular outline
disiincta Ooishi & Illg, 1977
Urosome 4-segmented. australiensis Gotto, 1970
Urosome unsegmented. kalocyntkiae (Fukui, 1965)
SUBFAMILY BOTRYLLOPHILINAE NOMEN CONSEBVANDUM
Kossmectridi Délia Valle, 1883, p. 252 (new family).
Ascidîeolidae, Gerstaecker, 1870-1871, p. 719 (part). — Canu, in Giard, 1888, p. 505 (part). — Canii,
1891a, p. 472, 475 (part) ; 1892, p. 107, 186 (part). — T. Scott, 1901a, p. 351-352 (part) ; 1901b,
p. 241-245 (part;. — Thompson & Scott, 1903, p. 255, 273 (part).
Ascidicolidés Brément, 1909, p. lxxxvi (part).
Schizoproctidae Aurivillius 1885a, p. 246-247 ; 1885b, p. 282.
Notodelphyidae, Norman & Scott, 1906, p. 201, 203 (part). — Schellenberg, 1922, p. 219-220. —Bar-
nard, 1955, p. 237 (part).
Botryllophilidae Sars, 1921, p. 66-67. — Wilson, 1932, p. 391. — Leigh-Sharpe, 1934, p. 216 (part).
— Pesta, 1934, p. 8. — Lang, 1948, p. 23, 25. — Sewell, 1949, p. 20, 145, 156-158. — Rose &
Vaissière, 1953, p. 90. — Bresciani & Lützen, 1962, p. 374. — Dudley, 1966, p. 155, 160. •—
Gotto, 1966a, p. 193. — Lützen, 1968, p. 101. — Stock, 1970, p. 1, 16-17.
Botryllophidae, Rose & Vaissière, 1953, p. 90.
Ascidieollnae, Schellenberg, 1922, 219-220 (part). — Brehm, 1927, p. 490 (part). — Bernard, 1955,
p. 237 (part).
Doropygidae, Sewell, 1949, p. 169-170.
The concept of a family Botryllophilidae would seem to be a fairly incisive one but there are some
complications in the history and a sériés of equivocations in assigning the genus Botryllophilus to a
higher category. Much of the difFicuIty stems from the fact that the genus was described by Hesse
(1864) and there are the usual problems of reconciling his statements and Ggures with any actualj spe-
cies. The identity of the principal host, Botryüus sp., is reasonably Grm, and the indications he gave
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
137
were descriptive enough that few authors hâve hesitated to recognize his genus, although he described
a whole sériés of species which cannot be evaluated with any degree of reliability. For the majority
he attributed as host “ un botrylle ” and, moreover, he did not fix a type. This was donc in the satne
year as the first description by Bâte (1864), who selected B. ruber Hesse, 1864. There has never
been a full-scale description nor complété set of figures presented for this species based on unequi-
vocally identified material and so comparisons and additions within the genus hâve rested upon a very
uneasy base. Moreover, there are some most peculiar variabilities in the morphology of most species.
Indeed, Lang (1948) was so impressed by the variabüity of a sériés of material he collected in Sweden
that he deoided to synonymize ail the species described, except for a couple of notably distinctive
forms, within the original species. There seem to be well validated cases of essentially the same mor-
phological type occurring either in compound or simple ascidians.
Della Valle (1883) described a species of Botryllopkilus, B. notopxis, but either did not know
of Hesse’s Work or, as did many reputable taxonomists, chose to ignore it. He proposed a genus,
Kossmechthrus, for his species, and was suffîciently impressed with the distinctiveness of its characters
that he also stated the genus should be assigned to a family of its own, termed by him Kossmectridi.
No author foUowed this usage. In 1885, Aürivillius, in describing the striking genus Schizoproctus,
also erected a family for, it, namely the Schizoproctidae. This genus, highly distinctive, clearly finds
it closest afFinities among the species of Botryliophilus and the 2 généra should be referred to the same
subfamily. Most workers active just after this time either overlooked these familial terms or simpiy
assigned Botryllopkilus and/or Schizoproctus to the Ascidicolidae or Ascidicolinae (Canu, 1892 ; Scott,
1901 ; ScHELLENBEKG, 1922, CtC.).
Sabs (1921, p. 66) used Botryllophilidae for Botryllopkilus, Schizoproctus, and a new genus
Pteropygus, without any diagnosis or discussion of the family. Most authors after Saks hâve used
the family name in his sense and attributed the concept to him. There are a dozen or so such citations,
and although the group is not such a conspicuous one that the matter is of any great conséquence, it
would appear most practical to continue this usage. We accept it here with the modification of
assigning the taxon subfamilial level within the Ascidicolidae.
As well as the somewhat clouded nomenclatural history, certain zoological considérations com-
plicatc the status of the taxon. The latter are the more interesting. Although few valid instances
are known, some of the males of Botryllopkilus species are uncannily similar to males of species of
Haplosloma. If no other taxa were known, there would seem to be very good grounds to include the
two généra in a single subfamily. But the two seem to stand at an interesting taxonomie divergence.
In the first place there are two types of males in Haplostoma (Ooishi & Illg, 1977) and the more modi-
fied type is readily distinguished from Botryllopkilus. Moreover, we are informed by Dr. J. Stock
(in correspondence) that there is a dimorphism of males within Botryllopkilus. We hâve seen no
pertinent material among our collections. More substantially, there is little doubt of the close rela-
tionship of Haplostonia and Haplostomslla and in the latter the males hâve taken on a much modified
anatomy (Ooishi & Illg, 1977), removing them considerably from the condition in Haplostoma-
Botryüophilus. In another direction, the male of Schizoproctus, depicted by Sabs (1921), is very
different from the male of Botryllopkilus, tending to show a doser correspondence to the features of
the female.
Diagnosis ;
Female :
The body in the adult is inflated, with varying degrees of expression of ingrédient segmentation.
The cephalosome bears the head appendages and the maxillipeds ; usually there is a very substantial
démarcation behind these appendages. The metasome usually consista of évident (sometimes quite
clearly demarcated) thoracic segments bearing the so-called swimming legs (thoracic segments 2-5).
The segment of the fourth legs is coalesced to a substantial degree with that of the fifth legs. Basi-
cally the urosome consists of the seventh thoracic segment, the abdominal segments and the terminal
anal somite bearing the caudal rami. Commonly there are substantial modifications because of the
Source : MNHN, Paris
138
PAUL L. ILLG AND PATRICIA L. DUDLEY
coalescence of 2 or more segments at the metasome-urosome boundary. The segment of the fourth
legs is coalesced to a substantiel degree with that of the fifth legs. In some forms the segment bearing
the génital apertures (thoracic segment 7) fuses essentially completely with the next anterior segment.
As a resuit the usual démarcation between metasomc and urosome may be obscure with no true major
hody articulation at this level. The urosome then consiste of the true abdominal segments, the number
varying according to species. Frequently additional annular cuticular constrictions lead to an appea<
rance of multiplication of the number of urosomal segments. The end piece of the urosome, hcre
called the anal somite, which bears the caudal rami, is doubtless some sort of a complex rather than
a simple telson (Dudley, 1966, p. 10) and thus surely also not a regular segment. The caudal ramua
bears 4 (typically) subequal, strong, clawed setae, and usually 1 or more additional setae.
The rostrum may or may not be conspicuously developed but there is usually some sort of
prominence between the base of the antennulcs.
The antennules are short, usually massive basally, with a short, slender terminal lash-like
component composed of rather few free or fused segments. Ail the segments bcar setae, and these
are so abondant as to suggest that the appendage dérivés morphologically from oiie of a much higher
degree of segmentation. It is very characteristic that some of the setae, parlicularly of the basal
segments, are borne singly or in groups on protubérances of the segments proper.
The antenna is conspicuous, relatively elongate and slender, with a strong articulation at the
middle, indicating a considérable freedom of action of the terminal portion relative to the base. At
this articulation there is often a considérable complication of the configuration of the cuticular pièces.
At one extrême the appendage is clearly 2-segmented. At the other, a separate article appears to be
delimited between the basal and terminal segments. Evidence is not available as to whether this is
a real segment or an articulating ring somehow related functionally to activities of the appendage.
However, the development is often so pronounced that the best descriptive désignation is as a 3-seg-
mented appendage. Ornamentation is restricted to the terminal article. There are some spines
inserted at- intervals, sometimes in shallow emarginations, on the médial margin, and a group of several
around the truncate apex. Probably these éléments are derived from setae, but there is a gradua¬
tion of modification resulting in a sériés of éléments varying from substantially setiform character
to well-developed spines. Most species hâve 7 or 8 of these éléments. There is typically some addi¬
tional ornamentation of spinules, near the bases of the spines or on the general surface in some sort
of pattern. These appendages usually show the asymmetry characteristic of many of the appendages
in the genus ; the différentiations tend to be in configuration of the segments and in the aspect of the
ornamenting éléments.
There is usually no élaboration or complication of the structure of the labrum, which simply
consists of the superior margin of the mouth opening.
The mandibles are symmetrical, always bearing basally a coxal masticatory lamella. The
segmentai composition of the appendage distal to the coxopodite is obscure. Essentially there is a
substantiel setiferous palp. Laterally on this at about mid-margin there are 3 setae, often on an
unarticulated lobe, or in some cases a lobe with 2 setae and a third at its base, these apparently repro-
senting the exopodite. About 5 setae around the more distal portion probably correspond, in part
at least, to éléments of an endopodite.
The maxillules are symmetrical and show again a coalescence of éléments. A médial lobe,
probably an endite of the coxopodite, bears a row of graduated setae, up to 7, often fewer, in number.
The distal portion of the appendage forms a palp with some lobes and setae which are probably indicative
of the basic composition. Medially and subterminally 2 setae probably correspond to the armature
derived from the endopodite ; this usually extends laterally. Some latéral lobes and setae may repre-
sent the exopodite and epipodite. One of these setae is distinctive in protruding proximally from a
latéral protubérance.
The symmetrical maxillae are reduced but are less modiefid than the other mouthparts. The
outline is triangular and several setae are set along the médial margin and at the apex. There are
several segments represented, 3 or 4 clearly seen in some species. The larger médial setae are borne
singly or in pairs on protrusions, usually without clear articulations of the setae on the bases. Smaller
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
139
setae insert singly on these bases. There is a characteristic apical pattern of a medially directed long
seta, with a very wide base and no articulation, this base serving for support of a much smaller distally
directed apical seta, this in turn accompanied by a latcrally directed small subapical seta.
The symmetrical maxillipeds are characterized by massive bases, and terminal articulated claws.
The exact number of segments has not so far been determined. Two well-niarked large segments form
the massive base. The much tapered articulation for the terminal claw involves an articulating ring,
sometimes rather complex, and whether ail this represents a single segment and whether the claw
ia a segment or alternatively a much transformed élément of ornamentation, remain to be determined.
There may be a very few reduced setules or spinules furnishing additional ornament on the appendage,
and in some cases there are patches or rows of spines, spinules, tubercles, or soine similar élément.
The first to fourth legs, the so-called swimming legs, vary from symmetrical to asymmetrical,
the latter in the majority of species, perhaps ail the species of the genus Botryllophilus. The patterns
of symmetry are specifically variable. This is also the case with the segmentation oftherami. However,
in ail cases the rami represent only one or at most 2 segments. These are typically set with ornamenting
éléments, with a great tendency for these to hâve the aspects of spines. In some species there are
surely some individual variations in the number of segments of the rami and also in the number of
ornamenting éléments.
The appendages of the sixth thoracic segment, the so-called fifth legs vary in the species and
their exact anatomy is not clearly understood as yet. They are always displaced very considerably
dorsally, and take a broad origin on the body segments without a definite articulation. It becomes
very difficult to say whether these pediform projections represent only the fifth legs or also involve
an extension into a protruding process of a portion of the body segment. The presence of a basal
seta very commonly would scem to indicate there is a basal segment, but this is completely coalesced
with the body proximally and with the reraainder of the appendage distally. Terminally there tend
to be 3, sometimes fewer ornamenting éléments, these usually short and slender, varying from spini-
fonn to setiform. In the consideraÛe gamut of aspects of these pediform projections there are
alternatively the aspect of a slender process, with the terminal ornaments rather prominent,on the one
hand, and at the other extreme the enlargement of the process into a subcircular voluminous lamella,
with the ornaments small or lacking.
The génital segment shows consistent structural features The insémination pore is midventral,
with diverging internai tubes leading laterally to séminal réceptacles near the oviducal apertures. The
latter usually involve a cuticular flap, in each, which may hâve some minor features of ornamentation.
There are characteristically 2 egg sacs, each firmly fastened to the apparatus at the gonopore and the-
refore taking a dorsolateral position. The fifth legs do not usually seem to form a firm attachment
to the sacs, although in the case of pediform processes, they closely subtend them, and in the case of
lamelliform processes they may completely envelop the egg sacs.
Male :
It is difficult to characterize the male at the subfamily level because so little is known. We do
hâve information on a male of Botryllophilus, astonishingly like that of Haplostoma, and a rather
typically cyclopoid swimming form. Sars illustrâtes very convincingly for Sckizoproctus inflatus
(1921, pl. XXXIV) an associated pair with the male represented as very much smaller than the female
but resembling her in general habitus, therefore hardly a typically cyclopoid form. We hâve seen
some material from late developmental stages in species of this genus which lead us to believe Sars
correctly depicted the male for S. inflatus. We are further informed by Dr. J. Stock, by letter, that
he has found a dimorphism among males of Botryllophilus. It is therefore prématuré to ofler an exten-
ded diagnosis of males for the subfamily.
In the material we hâve seen of males in Botryllophilus (fig. 52) there are some important consi¬
dérations in addition to the general form of the habitus. The antennules are furnished profusely
with aesthetascs, contributing to our impression of similarity between these males and those of some
species of Haplostoma. The mouthparts are much modified (fig. 52 b) with the masticatory lamella
Source : MNHN, Paris
140
PAUL L. ILLG AND PATRICIA L. DUDLEY
Fi«. 52. — Bolryllophiltu ap., Washington, male : a, habitus, latéral ; b, en face vie'vr of anterior cephalosome : Md =
mandible, M = mouthpart protninence, including possible rudiments ot maxillule and maxilla. Scales = 0.1 ratn.
Circies on anlennules indicate insertions of aesthetascs.
of the mandible lacking and the maxillule and maxilla reduced to minute vestiges. The first legs
show a modification of the endopodites, this tending to vary at the spécifie level. With this highly
developed dimorphism, there seems to be a conservatism in the antenna and a correspondence of the
number of ornaments in the 2 sexes. The latter feature has furnished us the only reliable guide so far
for differentiating the male we studied from the swimming males of Haplosloma species. None of
the latter has more than 4 éléments of armature on the distal antennal segment ; the male of Bolryl-
lophiluê has 7. In the males of Sars’ Schizoproetws type there seems to be fair correspondence in the
structure and ornamentation of the first to fourth legs, which also are symmetrical. We hâve found
no asymmetry in the males we hâve studied.
KEY TO GENERA, BASED ON FEMALES
1. Fifth leg displaced distinctly laterally and dorsally, of various shapes, always separated by both
wide ventral and wide dorsal spaces, not forming a distinct enclosure for the ovisac or ovisacs..
Botryllophilus
1.' Fifth leg a lamella, the pair taking origin close together so their dorsal medial margins are closely
adjacent, forming an enclosure or paired enclosures for the ovisac or ovisacs.... Sckizoproctus
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
141
Botryllophilus Hesse, 1864
Botrylbphilus Hesse, 1864, p. 345*348 (type species, B. ruher Hesse, by subséquent désignation). — Bâte,
1864, p. 308. — Hesse, 1865. p. 224 ; 1866, p. 85 ; 1869, p. 345-358. — Gerstaecker, 1870-1871,
p. 719, 774. — Aurivillius, 1886, p. 44-46. — Canu, 1891a, p. 473 ; 1892, p. 29, 33, 35, 66, 108,
203-209 (part). — T. Scott, 1900, p. 388-389 ; 1901a, p. 242-245. — Thompson & A. Scott, 1903,
p. 255. — T. Scott, 1907, p. 367-368. —Brément, 1909, p. lxix-lxx. — Hartmeyer, 1911, p. 1734
(part). — Chatton & Brément, 1915, p. 145. — Sars, 1921, p. 67-68. — Schellenberg, 1921, p. 9 ;
1922, p. 280, 281-287 (part). — Hansen, 1923, p. 25. — Harant, 1931, p. 370. — Wüson, 1932,
p. 392, 600, 602. — Neave, 1939, p. 460. — Lang, 1948, p. 8 (part). — Sewell, 1949, p. 145-146,
170. — Rose & Vaissière, 1953, p. 90. — Barnard, 1955, p. 240. — Gotto, 1960, p. 225. — Bres-
ciani & Lützen, 1962, p. 374. — Dudley, 1966, p. 155, 157, 160. — Gotto, 1960a, p. 193. —
Stock, 1970, p. 16-17.
Kossmechthrus Délia Valle, 1883, p. 248-252 (types species by monotypy, K. notopus Délia Valle, 1883).
— Canu, in Giard, 1888, p. 505.
Kosemcthrus Canu, 1886a, p. 311 ; 1886b, p. 1025. — Neave, 1939, p. 835.
Hlakeanus Wilson, 1921, p. 10-11 (type species, by original désignation, B. corniger Wilson, 1921).
— Hansen, 1923, p. 24-25. — Wilson, 1932, p. 391, 601. — Préfontaine, 1936, p. 76. — Neave,
1939, p. 436. — Sewell, 1949, p. 000.
non Ceratrichodes Hesse, Schellenberg, 1922 (synonym of Botryllophilus), p. 281, (cf. Lang, 1948,.p. 8)
Female :
The body in the adult is inflated, with varying degrees of expression of the ingrédient seg¬
mentation. The cephalosome bears appendages through the maxillipeds and is usually well demar-
catcd. The metasomal segments bear the 5 pairs of legs and exhibit the most inflation. There is
basically a clear-cut articulation between the segment of the fifth legs and the 6rst urosomal segment,
which is demonstrated as being thoracic by the presence midventrally of a considerably developed
apparatus surrounding the insémination pore and the latéral or dorsolateral apertures of the oviduct,
usually also with auxllliary cuticular structures. Rarely the génital segment fuses with the forebody
and there is no well-developed major body articulation. There seems to be a basic number of 4 true
abdominal segments between the segment of the génital apertures and the well developed article bearing
the caudal rami, here termed the anal somite and not considered as a true segment but some kind
of complex including the telson. In many forms abdominal annulation involving distinct inflections
of the cuticule présent the aspect of a multisegmented urosome, up to 8 such éléments being présent
among species examined. The evidence from internai anatomy, particularly with regard to the mus¬
culature, is so far lacking to makc an exact interprétation of this phenomenon. The caudal ramus
usually is set on a complication of the end of the anal somite. The ramus bears terminally 4 stout
curved claw-like éléments and 1 or more additional setae.
The rostrum may or may not be conspicuously developed but is typically présent.
The antennule is typical for the whole genus, of few, usually 4, segments, the basal portion mas¬
sive and the terminal much reduced in diameter. AH segments bear setae. On the basal segment
there is a row of protubérances furnishing support for the major setae and such protubérances also
exist on the more distal segments.
The antenna varies from 2-8egmented, to the appearance of having 3 segments, because of an
articulating région between the basal and terminal segments. The ornamentation, restricted to the
distal article, consists of marginal spines or seta, inserted at intervals, often in emarginations, and a
terminal row of éléments. No form has so far been found with less than 7 of such éléments. Many
of the species we hâve studied hâve exhibited an asymmetry in the appendages, this characterizing
the antennae and the swimming legs 1 to 4. In the antennae the asymmetry is not marked. There
Source ; MNHN, Paris
142
PAUL L. ILLG AND PATRICIA L. DUDLEY
may l)e subtle différences in the contour of the segments, particularly the distalmost. More apparent
is a different degree of development of the éléments of the armature of the distal segment. On one
side the éléments may tend more to a spiniform appearance, on the other they would then be more
setiform. We do not know of a case of different numbers on the 2 sides. This asymmetry is not
expressed în the other head appendages in the material we hâve observed.
The labrum is usually not distinctive. The mandible, maxillules, maxillae and maxillipeds
are not usually specifically distinctive and in general conform to the characteristics described for the
subfamily.
The first to fourth legs, the so-called swimming legs, are asymmetrical in most meml)ers eare-
fully examined. However, the pattern of segmentation of the rami, ranging from 1- to 2*segmentcd,
often is in a different pattern in the various members of the leg sériés. This variability can evidently
be very gréai, even within a rather local population of a single species, as was demonstrated by Lang
(1949), but unfortunately his statements and illustrations are not sufficiently detailed for a clear unders-
tauding of the situation without a re-examination of material from his sources. We agréé wifh Stock
(1970, p. 16) that the variability in these appendages is quite distinctly bound to certain limitations.
We also reject the sweeping synonmyization of practically ail previously described species that was
applied by Lang on the basis of his observations on the asymmetry.
The fith legs are of uniform basic construction throughout the genus, with a degree of develop¬
ment of specifically distinctive patterns of proportion and ornamentation. They hâve so far been
found to be symmetrioal. There are some specifically distinctive patterns of posture and configuration,
these extending to groups of species in some cases.
The génital segment of the female shows the insémination pore midventral, with diverging
internai tubes leading laterally to séminal réceptacles near the oviducal apertures. The latter bcar
a cuticular flap in each, with minor features of ornamentation, Thore are sometimes patterns of
sclerotization in the cuticle of the flaps or in the area around the oviducal apertures.
Males :
The male has been known for a fairly long time, but there are some confusions involved in the
identifications. The first decriptions and figures attributed to a male of the genus were those of
Hesse (1866). Even so rigorous a critic as Schellbnbebg found Hesse’s information recognizable
and considered he accounted for the male of Botryllophilus ruber. Canü (1892, PI. 20) figured a male
he ascribed to Aplostonm brevicauda ; this, as Schellenberg pointed out, is the male of a Botryüo-
philus. Furthermore, Canu presented a figure (1892, pl. 16) which he identified as a male of B. macro¬
pus. Bbesciani & Lützen (1962, p. 374, fig. 2, j) show this to be a young female, and they chose
to assign it to B. ruber. Scott (1901, p. 242, pl. XVII) presented a male identified as B. Pruber and
the observations were corroborated by Bresciani & Lützen (1962, p. 374-376, figs. 2, a-i) to the extent
that one can at least accept Scott’s generic désignation. The dimorphism, as shown by this male,
is very striking. Some statements of Stock (1970, p. 16-17) indicate there are still other patterns
to be described for the males within the genus.
On the basis of the males so far described and what we hâve found the body is distinctly cyclo-
piform, with substantial modifications. The antennule emphasizes a sensory function, bearing a great
number of aesthetascs arranged in a characteristic pattern. The mouthparts are modified, tending
to great réduction, with the masticatory lamella of the mandible lacking and the maxillule and maxilla
scarcely distinguishable as appendages. The first legs show a modification of the endopods, this
tending to vary at the spécifie level. The first 4 legs otherwise seem adequately developed natatory
appendages. A very significant point is that there seems to be a consorvatism in the antenna and
the number, at least of the éléments of ornamentation, appears to show a close correspondencè in the
two sexes, although the shapes and proportions of the segments themselves may vary somewhat,
The males we studied were not asymmetrical.
Because of the variations it is expected that discovery of males of additional species will bring
forth it is not fruitful to proceed further to try to formulate a generalized example for the male of the
genus.
Source : Mt'IHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
SPECIES OF BOTRYLLOPHILUS
Unfortunately Botryüophilus ruber Hesse, the type species of the genus, bas not been fully
described or illustrated from unequivocally determined material. Iii his original description, Hesse
presented figures, but of these only his PI. 12, fig. 1 and 2, which approximately depict the general
habitus of the female, may be said to convey any information. There are no précisé facts on the ana-
tomy of the appendages to be gained from his descriptions or from any of the figures he included. Canu
pointed out that the fifth leg of Hesse’s form was very long, and this can be inferred from the figure
accompanying the original description.
T. Scott, 1901b, figures a copepod he called Botryüophilus ruber Hesse. Saks, 1921, p. 68,
chose to differentiate the animal represented by Scott as differing from B. ruber, and referred to it
as B. sp. Some features were well presented, including some of the appendages of the male. Lang,
1949, presented some figures he attributed to B. ruber, but since he synonymized practically ail des¬
cribed species under the one narae, it is difiicult to use his présentation. Probably what he figures,
as B. ruber, sensu Lang, was the animal described by Saks as B. breuipes, which very possibly difîers
from the true B. ruber, in the light of Stock’s remarks (1970, see below). Lang’s material was from
the same région as Sars’ and from Saks’ typical host, Bolrylloides leachi. Ambiguity is compounded
by the fact that Hesse specified Botryllus sp. as the typical host of B. ruber. Many accounts subsé¬
quent to that of Hesse used the désignation B. ruber, derived from a wide array of different hosts.
To return to Lang’s treatment, his figures are of little practical value, but his statements indeed bear
out his daim that he found anatomical facts which essentially invalidated many previously presented
morphological discriminations.
Bresciam & Lützen, 1962, fig. 2, figured a male and young female of Bolryllophilus ruber sensu
Lang, again from the Swedish West Coast, and the host Botrylloides leachi. They pointed out some
minor disagreement with Scott’s présentation of the male, and indicated they were accepting the
conspecificity with their material. This then leaves open to question whether they actually were
dealing with B. ruber sensu Hesse.
Stock, 1970, p. 15-16, stated he had access to topotypic material of B. ruber, and he compared
varions details with his descriptions of specimens from the West Indies. From his statements one can
infer certain of the anatomical details characterizing B. ruber sensu Hesse, but not sufiiciently to gain
a thorough appréciation of the species. Dr. Stock informed us in correspondence, some time ago,
that he is in process of revisionary studies on European species of Botryllophilus. Until topotypes
of several species hâve been fully described and figured there remains a substantal sector in the genus
which cannot be definitely treated.
Central to the problem of the disposition of the previously described species is the fact that the
genus Botryllophilus is marked almost throughout, in the females, by a peculiar asymmetry of certain
appendages. It is particularly striking in legs 1 to 4, so much so as to imply strongly some sort of
différentiation of function. This condition is typically accompanied by an asymmetry in the antennae,
much less striking, and not very obviously suggestive of a functional diversification.
In the legs the asymmetry is expressed at several levels. It may refer to ail or only some pairs.
It tends to involve both rami, but not necessarily. In the endopodites the expression tends to be in
the degree of coalescence of the ingrédient segments, and, somewhat less coramonly, in the number
of the ornamenting éléments. The exopodites présent a much more complex picture. The asymmetry
may, rarely, be expressed only in the number of segments and ornamenting éléments. More commonly
the expodite is reduced to a single segment to begin with, and on this modification is superimposed
a striking différence in configuration on the two sides, and, also very commonly, by a striking dimor-
phism of the corresponding ornamenting éléments on the two sides. The most extremely modified
exopodite is like nothing seen elsewhere in ascidicole copepods. In the genus Schizoproclus the pheno-
menon has not so far been noted, and it certainly is not présent in 5. infUuus nor, apparently, in Sars’
S. (= Pieropygus) festitus ; this latter fact remains to be corroborated.
Source : MNHN, Paris
144
PAUL L. ILLG AND PATRICIA U DUDLEY
There is no evidence whether Hesse noted the phenomenon, although he described several species
in his genus. Stock (1970, p. 21) refers to material in his possession he considéra as topotypes of B.
ruber, and points out the legs 1-4 are asymmetrical, the condition extending to différences in numbers
of ornaments on most of the rami.
One of the most extended discussions of B. ruber is that of Lang (1949), in which among other
things the problem of asymmetry was discussed and dismissed with the conclusion that essentially
ail forms described in the genus Botryllophüus were referrable to the single species, B. ruber. The
material of Lang was from the Swedish west coast, from Bolryïloides leachi. Since he compared his
material to types of Sabs’ B. brevipes and found a good substantiation of the original description, the
conspecificity seems established. It is not at ail certain that these specimens refer to the Botryllo-
pkilue ruber of Hesse, contrary to Lang’s conclusion. Lang found in most of his specimens and one
of Sabs’ that the legs 1-4 are asymmetrical, but it is very likely that the condition he found is at a
different level from what Stock referred to in his specimens. Sabs depicted the appendages of his
form as very simple with ornaments little modified, except perhaps as to length, and, indeed, in the
direction of some general réduction. Lang considers the différences he found in most legs to amount
to a very marked asymmetry. His drawings and indeed his discussion do not convey a great deal
of information but would lead one to conclude that Lang did not really appreciate the nature of asym¬
metry as it is found in the majority of species. He found that sometimes a weak articulation con-
veyeda 2 articled aspect to a ramus, and he found some variability in the numbers of éléments.
Actually his material would seem to us to represent a species with a very low level of asymmetry. As
he more or less directly pointed out bimself, he was dealing with a rather exceptional order of indivi¬
duel variation. His conclusions therefore hâve little bearing on the problems of asymmetry in the
majority of species. Doubtless his misunderstanding of this basic point was crucial to his completely
unacceptable sweeping conclusion as to the conspecificity of ail previously described material.
The literature on asymmetry is useful to some degree in providing a few dues with regard to
older species. Hesse did not make any allusion and his figures and descriptions are too uninformative
to allow any inference as to whether he was or was not aware of the phenomenon. Della Valle
certainly was aware, and invoked the asymmetry of the legs in his description of B. nolopus. Canu
(1892) knew of the situation from Della Valle’s paper and expressly stated that ail of the legs of his
B. macropus were syrometrical. Such authors as the Scotts and a fairly long list of others of the
epoch did not seem to appreciate the phenomenon.
Bbément (1909) invoked asymmetry and “ resemblance ” in his descriptions of B. brevipes
and B. banyulensis, referring to différences in configuration in the successive pairs of limbs as well as
to differing numbers of segments and ornaments. He understood the phenomenon very well and had
a good grasp of the taxonomie problems in the genus.
ScHELLENBERG (1922) considcred the matter carefully and obviously noted aspects of the con¬
dition in his own material. He carefully described some details and made some spéculations as to
possible underlying influences. We are inclined to agréé he was right in this regard to the extent that
some features in the behavior of the animal in relation to its naturel history as a symbiont may very
well dépend on the well developed morphological différentiation. Lang’s airy dismissal of Schel-
lenberg’s careful observations and clearly tentative conclusions reflects oniy his own lack of précisé
information.
We are firmly of the opinion already stated by Stock, 1970, that Lang went much too far in
synonymizing so many species in B. ruber. There are some morphological features of sufficient relia-
bility to differentiate a considérable of the recorded forms. On the other hand, as we hâve found
throughout the ascidicolids, but here even more pronouncedly, there are some eonsistent morpholo¬
gical plans of suspiciousiy wide géographie dispersion and diversity of hosts. Superimposed in Botryl-
lophilus is a strong tendency to great individual variability of varions characters, particularly thosc
deriving from the thoracic appendages. It will be necessary to study in detail large suites of specimens
and probably, also, to scrutinize closely, in selected strategie localities, the entire complément of species
to place the genus in a sound taxonomie State. Our intensive studies were in a locality (San Juan
Àrchipelago, Washington) fortunate for many features of occurrence of simple ascidians and a great
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
145
variety of ascidicoles, but Jt îs a less than idéal locality for botryllophüins because the fauna of com-
pound ascidians is impoverished and in various regards somewhat anomalous. Further, the occur¬
rence of several botryllophüins in hosts of decided immaturity leads us to suspect that for such spe-
cies we hâve not yel located the most characteristic hosts. We hâve not been able to assign a satis-
factory identification to the most common species available to us because it corresponds morpholo-
gically to the group of forms from the west coast of Europe centered around B. ruber Hesse and “ B.
brevipes ” Sars. We await the resolution of this taxonomie problem before we feel we can présent
our anatomical findings and a description we hâve compiled of a suite of the developmental stages.
The material from Europe at our disposai is insufficient for us to attempt the disposition of a majority
of the species in the genus. We hâve some spécimens which lead us to conclude a number of the spe¬
cies described hâve substantiel validity. We list below the species so far described with our commenta.
Botryllophilus ruber Hesse, 1864
Botryllophilus ruber Hesse, 1864, p. 345-347, pl. 12, fig. 1-12 (type locality Rade de Brest, France, from
Bothyüus (sic) slellatus). — Bâte, 1864, p. 308. — Gerstaecker, 1870-1871, p. 719, 774. — Canu,
1892, p. 205,207. — (?) Thompson & Scott, 1903, p. 255,273. — Norman & Scott, 1906, p. 203. —
Brément, 1909, p. lxxiv, lxxvii, lxviii. — Hartmeyer, 1911, p. 1734. — Schellenberg, 1922,
p. 283, 294. — Harant, 1931, p. 370. — Leigh-Sharpe, 1935, p. 48. — Lang, 1948, p. 3, 9,10,
11-14, 15-16, ? figs. 1-6 (part). — Sewell, 1949, p. 145, 170, 177, 188. — Gotto, 1952, p. 674 ;
1954, p. 666 ; 1960, p. 216, 221 ; 1961, p. 153. — (?) Bresciani & Lützen, 1962, p. 374-376, text-
fig. 2. — Gotto, 1966a, p. 193. — Stock, 1970, p. 21-22, 24. — Hamond, 1973, p. 350.
non B. ruber T. Scott, 1900, p. 388 ; 1901b, p. 242-245, pl. XVII, fig. 15-27 (= B. sp., Sars, 1921, p. 68).
This species requires extended redescription and illustration and estimation of the range of
variation in the morphological characters and host incidence. Stock (1970, p. 21-22) has providéd
a number of details from topotypes but a fuller treatment is needed. With this uncertain base of
comparison it becomes difficult or impossible to allocate a substantial number of the subsequently
described species.
Botryllophilus macropus Canu, 1891
Botryllophilus mucropus Canu, 1891a, p. 473 (type locality, Wimereux, France, from Lithonephria
eugyranda Giard) ; 1892, p. 66, 67, 206-208, pl. 16, fig. 1, 3, 5-14). — Brément, 1909, p. lxvi,
Lxvii, LXVIII. — Chatton & Brément, 1915, p. 145. — Schellenberg, 1922, p. 283,194. — Harant,
1931, p. 370. — Sewell, 1949, p. 145,188. — Stock, 1970, p. 21, 24.
BolryUophilus ruber, Lang, 1948, p. 9, 10,13, 14, 15 (part). — Bresciani & LUtzen, 1962, p. 374 (part).
(?) Botryllophilus canopus Canu (sic), Hartmeyer, 1909-1911, p. 1734.
Canu characterized this species well and there is no difficulty in maintaining its status. It is
interesting in departing from the asymmetrical pattern, and again, perhaps in corrélation, for occurring
in a species of solitary ascidian.
Botryllophilus banyulensis Brément, 1909
Botryllophilus banyulensis Brément, 1909, p. lxxi-lxxviii, fig. IX-XI (type locality, Port-Vendres,
Golfe du Lion, France, from Parascidium areolatum délia Chiaje). — Schellenberg, 1922, p. 282,
293. — Harant, 1931, p. 370. — Sewell, 1949, p. 183.
10
Source ; MNHN, Paris
^46 PAUL L. ILLG AND PATRICIA L. DUDLEY
Botryllophïlus ruber Lang, 1948, p. 9, 10, 14, 15 (part).
The description and figures by Brément are excellent ; it would appear to be a valid species.
Botryllophilus brevipes Brément, 1909
Botryllophilus brevipes Brément, 1909, p. lxx-lxxii, fig. VI-VIII (type locality, Port-Vendres, Golfe
du Lion, France, from Amaroucium lacteum D'-asche). —• Hartmeyer, 1911, p. 1736. — Schellen-
berg, 1922, p. 282. — Harant, 1931, p. 370. — Sewell, 1949, p. 183.
Botryllophilus ruber, Lang, 1948, p. 9, 10, 14, 15 (part).
The description and figures by Brément are excellent ; we consider it a valid species.
Botryllophilus bergensis Schellenberg, 1921
Botryllophilus bergensis Schellenberg, 1921, p. 9-11, fig. 8-9b (type locality Bergen, Norway, from
Leploclinides faerôensis (Bjerkan)) ; 1922, p. 282, 293.
Botryllophilus ruber, Lang, 1948, p. 9, 10, 14, 15 (part).
The asymmetrical thoracic appendages depicted by the author indicate some correspondence
with statements of Stock about B. ruber. Fuller description of both forms will be necessary to esta-
blish if this species can be maintained as a valid one.
Botryllophilus norvegicus Schellenberg, 1921
Botryllophilus norvegicus Schellenberg, 1921, p. 6-9, figs. 5-7 (type locality, Trondhjemfjord, from
Pelonaia corrugaia Goodsir) ; 1922, p. 282, 294. — Harant, 1931, p. 370. — Lang, 1948, p. 9,
10,13,16. — Gotto, 1960, p. 225.
Blakeanus corniger Wilson, 1921, p. 11-13, pl. 5, fig. 39-43 (type locality. Long Island Sound, U.S.A.,
from Cynthia carnea Verrill). — Wilson, 1932, p. 391, fig. 240.
Blakeanus groenlandieus Hansen, 1923, p. 24-25, pl. II, fig. 8a-8d, pl. lîl, fig. la-lh (type locality»
West Groenland, from Cynthia rustica L.). — Préfontaine, 1936, p. 76. — Sewell, 1949, p. 194.
— Préfontaine & Brunei, 1962, p. 253.
This striking species is valid. We bave a number of collections indicating it is rather wides-
pread in northern waters. It is a very large form, perhaps to some degree in corrélation with its occur¬
rence in the branchial cavity of solitary ascidians. A degree of asymmetry is persistent, although
not as extreme as in some of the small species.
Botryllophilus brevipes Sars, 1921
Botryllophilus brevipes Sars, 1921, p. 68-69, pl. 33 (type locality, Espevaer, Norway, from Botryllus sp.).
— (?) Wilson, 1932, p. 392, fig. 241. — SeweU, 1949, p. 145, 157, 158, 192.
Botryllophilus ruber, Lang, 1948, p. 8, 9, 10,11, 12, 13, 14, 15 (? fig. 1-6) (part). —Bresciani & Lützen,
1962, p. 374-376, fig. 2.
As was usual with Sars this form is very well illustrated and his présentation seems well substan-
tiated by the subséquent reports of Lang and of Bresciani & Lützen. Lang’s synonyray of this
form with B. ruber does not seem to us justifiable, but the question must remain open until B. ruber
is better known. Brément’s spécifie name has precedence over that of Sars. It seem to us unlikely
that the species of the two workers are conspecific. It may well be that B. bergensis Schellenberg
may turn out to be a synonym, in which case his name, as older, would take precedence.
Source : MNHN, Paris
ASdDICOLIDAE AND SUBFAMILIES
147
Botryllophilus aspinosus Schellenberg, 1922
Botryllopkilus aspinosus Schellenberg, 1922, p. 282-284, 293, fig, 1-2 (type locality, Plymouth, from
Polycarpa pomaria (Savigny) ; Angola, from Styela kupferi Michaelsen). — Harant, 1931, p. 370.
— Sewell, 1949, p. 188. — Barnard, 1955, p. 240-241, fig. 7, 9.
Botryllopkilus ruber, Lang, 1948, p. 9, 10, 14, 15 (part).
This appears to us to be a valid species, wel! characterized for the period at which it was des-
cribed. Redescription on a more detailed basis is now necessary to allow for treatment of the genus.
Botryllophilus inaequipes Hansen, 1923
Botryllopkilus inaequipes Hansen, 1923, p. 25-26, pl. III, fig. 2a-2f (type locality. Davis Strait, West
Groenland, bottom sample). — Sewell, 1949, p. 194.
Botryllopkilus ruber, Lang, 1948, p. 9, 14, 15 (part).
This species is probably not B. ruber Hesse, so it becomes another possibility in the disposition
of Sars’ b. brevipes. A much more detailed description of the Greenland form and détermination
of its normal host will be required.
Botryllophilus bandalli Stock, 1970
Botryllophilus randalli Stock, 1970, p. 17-22, fig. 14-18 (type locality, Puerto Rico, from Eudistoma
olivaceum (Van Name).
Stock, well aware of the problems in the genus, referred to topotypic specimens of B. ruber
in establishing the substantial différentiation of his valid species.
INDETERMINABLE SPECIES
The remaining species in the genus are indéterminable by us with présent information. It is
possible that fuH redescriptions of some of the species from remoter localities, such as those of Schel¬
lenberg and Sewell, wiU validate their species. Della Valle’s form possibly could be mainteined
with fuller information. The species described by Hesse seem to offer no possibility for constructive
treatment except for B. ruber. The other names should be dealt with by the protocol for disposing
of forgotten names (International Code of Zoological Nomenclature, 1961) and suppressed as far as
future nomenclatural considérations are concerned.
Botryllophilus virescens Hesse, 1864
Botryllophilus virescens Hesse, 1864, p. 347-348 (type locality, côtes de France [rade de Brest], host
not specified). — Bâte, 1864, p. 308. — Gerstaecker, 1870-1871, p. 774. — Canu, 1892, p. 207.
— Hartmeyer, 1911, p. 1734. — Sewell, 1949, p. 145.
Botryllophilus ruber, Lang, 1948, p. 9, 14, 15 (part).
Source ; MNHN, Paris
148 PAUL L. ILLC. AND PATRICIA L. DUDLlîY
?Bolry(lophüus vert, nobis, ? B. viridis, nobis, Hesse, 1865, p. 223-224 ; 1866, p. 79-80, pl. 4, fig. E,
E1-E7. — B. i'iridîs Gerstaecker, 1870-1871, p. 774. — Hartmeyer, 1911, p. 1734.
Botryllophilus pallious Hesse, 1865
Botryllophiluê palUdus Hesse, 1865, p. 224-225 (type locality. Rade de Brest, France, in “ Botrylle ”,
attached to Fucus serratus). — Gerstaecker, 1870-1871, p. 774. — Hartmeyer, 1911, p. 1734.
Bolryllophilus ruber, Lang, 1948, p. 9, 14, 15 (part).
Botryllophilus armatus Hesse, 1869
Bolryllophilus armatus Hesse, 1869, p. 293-294 (type locality, “ côtes de France ”, în “ Botrylle ” on
frond of Fucus vesiculosus). — Gerstaecker, 1870-1871, p. 774. — Hartmeyer, 1911, p. 1734.
Botryllophilus brevis Hesse, 1869
Botryllophilus brevis Hesse, 1869, p. 295 (type locality, from “ Botrylle ” on a Peclen). — Gerstaec¬
ker, 1870-1871, p. 774. — Hartmeyer, 1911, p. 1734.
Botryllophilus pubpurescens Hesse, 1869
Bolryllophilus purpurescens Hesse, 1869, p. 294-295 (type locality, rade de Brest, France, from a
“ Botrylle ”, on an annelid tube). — Gerstaecker, 1870-1871, p. 774. — Hartmeyer, 1911, p. 1734.
Botryllophilus propinquus Hesse, 1872
Botryllophilus propinquus Hesse, 1872, p. 25-27 (type locality. Rade de Brest, France, from compound
tunicate attached to Pecten maximus). — Hartmeyer, 1911, p. 1736.
Botryllophilus notopus (Délia Valle, 1883)
Kossmechthrus notopus Délia Valle, 1883, p. 248-252, 253, fig. 11-23 (type locality, Naples, from Dis-
toma panceri Délia Valle).
Kossmechthrus notopus, Canu, 1886a, p. 311 ; 1886b, p. 1025.
Botryllophilus notopus, Canu, 1892, p. 207. — Brément, 1909, p. 74, 78. — Hartmeyer, 1911, p. 1735,
1736. — Schellenberg, 1922, p. 282, 294. — Harant, 1931, p. 370. — Sewell, 1949, p. 145, 183.
Botryllophilus ruber, Lang, 1949, p. 9, 10, 14, 15 (part).
Botryllophilus africanus Schellenberg, 1922
Botryllophilus africanus Schellenberg, 1922, p. 284-285, 293, fig. 3-4 (type locality, Angola, from Afocro-
clinum angolanum Michaelsen). — Barnard, 1955, p. 240. — Stock, 1970, p. 21, 24.
Botryllophilus ruber, Lang, 1948, p. 9, 10, 14, 15 (part).
Source : MNHN, Paris
ASCIDICOLIDAIÎ AND SUBPAMIIJES
149
Botbyllophilus indicus Sewell, 1949
Botryllopkilus indicii^ Sewell, 1949, p. 146 (type locality, Nankauri Harboiir, Nicobar Islands, weed
washings). — Stock, 1970, p. 22, 24.
There seems a strong pos-sibility that this species can be maintaincd, but a much more detailed
description and clearer illustrations of the appendages are needed.
Botbyllophilus sp. Brément, 1909
Botryllophilus sp. Brément, 1909, p. lxxviii. — Schellenberg, 1922, p. 294. — Lang, 1948, p. 16.
Botbyllophilus sp. Sars, 1921
Botryllophilus sp. Sars, 1921, p. 68 for.
Botryllophilus ruber, T. Scott, 1900, p. 388 (Loch Fyne, Scotland, from débris) ; 1901b, p. 242*245,
352, pl. XVIII, fig. 15-27. — Lang, 1948, p. 16 (part). — Sewell, 1949, p. 145.
Botryllophilus sp. Schellenberg, 1922
Botryllophilus sp. Schellenberg, 1922, p. 294 (from Plettenberg Bucht, S. Africa, from Polycitor renieri
Hartmeyer). — Lang, 1948, p. 16.
Botryllophilus sp. Schellenberg, 1922
Botryllophilus macropus (“ petits ”) Canii, 1892, p. 206, pl. XVI, fig. 4, 5 (Boulonnais, English Channel,
from Polyclinum luteum Giard).
Botryllophilus sp. Schellenberg, 1922, p. 294.
Botryllophilus ruber, Lang, 1948, p. 13, 15 (part). — Bresciani & Lütsen, 1962, p. 374 (part).
Botryllophilus sp. Dudley, 1966
Botryllophilus sp. Dudley, 1966, p. 157, 160 (Friday Harbor, Washington, from Aplidium sp.).
Botryllophilus sp. Stock, 1970
Botryllophilus sp. Stock, 1970, p. 22-24, fig. 19-21 (from Puerto Rico, in Didemnum sp.).
ScHizoPBOCTus .\urivillius, 1885
Schizoproctus Aurivillios, 1885a, p. 247-248 (type species, by monotypy, S. inflalus Aurivillius, 1885a) ;
1886, p. 44-46. — Sars, 1921, p. 71-72. — Hansen, 1923, p. 26. — Neave, 1940, p. 137. — Wilson,
1932, p. 600. — Lang, 1948, p. 8. — Gotto, 1960, p. 225. — MacGinitie, 1955, p. 114, 150. —
MacGinitie & MacGinitie, 1968, p. 48.
Source : MhtHN, Paris
150 PAUL L. ILLG AND PATRICIA L. DUDLEY
Botryllopkibis {part) Canu, 1892, p. 203. — Brément, 1909, p. 75, 76, 78. — Hartmeyer, 1911, p. 1734.
— Schellenberg, 1922, p. 281. — Harant, 1931, p. 370.
Pteropygus Sars, 1921, p. 70 (type species, by monotypy, P. veslUus Sars, 1921). — Wilson, p. 600.
— Xeave, 1940, p. 1029. — Sewell, 1949, p. 191. — Gotto, 1960, p. 225.
Sckizoproclus is a well established genus, readily rccognizable since its establishment. It shows
a high degree ot correspondence to Boiryllophilus and the diflerences between the 2 généra are those
of degree rather than of kind. A number of perceptive workers did not hestiate to synonymize the
2 généra. Sars maintained the séparation and wcnt so far as to establish a new genus, Pleropygits,
for a related form he found in Swedeii. We feol that maintainiiig a third genus in this situation is
not supportable. Further, adding P. veslilus Sars to Sckizoproclus rc-enforces the generic concept
and also emphasizes the différentiation between the 2 sets of species. It is of interest that botli the
species we assign to Sckizoproclus corne from solitary ascidians, from northern localities, and do not
show the asymmetry encountered in so many species of Boiryllophilus. They are also among the giants
of the ascidicole copepods. The species are readily differentiated, following characters presented by
Sars.
KEY TO THE SPECIES
1. Urosome including 5 articles posterior to the génital complex. inflatus, Aurivillius
1'. Urosome including 3 articles posterior to the génital complex. vestitus, Sars
ScHizopRocTvs INFLATUS Aupivillius, 1885
Sckizoproclus inflcUus Aurivillius, 1885a, p. 248-250, pl. 9, fig. 21-32 (type locality, Hackluyts Heade,
Spitzbergen, from Pkallusia sp.) ; 1886, p. 44-40, pl. 1, fig. 7. — Hartmeyer, 1911, p. 1735. —
Sars, 4921, p. 72-73, pl. xxxiv. — Hansen, 1923, p. 26-27. — Stephensen, 1932, p. 3-4. — Lang,
1948, p. 8. — Sewell, 1949, p. 192, 193, 194. — Gotto, 1960, p. 225. — MacGinitie, 1955, p. 114,
150. — MacGinitie & MacGinitie, 1968, p. 450.
Boiryllophilus inflatus, Canu, 1892, p. 207-208. — Brément, 1909, p. 75, 76, 78. — Schellenberg, 1922,
p. 283, 293. — Harant, 1931, p. 370.
Doropygus gibber, Vanhôffen, 1897, p. 292.
Distribution : Arctic Océan, Atlantic : Greenland, Norway.
Hosts : Pkallusia sp., Ascidia dijmphniana (Traustedt), A. obliqua .\lder, Boltenia ovifera (Linnaeus),
Molgula groenlandica.
This species was described from the Arctic Occan and subséquent records show it ranges faîrly
widely there. Sars’ record added it to the fauna of Norway and we hâve other evidence of its occur¬
rence in the northern Atlantic fauna.
ScHizopRocTus VESTITUS (Sars, 1921), New Combination
Pleropygus festilus Sars, 1921, p. 70-71, pl. xxxiii (type locality, Risôr, Norway, from Pkallusia obli¬
qua). — Sewell, 1949, p. 191. — Gotto, 1960, p. 225.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIliS
151
INDETERMINABLE GENÜS, BOTRYLLOPHILINAE
Parabotryllovhilus Leigh'Sharpe, 1934
Paraioln/llophilus Leigh-Sharpe, 1934, p. 216 (type species, by monotypy, P. arrisio Leigh-Sharpe,
1934).
The animal as described cannot be assigned to the Botryllophilidae, although placed there by
the author. We know of no other place in the classification to receive it. The description seems to
ns to be a chimaera of some sort ; perhaps drawings ot 2 organisme were somehow combined. We
find the présentation insufiiciently detailed to place the animal anywhere in the Ascidicolidae.
Entebognatiiinae, New Subfamily
Ascidicolidae, Giesbrecht, 1901, p. 77 (part). — Caïman, 1908, p. 172, 182 (part) ; 1909, p. 103 (part).
— Stebbing, 1910, p. 550 (part). — Gravier, 1912b, p. 70 (part) ; 1913, p. 66 (part). — Chattou
& Brément, 1915, p. 143 (part). — Brehm, 1927, p. 490 (part). — Stock, 1959, p. 74 (part) ;
1966, p. 211 (part).
Ascidicolinae, Schellenberg, 1922, p. 227-230 (part).
Enterocolinae, Chatton & Harant, 19241), p. 361 (part).
Female :
The Lody is inflated and the shapes suggest those found among the haplostomins and enterocolins,
with prominent pediform projections corresponding to the fifth legs and tending to enclose the anterior
ends of the egg sacs. The segmentation of the body varies from very clear eut to much reduced, with
only indications by distributions of the appendages and a number of vague cuticular inflections.
The antennules and antenna conform in a basic way to the ascidicolid generalized type but
with the modifications deriving essentially from extrême réduction in segmentation and ornamentation.
There is little or no représentation of a rostrum. There is a mouth cône apparently representing the
labrum and labium. Essentially there is a mandible wth well-developed masticatory lamella (Ente-
rognalhus), this appendage possibly graduating to obsolescence or disappearance (Zanclopus).
The maxillule and maxilla vary from retrgressed but recoignizable ascidicolid dérivation [Ente-
rognathus) to a single pair, much reduced {Zanclopus). Maxillipeds are probably absent. The single
pair of conspicuous mouthparts of Zanclopus are entirely problematicai. There is no evidence for
determining exactly their identity. They could as well represent maxillae or maxillipeds, but in any
case are much modified by réduction.
The legs of pairs 1 to 4 are much modified, but show ascidicolid features. They are biramous
but with much suppression of segmentation. Intercoxal plates are not developed. The usual latéral
setae of the coxopodites are présent (but not so in one of the species of Zanclopus, and in Enterognatkus
the médial spine of the first basipodite is présent). The exopodites terminate in falciform hook-pro-
eesses, not uniike some seen elsewhere among ascidicolids. The much modified endopodites, orna-
mented with processes which somehow probably represent some of the usual éléments of armature,
are distinctive, but not exceptional when considered in relation to the famîly as a whole.
The pediform processes involving the fifth legs are expanded curved lamellae such as are found
in many ascidicolids. In the case of Enterognathus the ornamentation is unique, consisting of 4 well-
formed setae. The insémination apparatus of the segment next posterior is comparable to that found
throughout the ascidicolids.
Source : MNHN, Paris
152
PAUL L. UXG AND PATRICIA L. DUDLEY
The caudal rarai are not greatly distinctive ; they are somewhat modiiied in having a reduced
armature — but there are perhaps 4 éléments represented in the complément of Zanclopus cephalodisci.
Male :
The males are rather generalized cyclopiform in appearance, but with specializations. As in
many of the ascidicolids the antennules are more developed than in the females, with 6 to 8 segments
présent. There are fairly numerous setae, but no distinctive feature among them and no notably
salient development of aesthetascs. In both Enterognathus and Zanclopus this appendage could be
claimed as offering strong support for inclusion of the généra in the Âscidicolidae. The antenna is
3-segmented and suggests that of some of the more dérivative ascidicolids, especially that of Entera-
cola. There is a well-developed seta on the basal segment in Enterognathus, also a feature of Entera-
cola. The second segment is unornamented. The third beats terminal éléments, somewhat between
spiniform and setiform. In Zanclopus cephalodisci there is an apical trio of distinctly curved strong
setae. The presence of a curved or clawed élément apically on the antenna is rare in ascidicolids and
a very strong character in the Notodelphyidae. The persistance in Enterocola and Enterognathus
could quite possibly be regarded as a conservative character.
The mouthparts are reduced to lacking, totally so in Enterogrxathus. There remains Calhan’s so
far unverified représentation of a pair of mandibles in Zanclopus cephalodisci and a pair of protubérances
more or less in the position of the more posterior mouthparts. In any case the diagnostic fact for
the males of the subfamily is a very great réduction of mouthparts, a feature widespread in the family.
Legs 1 to 4 are biramous with the rami trimerous. The ornamentation as depicted so far shows
some slight variation from a typical cyclopoid pattern but there does not appear to be any diagnostic feature
deriving from this which could be presented with any comfortable degree of reliability.
Fifth and sixth legs conform well enough to a general characterization for ascidicolids and do
not offer distinctive features. The fifth legs are bimerous basically, each article set with at least one
prominent seta. There are supplementary spiniform processes of the distal article in Enterognathus,
these replacing setiform éléments of earlier developmental stages. If indeed, 3 setae are represented
this would be a rather primitive expression. The sixth legs of Enterognathus seem to hâve 3 setae,
an exceptionally high number.
The caudal rami are again very generalized, with 6 éléments of armature, 4 of these terminal,
as in typically cyclopoid maies.
Varions observera pointed out strong similarities of Enterognathus with the Enterocolinae, but
the conformity is contradicted by the presence of the substantiel mandibular lamella and vestigial
palp in Enterognathus. Enterocolinae lack the mandible. Zanclopus is in a more ambiguous position
because the nature of the mouthparts is not apparent, but it should in any case accompany Entero-
gruUhus in the classification.
KEY TO GENERA (BASED ON FEMALES)
1. Body with segmentation distinctly apparent ; mouthparts for the most part well-developed, with
some apparent setiferous lobes. Enterognathus Giesbrecht
1'. Body inflated, with segmentation not strongly demarcated ; mouthparts much reduced, no seti¬
ferous lobes apparent. Zanclopus Caïman
Enterognathus Giesbrecht, 1900
Enterognathus Giesbrecht, 1900, p. 61, 76-78 (type species, by monotypy, E. comatulae Giesbrecht,
1900). — Caïman, 1908, p. 171, 182 ; 1909, p. 123. — Gravier, 1912b, p. 70 ; 1912c, p. 244, 245 ;
Source ; MhiHhl, Paris
ASCIDICOLIDAE AND SUBFAHILIES
153
1913, p. 59, 70-74. — Farran, 1913, p. 19. — Chatton & Brément, 1915, p. 143, 145. — Schellen-
berg, 1922, p. 295. — Chatton & Harant, 1924b, p. 361. — Brehm, 1927. — Wilson, 1932, p. 596
— Neave, 1939, p. 241. — Grainger, 1950, p. 636. — Changeux & Delamare-Deboutteville,
1956, p. 106-107, 155. — Stock, 1959, p. 74 ; 1966, p. 211, 215-216.
The inflated body and the salient appendages of this form strongly evoke the concept of an
ascidicolid. The réduction of the thoracic legs is very similar to the pattern in enterocolins. The
antennule has few segments, but is setiferous. The antenna is trimerous and rather ascidicolid in one
species, more reduced in the other. There are lahrum, labium and paragnaths. The mandible is reduced
but maintains a well-developed masticatory lamclla. There is a réduction in the remaining mouth-
parts but the hindermost has a somewhat prehensile aspect, with an articulated terminal spiniform
piece set on a massive basal segment. The 2 species so far known can be readily separated by a number
of salient charactors.
1. Antenna 3-segmented ; pediform processes involving the fifth legs set with 4 setae on the distal
margin . comatulae Giesbrecht
1'. Antenna 2-segmented ; pediform processes involving the fifth legs set with 3 setae on the distal
margin . laleripes Stock
Entekocnathus comatulae Giesbrecht, 1900
(Figures 53, a-c)
Figured, but not described : Mcintosh, 1866, p. 611, fig. 5 ; 1875, p. 140.
Enlerognathiis comatulae Giesbrecht, 1900, p. 62-73, pl. 5, fig. 1-33 (type locality, Gulf of Naples, Italy,
from Antedon rosaceus). — Caïman, 1908, p. 177. — Gravier, 1912b, p. 70 ; 1912c, p. 243 ; 1913,
p. 70, 74. — Farran, 1913, p. 19. —Chatton & Brément, 1915p. 145,. — Scheilenberg, 1922, p. 295.
— Grainger, 1950, p. 636. — Changeux & Delamare-Deboutteville, 1956, p. 155. — Stock,
1959, p. 74 ; 1966, p. 215, 216.
Distribution : Mediterranean, British Isles.
Hosts : Anledon bifida (Pennant), A. rosacea (Linck), A. medilerranea Laniarck.
We hâve reviewed specimens and the bterature on the species of EnJerognatkus. The first
described species, E. comatulae was so well illustrated by Giesbrecht and some later workers that
we are not presenting further figures. The appendages conform without difilculty to the range of
variation found within the ascidicolids, but with indications of dérivation from a very primitive condi¬
tion for some of the characters. As is usual there is great dimorphism and the charaeters of the female
are much more distinctive than those of the male, which is a very generalized gnathostome type. Both
male and female lack the eye. The head is perhaps somewhat produced between the antennular bases,
but there is not really a rostrum. The labrum is a prominence with a posteriorly directed rounded
lobe, but without ornamentation. There is a labium-like structure, called by Giesbrecht the lower
lip, and on the sides, at the bases of the mandibles are prominent paragnaths. These latter are well
known in the notodelphyids ; we hâve now seen a very similar type in an archintodelphyid species ;
and above we hâve recorded the presence of paragnaths in Ascidicola rosea. They seem to be lacking
in most of the aseidicolid subfamilies, although it is diffieult to déterminé just how they might be repre-
sented in the very modified and eomplex mouth apparatus of most forms. In possession of the para¬
gnaths and some other features we would say the mouth région of Enlerognalhus shows some genera¬
lized characters, these tending, however, to modification in the direction of régression.
In the female the antennules are very reduced, but without other specialization, and give little
Source : MNHN, Paris
Fio. 53. — Enlerognathut comatulae Giesbrecht, nauplius (a-c) ; Zanclopus ctphalodiaci Caïman, female (d-o) ; male
(p-r) : a, antennule ; b, antenna ; c, mandible ; d, habitua, dorsal ; e, urosome ; f, head, en jace, showing cepbalic
atructurea and appendages ; g, head, en face, anotber apecimen ; h, antennule ; i, antenna ; j, antenna o! opposite
aide ; k, firat leg ; 1, second Icg ; m, third leg ; n, fourth leg ; o, urosome and pediform projection ; p, antennule ;
q, antenna ; r, head and mouth area en face.
Source : MtJHN, Paris
ASCIDICOLIDAE AND SUBFAMIHES
155
information as to the systematic position. They could well be referred to an early copepodid stage
of varions of the ascidicoles we hâve studied. The antennae are also much simpHfied but could also
be derivable from a generalized developmental form. They are much like those seen in the male of
Enlerocola species. The mandibles furnish the strongest grounds for our disposition of the genus in
the Ascidicolidae. The strong coxa with very effective masticatory lamclla is of Lasic gnathostome
type. Paradoxically, the palp is reduced to a minuscule projection, probably incorporating a single
reduced seta. This aspect might readily be derived from a stage like that seen in Ascidicola, and is
close to the condition of Buprorus. The position of the latter in our scheme is somewhat ambiguous,
but basing our argument on the condition of the mandible, we might argue that both the Buprorus
line and the Enterognathus line represent very early dérivative offshoots of a basic ascidicolid stock,
the daughter sériés taking rather indépendant evolutionary pathways of morphological adaptation.
The maxillule is bilobed, as in some ascidicolids, with a médial portion, probaÛy the endite, bearing
a few setae, and with the palp reduced to an unscgmented setiferous knob. The bimerous maxilla
is much specialized, and certainly reduced, but retains a strong indication of a very basic structural
pattern among the ascidicoles. The massive basal segment bears a single setiferous process which
corresponds well to one of the endites of a more developed maxilla. The terminal piece doubtless repre-
sents a coalescence of distal éléments and retains the strong medially directed claw process of the pri¬
mitive third segment (as in Nolodelpkyopsis) or the usual second segment of somewhat modified forms
{Nolodetphys, many other notodelphyids. The condition is strikingly similar to that in Ascidicola.
The lack of the maxilliped is a character of two other ascidicolid subfamilies, the Enterocolinae and
the Enteropsinae. In both of these, however, the mandible is also lacking in the adult. The obser¬
vation of Stock, 1966, p. 215, fig. 1, h) that the pair of appendages we are calling maxillae are United
by a connecting lamella in his species E. lateripes, injects a somewhat perplexing note in attemps to
identify the mouthparts. From ail past expériences with copepod anatomy, as Stock points out,
a pair of mouthparts so constructed should be the maxillipeds. We lack the information from deve¬
lopment necessary to décidé the exact position, and therefore, the identity of these appendages in the
mouthpart sériés. Stock elected to leave the resolution for future investigation and called the appen-
dage the posterior mouthpart. In the subfamilies which lack the maxilliped, the maxilla is built with
very convincing resemblance to this mouthpart of Enterognathus. Maxillipeds of somewhat similar
aspect occur in the Haplostominae and Botryllophilinae, but the resemblance would be rather far-
fetched. We continue the tradition of the older observera but acknowledge the weight of Stock’s
observation. As to the utility of the appendage as a taxonomie character we would say it serves very
well in either case to place Enterognathus in the ascidicolid sériés. The potentiel for modification of
either the maxilla or the maxilliped to evolve into such a structure seems to us well demonstrated by
the anatomies now known. The thoracic legs show many éléments seen among ascidicolids, but the
exact pattern is unique. The remarkable exopodites hâve a striking resemblance to those of Styeli’
cola ligkti, this aspect enhanced by the similar development of a very large seta adjacent on the latéral
margin of the basipodite. The ramus of S. lighti has a much more complex articulation and doubtless
functions in a very different manner. The endopodites suggest very much the condition found in
some enterocolins. The so-called fifth leg probably incorporâtes a portion of the body proper as is
the case in the enterocolins. There is no other instance of similar ornamentation among the other
ascidicolids. However, the evidence from development of the male, as presented by Giesbrecht,
would seem to indicate that early stages show a fifth leg very comparable to that seen in other asci¬
dicoles ; in the female, however, the arrangement is unique from an early stage. There is no appea-
rance of a basa! seta, so usual among the ascidicoles, and occurring here in the male, and the free élé¬
ment is ornamented with 4 setae, an unusual, and probably very primitive feature. The insémination
pore, canals and associated apparatus of the female are very much as in the majority of the ascidi¬
colids. The habitus and the appendages of the male which are présent correspond very well to the
structural characteristics of generalized ascidicolids without any difficulty. There are no mouthparts.
The plans of the fifth and sixth legs conform very well to those of a number of ascidicolids.
It would be very useful to hâve a full account of the development of this form. At Naples,
Dudley was able to make some observations on nauplii and obtained some copepodids, but there
Source : MNHN, Paris
156
PAUL L. ILLG AND PATRICIA L. DUDLEY
was iiisuflicient material for a thorougli study. Giesbrecht presented some information from laie
copepodids. Our nauplJus showed many primitive characteristics, and as in inost ascidicoles the
development was lecithotrophic. In the first nauplius the larva was of good size (0.264 mm in length)
and the features of the appendages were very suggestive of those of a primitive notodelphyid, such
as Notodelphys. The general impression would be of a somewhat more generalized nauplius than has
so far been seen in ascidicolids. However, such observations as we hâve indicate a very fast passage
through the naupliar sequence, and very probably an abbreviated number of naupliar stages, perhaps
about 4. The nauplius contains a substantial amount of yellow pigment, somc of it associated with
the yolk, and lacks an eye. The uniramous antennule (fig. 53, a), very much like that of Notodelphys,
is trimerous, uniramous. The basal article lacks armature, but there is a short ventral row of fine
spinules on the distal margiii. The second article has a distal ventral plumose seta of moderate length,
two small thorn-like spines inserted on the ventral margin at about equal intervals, and 3 short rows
of fine spinules on the surface. The distal article is the longest, substantially longer than widc. It
hears 2 apical setae, very unequal in length. The major seta, about 3 times as long as the other, bears
a short projection near its base, which probably is an aesthete. Both setae are plumose.
The antenna (fig. 53, b) is biramous, with the segmentation obscure. The protopodite is possi*
bly bimerous, with an articulating basal élément, which however may only represent a protrusion
of the surface of the head. The major article of the protopodite is J*shaped, so that although the
uniinerous endopodite forms a sharp articulation, the positions of the setae indicate that the basal
article of the exopodite is directly continuons with the longer arm of the protopodite. There are orna-
menting rows of spinules and 2 short thorn-like spines equispaced on the médial margin. The some¬
what ovoid endopodite bears a short seta at about midway on the médial margin and 2 subequal long
plumose setae at the apex, which is constricted to a rounded contour. There are 3 clearly set off
exopodite articles, each bearing a long plumose seta at the mediodistal corner, as does also the fused
basal article. The distal article bears 3 setae, a proximal médial one very slender and rather short,
and a long terminal seta accompanied by a much shorter rather stout latéral seta. Ail these setae
are plumose.
The mandible (fig. 53, c) has somewhat the appearance of the antenna, with J-shaped proto-
podite, this also with a possibly basal article. The distal article of the protopodite bears 2 short thorn-
like setules equispaced on the médial margin, with a short row of spinules at the base of the distal one.
The endopodite is bimerous, the distal article much the longer, although the width is uniform throu.
ghout the ramus. On the basal article there is a short spine at about midway on the médial margin-
There is a short row of fine spines on the surface just distal to this. The distal article bears a short
setule, without ornament, on the médial margin well distal to the midpoint, and 2 subequal apical
plumose long setae. The exopodite is probably tetramous, as indicated by the setation, but there is
no articulation between the basal portion and the exopodite. At the distal médial corner of the basal
article portion there is a long plumose seta, corresponding to 3 more équivalent setae, progressively
slightly shortened, each inserted distomedially on its article. Accompanying the seta of the terminal
article there is a very short acute apical setule.
Reference to our studies of Enlerocola and Enteropsis hâve demonstrated many points of simila-
rity with the nauplius of Enterognathus and with very few substantial différences, if any.
Enterognathus lateripes Stock, 1966
Enterognathus lateripes Stock, 1966, p. 211-216, fig. 1-2 (from Eilat, Guif of Aqaba, Red Sea, from
Helerometra saoignyi (J. Muller) ; Oligometra serripinna (P. H. Carpenter) ; Decametra chedwicki
A. H. Clark.
Distribution : Red Sea.
Hosts : Helerometra savignyi (J. H. Miiller), Oligometra serripinna (P. H. Carpenter), Decametra ched¬
wicki A.. H. Clark.
Source ; MNHN, Paris
ASCIDICOI.IDAE AND SL'BFAMILIliS
157
Zanclopus Caïman, 1908
Zuuclopus Caïman, 1908, p. 178, 182 (types species, by monotypv, Z. cephalodUci Caïman, 1908). —
Stebbing, 1910, p. 550. — Gravier, 1912a, p. 1440 ; 1912b, p. 243 ; 1913, p. 68, 69, 73. — Chat-
ton & Brément, 1915, p. 143, 145. — Schclienberg, 1922, p. 295. — Chatton & Harant, 1924b,
p. 361. — Brehm, 1927, p. 490. — Wilson, 1932, p. 601. — Neave, 1940, p. 691. — Barnard,
1955, p. 237, 242.
Zunelopus (lapsus) Gravier, 1913, p. 74.
This interesting copepod, an internai parasite of Cephtdodiscus, was assigned by the author
to the .4scidicolidae and he emphasized the resemblance to ErUerognathus. The genus is readily difîe-
rontiated because of the great réduction of mouthparts, none of them being positively identifiable.
Calman claimed to find a rudimentary mandible terminating in a scythe-shaped blade. The general
aspect of the animal and the thoracic appendages are more or less ascidicolid. We will discuss more
in detail below the observations we hâve been able to make on Calman’s types.
The 2 species so far described are similar to each other, confirming the generic diagnosis of Cal¬
man. They are readily separated, as Gravier, the describer of the second species points out, by
substantial différences in the oral appendages and slighter différences in the ornamentation of the tho¬
racic legs. Gravier did not find identifiable mandibles in his species. The position of the genus
therefore becomes ambigiious. Collection of fresh material and more detailed dissection and examina-
tion of the appendages will be necessary before the genus can be firmly assigned as to taxonomie position.
Zanclopus cephalodisci Calman, 1908
(Figures 53, d-r)
Zanclopus cephalodisci Calman, 1908, p. 178-182, pis. 18,19 (type locality, .4gulhas Bank, ofî Cape of Good
Hope South Africa, from Cephalodiscus gilchristi Ridewood). — Stebbing, 1910, p. 550. —
Gravier, 1912b, p. 70 ; 1912c, p. 243-245 ; Gravier, 1913, p. 72-74. — Schellenberg, 1922, p. 295.
— Barnard, 1955, p. 242, figs. 7d, 7e.
Distribution : 0£f Cape of Good Hope, S. Africa.
Host : Cephalodiscus gilchristi Ridewood.
This interesting copepod was fairly well described as to superficial characteristics by the author.
We hâve made some observations on his material now deposited in the British Muséum (Natural His-
tory). Unfortunately, the State of préservation does not allow exact détermination of a number of
details, particularly of the appendages.
There are some females preserved entire. In these the body is seen to conform weli enough
to Calman's présentation (fig. 53, d : female, dorsal view : fig. 53, e : urosome). We are able to présent
some information about the head appendages (fig. 53, f : female, apex of head and mouth area, ventral ;
53, g : another specimen, similar). The dissected material includes antennules (fig. 53, h) and antennae
(figs. 53, i, j). We corroborate these are extremely reduced, and by this fact indicate the affinity,
but hardly offer substantial évidence for the position of the animal in the general scheme. Calman
reported he made out in satisfactory fashion a rudimentary mandible. His statements and figures
only indicate the position and some rather general outlines. The state of the material did not offer
our examination further details ; in fact we cannot actually confirm the existence of the mandible.
We find (figs. 53, f, g) the presence of only a single pair or mouthparts, these situated well posterior
to the well-developed oral cone.Without anatomical details or observations on development it is impos¬
sible to fix an exact identification on these appendages.
Source : MNHN, Paris
158
PAUL L. ILLG AND PATRICIA L. DUDLEY
The thoracic legs (figs. 52, k-n, first to fourth legs), in our examination, confirmed Calman’s
présentation. As he suggested they most closely approach those of Enterognalhus, with indeed quite
remarkable conformity. We présent in our figures what we could make out in additional details of
apparent articulations and the insertions of setae and spines. In the figures of the endopodites some
of our lines would seem to indicate that the spine-like ornaments are articulated on their articles ;
such is not the case, ail are in direct continuation with the supporting élément.
On a rather crumpled préparation of the urosome {fig. 52, 0) we confirmed that the so-called
fifth leg is as was described, and that it bears 2 well-developed but small setae on the margin. The
caudal ramus bears an apical assemblage of 4 somewhat differentiated éléments.
On the préparations of the males we could confirm somc of Calman’s présentations in general.
The antennule (fig. 53, p) offers no problem in placement in the ascidicolid sériés. The antenna
(fig. 53, q) is also a reasonable example of an ascidicolid general type, with an interstingly differentiated
set of apical éléments. In the mouth area (fig. 53, r) we could not confirm the daim of presence of
mandibles, indeed are not certain there is any definitely developed oral appendage. The labrum and
labium form a strong oral cône. It must be said that the material is in very poor condition and our
observations are in no way conclusive. We could not find préparations to make observations on the
thoracic appendages and urosome of the male, but see no reason at ail to contest Calhan’s quite rea¬
sonable présentation. The legs do not offer strong evidence for the position of the genus, but there
is no apparent fact contradictory to Calman’s conclusion.
Zanclopus antarcticus Gravier, 1912
Zanclopus antarcticus Gravier, 1912a, p. 1440 (type locality. Ile Jenny, [Marguerite Bay], latitude 68*» S.,
Longitude [Paris] 70.20® W., from Ceplu^discus anderssoni Gravier, 1912) ; 1912c, p. 240-245,
fig. 1-4 ; 1913, p. 68-74, fig. 53-62. — Schellenberg, 1922, p. 295.
Distribution : Antarctic.
Host : Cepkalodiscus anderssoni Gravier.
In his description Gravier characterizes this form very well and corroborâtes Calman’s dia-
gnosis of the genus as far as the generalities of body outline and form of thoracic legs are concerned;
his idea of the presence of mouthparts and the possible identities substantially differs. The resolution
of the situation is not possible with présent information.
The addition of Gbavieb’s species adds a certain confirmation to the long-standing assignment
of the genus to the Ascidicolidae. From information now available it does not belong to any of the
ascidicolid subfamilies previously proposed and its position remains, as originaily suggested in close
relation to Enterognalhus. We tentatively assign it to the subfamily of the latter, a solution which
at least has the rnerit of finding a place in the overall copepod scheme for this interesting animal. We
maintain the serions réservation that future thorough investigation may well challenge this disposition.
We agréé with the assertion of Gravier that the many parallels with the annelidicole poecilostome
généra leave the question a very open one. However, finally, there remains the very firm suggestion
toward ascidicolid afllnity provided by the antennule of the male of Zanclopus cephalodisci.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMIJ.IKS
159
III. — DEVELOPMENT
In his revisionary treatment of the former Notodelphyoida, Lang {1948) suggested that the
Enterocolidae and its relatives should be incorporated into the Cyclopoida Poecilostoma. Among
his substantiating evidence, he presented a comparison of the biology and development of Mycho-
philus roseiis, first nauplius stage, Nolodelphys agilis and Ascidicola rosea. He stated “ the first nau-
plius stage in Mychophilus differs in one or two respects from the corresponding stage in Nolodelphys
and Ascidicola and seems to show greater similarity to the first nauplius stage of the poecilostomes.
It cannot, however, be determined with certainty how this likeness is, as the first nauplius stage of
the poecilostomes is only known in a few forms and as the figures relatingto this stage given by various
zoologists differ from each other in more than one respect We thoroughly agréé that the study
of the poecilostomes is incomplète. Some original observations made by us (Dudley, Ph. D. thesis,
1957 ; 1966, p. 155-156) show immediately that important différences exist among poecilostomes,
as well as among gnathostomes, because of a différentiation in either sériés of planktotrophic and leci-
thotrophic developmental forms. Since ail ascidicolous copepods known in development so far are
lecithotrophic it is important to compare these with lecithotrophic poecilostomes. Such forms are rare,
and we hâve been able to study only one. The literature has furnished only preliminary information
on such forms. At this point, therefore, the position of the lecithotrophic poecilostomes is uncertain,
on developmental evidence. Further, there is no appendage in the oral apparatus of the adult poeci¬
lostomes which actually corresponds structuraUy to the appendages of the ascidicolids. The mere
lack of an appendage in either sériés is not a positive relating character. Furthermore, the lack of the
prehensile modification of the antennule in ascidicolid males i? equally uniformative (cf. Dudley,
1966, p. 156-157). The fact that the nauplii of the ascidicolids (ail lecithotrophic) hâve certain défi-
nite similarities with poeciolostome lecithotrophic nauplii is principally a resuit of parallel adaptations
in this pattern of development.
Ail lecithotrophic cyclopoid nauplii that hâve been reported or that we hâve studied hâve
definite characteristics in common : The development is consistently extremely rapid, ranging from
1 1/2 days for completion of the entire naupliar sequence {Enleropsis capiiulatus) to 6 days (Pseudomyi-
cola sp.). Ail such nauplii are distinctive for the very thin cuticle which shows little or no external
evidence of internally developing postmandibular appendages during the later naupliar stages. There
are actually postmandibular rudiments présent from the first naupliar stage, and by late stages the
appendages through the third legs are very definitely differentiated, but the external expression is in
simple fold-like protrusions of the exuviae, indicative only of the margins of the appendages. The
basic naupliar appendages (antennule, antenna, mandible) hâve a particularly simple pattern of struc¬
ture, notably expressed in the lack of feeding modifications or accessory structures. Probably the
function of these appendages combines sensory and locomotor considérations. In the notodelphyids
the gut, although fully developed, is not patent, nor in direct communication with the stomodaeum
or proctodaeum, and we assume from external observations that this feature is consistent throughout
the lecithotrophic nauplii of the ascidicolids.
Further studies of the diversity of naupliar characters in the ascidicolids as we presented them
above, show a solficient range of variability that generalizations about the nauplii of the ascidicole sériés
(notodelphyids plus ascidicolids) wül doubtless corne to show extensive overlap with a similar généra-
Source ; MNHN, Paris
m
PAUL L. ILLG AND PATRICIA L. DUDLEY
lization for the lecithotrophic poecilostomes when they corne to be equivalently analyzed. In our
comparison below of the descriptive features of the appendages of the first nauplius of ascidicolids
oiir inclusion of notodelphyids and the poecilostome Pseudomyicola sp. will demonstrate such an over-
lap,
So far there exist reports on accurate déterminations of the numbers of naupliar stages only
for the genus Aseidicola (4 stages, Gotto, 1957), various généra of Notodelphyidae (5, rarely 4, Dudley,
1966), and for a few lecithotrophic poecilostome forms, namely Mytilicola (2, Pesta, 1907 ; Caspers,
1939) and the généra Sabellackeres (4) and Gastrodelphys (2) in the Gastrodelphyidae (Dudley, 1964).
Excellent partial accounts of life historiés hâve been reeorded, including details of one or more naupliar
stages, as in the monograph of Canu (1892). Our iindings are also fragmentary, but with the existing
background we hâve indications of the major features. There seems to be a strong tendency to per-
sistence of a sequence of naupliar stages. For many cases, 4 or more deGnite naupliar stadia hâve
been deterniined.
Naupliar appendages of Bolryllophüus sp., “ Aplostoma ”, Notodelphys sp., Pseudomyicola sp.
(Dudley, 1966) and Aseidicola, Enterocola, Enteropsis, above : The antennule is 3-segmented in the
First nauplius throughout the sériés of lecithotrophically developing cyclopoids. The basal 2 segments
approximately equal the length of the terminal. The basal article is unarmed. The second article
bears 3 setae in Notodelphys and Pseudomyicola, 3 setules in Aseidicola, 2 setae in “ Aplostoma ”, 1 seta
and 2 setules in Enteropsis, 1 seta and 1 setule in Enterocola and 1 seta only in Bolryllophilus. The
terminal article bears 1 long and 1 shorter seta in ail forms and an aesthete in ail except ” Aplostoma ”,
Aseidicola, Enterocola, and Pseudomyicola.
The antenna is composed of a bimerous protopodite without any gnathal modiGcations, throu¬
ghout and the segmentation of the rami varies. The exopodite is tetramerous (some Enterocola, Botyl-
lopkilus “ Aplostorrut ”, Pseudomyicola), or obscurely tetramerous, that is, with the basal article comple-
tely fused with an extension of the basipodite {Notodelphys, Enteropsis, Aseidicola, Enterocola). The
endopodite is unimerous.
The coxopodite is unarmed throughout. The basipodite has 1 médial setule in Notodelphys
and Aseidicola and 2 médial setules in Pseudomyicola. In the others it is unarmed. Each of the
3 basal articles of the exopodite bears 1 long médial seta in ail cases. The terminal article of the exopo¬
dite bears 2 short setae or setules and 1 long seta in Notodelphys, Enterocola, “ Aplostoma ”, Botryllo'
philus and Pseudomyicola. There is only 1 short seta or setule and 1 long seta in Enteropsis and Asci-
dicola. The endopodite bears 1 médial setule and 2 terminal setae in ail. In Pseudomyicola the médial
seta and 1 terminal seta are stiffened and are margined by a hyaline Gange.
The mandible is composed of a bimerous protopodite, without gnathal modiGcations, tetra¬
merous exopodite and bimerous endopodite in Notodelphys, Enterocola, “ Aplostoma ”, BotryUophilus,
Aseidicola, and Pseudomyicola. In Enteropsis the exopodite is unsegmented and fused with the basi¬
podite.
The coxopodite bears a médial spine-like element in Aseidicola and is unarmed in ail others.
The basipodite in Notodelphys bears 1 setule and 1 minute pointed process. In Enterocola there is 1
médial unarticulated pointed process. In Pseudomyicola there is 1 setule and 1 spine. In ail the
others the basipodite is unarmed. The exopodite bears 4 long setae and 1 short terminal seta in Noto¬
delphys ; 4 long seta and 1 unarticulated spiniform process in Enterocola. In ail the others this ramus
bears only 4 long setae. The basal article of the endopodite bears 1 médial setule in ail except Pseu¬
domyicola, in which the article is unarmed. The second article bears 2 setules and 1 seta in Notodel¬
phys ; 1 setule and 2 setae in Enteropsis, Enterocola, and BotryUophilus ; 1 seta and 2 Ganged spines in
Pseudomyicola ; 2 setae in “ Aplostoma ”. In Notodelphys, and ail other notodelphyids, the mandi-
bular endopodite has a very characteristic form ; the endopodite is only as long as the basal article of
the exopodite and the setae are very reduced. In all the others the endopodite is much longer, reaching
to the second or even to the end of the third article of the exopodite and the terminal setae are as long
as those of the exopodite.
Lang’s (1948) présentation for the Grst nauplius of Mychophilus is in essential agreement in
terms of the basic pattern of segmentation.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
161
Although, as demonstrated by Dudley (1966) there is a very reliable salient characteristic
for récognition of notodelphyid nauplii, namely a distinctive outline of the endopodite of the mandible,
there is no equally reliable differentiating character for the first nauplii of the ascidicolids, nor can
they be easily separated from the poecilostome sériés. However, it can be shown by use of the charac-
ters of the early copepodids that the ascidicolid group is cohesive and shows a greater afflnity with the
iiotodelphyids than with any poecilostome genus.
In the poecilostomes (Mytilocola, Pesta, 1907 ; Caspbrs, 1939 ; Sabellacheres, Dudley, 1964 ;
Pseudomyicola sp., Dudley, 1966) the identifying cephalic mouthparts appear at the first copepodid
stage and immediately establish the divergence from ail the gnathostomes. It is interesting to note
in Sabellucheres there is a vestige of the antennal exopodite in the first copepodid, which is lost in the
ni'xt moh. This feature is also characteristic of all notodelphyids and ascidicolids for which we hâve
information, and also occurs in Cyclops {s. lot.) and in Oilhona, so might be regarded as a generalized
cyclopoid characteristic, although it definitely does not occur in some poecilostomes. The mandible
lindergoes metamorphosis during the last naupliar stage in typical peocilostomes and is lost entirely
in the last naupliar molt in Mylilicola.
The resulting condition of the appendage, in any case, is similar to that in the adult. In the
notodelphjids and ascidicolids, the mandible of the first copepodid is characteristic for each particular
genus, and represents a transformation from the naupliar appendage, but always undergoes subsé¬
quent modification through ensuing copepodid stages to attain the adult characteristics. This modifi¬
cation may indeed be as extreme as the complété loss of the mandible at the third copepodid stage in
Enteropsis and Enterocola. Other characteristic modifications of the mandible and of the legs tend
also to appear at the third copepodid, a feature of the whole assemblage, representing a second major
metamorphosis, related to the event of infection of the host. This is a different pattern from the sequence
of poecilostome development at all stages.
Tracing further development of lecithotrophic poecilostomes shows distinct différences emerging
from the pattern of development in ascidicolids, particularly with the appearance at the first copepo¬
did of the identifying mouthparts. Pseudomyicola àoe:s notpossess any evidenceofan antennal exopodite
at the first copepodid stage) we know some poecilostomes do {Sabellacheres, Dudley, 1964). Myti-
licola goes further than Pseudomyicola in réductions before the molt to the first copepodid stage, since
in this molt it loses the mandible. Mylilicola and Pseudomyicola also show similarities in not further
modifying the appendages of the cephalosome appeciably after the first copepodid except for the diffé¬
rentiation of the dimorphic maxilliped in the late male stages. Ascidicolids, then, show trends of
réduction that are distinctiy different from those in Mylilicola and Pseudomyicola, representing the
poecilostomes.
Such nauplii demonstrate there is never any sign of a maxilliped in the species of Enlerocola shown
by Canu and us, and in the Enleropsis species studied. The mandible can be traeed into the first
copepodid stage and it can be shown that this appendage disappears during copepodid molts in Ente-
ropsis and Enterocola. The exopodite of the antenna is similarly lost during the first copepodid stage
in all notodelphyids and ascidicolids we hâve studied. Exactly this also happens in Cyclops (Dietiuch,
1915) this feature thus serving to tie all the gnathostomes together. Further strong evidence for rclating
the ascidicolids and the notodelphyids would corne from the following : In Notodelphjs, Ascidicola,
Enlerocola, Bolryllophilus, Enleropsis, and “ Aplosloma ” (Canu) the naupliar appendages are strikin-
gly similar. The antennule bas 3 segments ; the antenna has a protopodite of 2 segments, a basically
tetramerous exopodite and a unimerous endopodite ; the mandible is biramous with bimerous proto¬
podite, basically tetramerous exopodite and bimerous endopodite. Lang’s présentations concerning
the nauplii of Mychophilus incline us to believe these are in essential agreement. The patterns of
setation throughout, while showing some minor différences are sufficiently consistent as to offer to us
reinforcement of this agreement.
Although the above references furnish some indications of the structure of nauplii and late cope¬
podids of some of the species in the Ascidicolidae, the only early substantiel published study of develop¬
ment is that of Canu (1892). He furnished basic and significant information on the life historiés of
Enlerocola fulgens and “ Aplosloma ” breoicauda. The literature has furnished some descriptions of
11
Source : MNHN, Paris
162
PAUL L. ILLG AND PATRICIA L. DUDLEY
males based on subadult stages, and these refer in part then to life history fealures. \Ve refer to such
information under the individual general above.
In collections from the host ascidians and from cultures from developing egg masses we hâve
been able to obtain the first nauplius and the first three copepodid stages of Ascidicola rosea ; the first
nauplius and ilrst 2 copepodids of Enterocola fertilis ; the lirst nauplius and a fourth copepodid of
Enterocola latieeps ; the first nauplius and first 2 copepodids of EnUropsis capitulatus ; and the fourth
and fifth copepodids of EiUeropsis superbus. Although we hâve not observed a complété life history,
the descriptions below show great basic conformities and give additional evidence for our concept
of the family Ascidicolidae.
Development of Ascidicola rosea
(Figures 54, 55)
The nauplius is a lecithotrophic larva and has the basic characteristics of cyclopoids of non-
feeding type. The antennule (fig. 54, a) is uniramous and functionally trimerous. The basal article
lacks armature and ornamentation ; it may be only a mechanically difîerentiated element. The
second article bears a seta, which is about 1/3 the length of the article, on the ventral face, at the distal
margin. Proximal to this, on the ventral margin, there are 2 shorter, equispaced setae on projections
of the margin. Three rows of spinules ornament the médial surface. The terminal article is somewhat
longer than the second and tapers slightly. On the apical margin there is a long, ventral seta and
somewhat removed from this there is a dorsal seta which is about 1/3 as long. There is no aesthete.
The antenna (fig. 54, b) is biramous, the protopodite bimerous, the endopodite unimerous, and
the exopodite tetramerous, but not articulated on the protopodite. The basal article of the proto¬
podite lacks armature and ornamentation. The distal article has a small marginal setule near the
articulation with the endopodite. The endopodite, which reaches only slightly beyond the distal end
of the basal article of the exopodite, bears 1 setule in an emargination at the middle of the médial
margin, and 2 long terminal setae. The basal article of the exopodite shows no clear articulation with
the protopodite but probably is represented by an extension of the latter terminating in a long, arti¬
culated médial seta. This extension is longer than the 3 terminal articles of the exopodite. The
second and tbird articles of the exopodite are wider than long and each bears a long seta at the distal
médial corner. The terminal article is rounded, with an emargination on its médial face ; a short seta
is inserted in this. Apically there is a long seta. Rows of spinules are found on the articles of the
ramus.
The mandible (fig. 54, c) is biramous, with bimerous protopodite, bimerous endopodite and
tetramerous exopodite. The basal article of the protopodite has a distal médial setule. The second
article is unarmed. The endopodite is long, subrectangualar and reaches to the end of the third article
of the exopodite. The basal article beafs a small setule at the distal médial corner. The second article
bears a minute setule just beyond the middle of the médial margin and 2 long apical setae. The exopo¬
dite is poorly articulated with the protopodite. The first 3 articles each bear a long seta at the distal
médial corner. The terminal article is rounded and bears a long apical seta. There is a row of spi¬
nules on the basal article of the exopodite. Ail of the long setae of the antenna and mandible bear
long coarse hairs.
In the successive naupliar stages, setae are added to the antennule, and the other appendages
begin the régression which leads to the form of the appendage in the copepodid stages. In the last
nauplius, containing internally the structures of the first copepodid, the following culicular structures,
whieh are still naupliar, can be made out.
The antennule is still trimerous, the basal article unornamented ; the second article has reduced
setae, consisting of a very short seta at the distal end of the article and 1 setule. Therefore, during
the naupliar stages, 1 seta has been lost from this article. The terminal article still bears a distal
elongate seta, accompanied by a short seta, but 5 or 6 setules hâve been added to the ventral terminal
margin and 1 short setule to the dorsal terminal margin.
Source : Mt-JHN, Paris
,G 5/. - Ascidicola rosea : f.rst nauplius fa^) ; firet copepodiü (d-n) ; second copepod.d o-p) : a, antennule ; b. antenna ;
c, niandible ; d. habitus, latéral ; e. antennule ; f, antenna ; g, mand.ble ; b, max.llule ; n.axilla , j, maNinipcd ,
k, first leg ; I. second leg ; m, ihird leg ; n, caudal ramus ; o. babil us, latéral ; p, antennule. >,cales tor a, b, c. d,
k, 1, m, n, 0 = 0.1 mm ; scalcs for e, f, g. b, i. j = 0.05 mm.
Source ; MNHN, Paris
Fie. 55. — Ascidicola roa^a : second copepodid (a-i) ; third copepodid (j-q) : a, antenna ; b, mandible ; c, maxillule ;
d, maxilla ; e, firet leg ; f, second leg ; g. third leg ; h, fourth leg, with dolted line indicating forming fourth leg
o{ third copepodid j i, terminal somite ot urosome and caudal rami, dorsal ; j, habitus, latéral ; k, antennule ;
1, antenna ; m, flrst leg ; n, second leg ; o, third leg ; p, fourth leg ; q, temunal somite of urosome and caudal rami,
ventral. Scales for a, b, c, d, k, 1 = 0.05 mm ; other scales = 0.1 ram.
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
165
In the antenna, the protopodite and exopodite are the same as in the first naupliar stage, the
médial seta of the endopodite has been much reduced.
In the mandible the protopodite and exopodite are the same as in the first stage, the médial
seta of the endopodite is reduced.
Cuticular Unes indicate the margins of the maxillule, maxilla and maxilliped, which are deve-
loping internally. The first and second legs are visible on the cuticle as protruding sacs within which
the well-formed legs are differentiating. The posterior end of the last nauplius is bilobed. Each lobe
bears 1 long seta and 5 short setae.
When the late last nauplius is examined in life, features of the first copepodid can be seen inter-
nalty. It would appear that the antennule will hâve 5 articles. The terminal article of the naupliar
appendage will divide into 2 articles in the copepodid. Details of the first 2 legs, the presence of
third leg rudiments and the developing caudal rami are visible. Ail the naupliar stages lack an eye.
This feature has been remarked on both in regard to the adult and the nauplius by various authors,
and we can confirm the observation.
In this species we hâve been able to study the first 3 copepodids which were obtained by
rearing the nauplii. We obtained large numbers of first and second copepodids, but only a single
third copepodid. The first copepodids were moltingto second stages after about 24 hours. The single
copepodid to molt to the third did so after about 96 hours. We are convinced that the second cope¬
podid is the infective stage.
In the first copepodid the body (fig. 54, d) is composed of the cephalosome, metasome and uro-
some. The overall length is 0,35 mm. By analogy with later developmental stages and the observed
anatomy we concluded the cephalosome to include the portion of the body anterior to the segment
of the first legs and there appears to be an actual articulation. The metasome has 2 leg-bearing seg¬
ments, these very well-defined and each with a dorsal plate-like sclerotization. The urosome consists
of a segment bearing rudimentary third legs, 1 leg-less segment and the anal somite with its terminal
caudal rami. There is no salient rostrum. The antennule (fig. 54, e) is obscurely 4-segmented. There
is a slight taper from base to apex. The basal segment is about twice as long as any other. The arma¬
ture is as follows : article I — 3 setae ; II — 3 setae ; III — 2 setae ; IV — 6 setae, 1 aesthete. There
is no vestigial seta on the apical segment as seen in the first copepodids of Enterocola and Enteropsis.
The antenna (fig. 54, f) is obscurely trimerons and early in the stage is clearly biramous, but
very soon the internai tissue of the expodile régresses and the cuticle crumples as shown in the figure.
The basal article has a large flattened spinulose seta at about the middle of the outer margin, and
there is a row of spinules distally on the outer margin. The terminal segment, approximately equal
in length of the other 2 combined, bears 1 terminal flexible hook-like élément and 4 fine setae. In
addition there are 2 setae at the proximal third of the outer margin.
The mandible (fig. 54, g) early in the stage is biramous, but very soon the tissue in both rami
régresses and the cuticle tends to crumple. However, in the later portion of the same stage one can
discern the palp of the second copepodid forming within the cuticle, demonstrating that this palp
includes the endopodite of the appendage, and there is also a possibility that the basal portion of the
palp is the basipodite. The coxopodite has a toothed médial margin, but the teeth are not so complex
as in later stages and the article is narrow.
The maxillule (fig. 54, h) is bilobed and has an outline similar to that of the adult. Nearthe
distal médial corner of the basal segment there is an endite bearing 2 unarticulated setae. The apical
segment is drawn out medially as a narrow setiform process and there are 3 setae on the apical margin.
The apical segment thus lacks three setae présent on this segment in the adult.
The maxilla (fig. 54, h) is bimerous, the distal article about 1/4 the mass of the basal. On the
latter the médial margin is produced at about its middle to a rounded process terminated in 2 unequal
pointed processes. The distal article has a sloping antérolatéral margin. This terminâtes at the ante-
rodistal corner as a pointed process. A slightly smaller spiniform seta is set on the base of the process
and 2 somewhat smaller spiniform setae are sel at intervals on the distal margin.
The maxilliped (fig. 54, j) is a very sraall, narrow lobe with 2 apical setae and a médial setule.
The first leg (fig. 54, k) is biramous, with unimerous protopodite and rami. The subquadrate
Source : Paris
166
PAUL L. ILLG AND PATRICIA L. DUDLLY
intercoxal lamella is well developcd. The protopodite lias a small seta at the distal latéral corner iiear
the articulation of the exopodite. There is also a pointed protrusion at the distal médial corner ;
but no articulated spine is présent as yet. The exopodite, flattened and considerably longer than
the endopodite, has 3 short latéral spines, 1 very long terminal spine, 1 terminal seta and 2 inedial
setae. The endopodite has 1 latéral seta, 2 terminal setae and 1 distal médial seta.
The second leg (fig. 54,1) is biramous with unimcrous rami. The intercoxal lamella is subqua-
drate. There is a relatively long seta on the latéral margin of the protopodite, proximal to the arti¬
culation of the exopodite. The exopodite bears 3 latéral spines, 1 terminal spine, 1 terminal seta and
2 médial setae. The endopodite, about 2/3 as long as the exopodite, bears 1 distal latéral seta, 2 ter¬
minal setae and 2 distal médial setae.
The third leg {fig. 54, m) is non-functional and the basal articulation with the body is unde-
veloped. Each leg consists of 2 flaps which encapsulate the developing rami of the leg of the second
copepodid. The cxopodal portion has 3 setiform projections and the endopodal 2.
The second urosomal segment and the anal somite proper hâve no ornamentation. The caudal
ramus (fig. 54, n) is slightly shorter than the anal somite and subrectangular in outline. There are
6 setae on the ramus, 1 at the proximal latéral third, 1 on the dorsal surface at about the distal third
and 4 terminal. The médial terminal seta is the longest, and is slightly more than twice the length
of the ramus.
The second copepodid (fig. 54, o) has the body composed of cephalosome, bearing appendages
through the maxilliped, metasome of 3 leg-bearing segments, and a urosome of 2 segments and the anal
somite ; with terminal caudal rami. The overall length is 0.36 mm. There is no defined rostrum.
The antennule (fig. 54, p) is clearly 5-segmented, with only a slight taper. The basal segment
is the longest. The armature is a follows : Article I — 3 setae ; II — 4 setae and a hook ; III — 2 setae ;
IV — 3 setae, 2 hooks ; V — 6 setae, 2 aesthetes and 1 hook.
The antenna (fig. 55, a) is trimerous and uniramous. The exopodite has been lost at the molt.
Each of the 2 basal segments bears an outer distal spatuliform spinulose seta and a row of spinules on
the distal external margin. The apical segment, about equal to the combined lengths of the 2 basal
segments, and somewhat narrower, has 4 apical setae and a narrow hook, articulated terminally. At
the proximal third there are 2 short setae. Curved rows of spinules ornament the surface at the proxi¬
mal third and distal fifth. The mandible (fig. 55, b)issimilartothat oftheadult. There is a coxal lamella,
with complex articulated spines directed medially, and a unimerous palp, which bears 4 unequal setae
on its flattened anterior margin. The setation of the palp thus conforms to the adult in number but
not in the form of the setae.
The maxillule (55, c) is bilobed, with a poorly defined articulation between the base and the palp
lobe. An endite-Iike lobe continues directly from the base without articulation. The palp bears
7 setae and a setule on the latéral and apical margins. The endite has only 4 setae and thus has lost
the basal seta seen in the appendage of the first copepodid. This basal seta is also absent in the adult,
but several marginal setae are added.
The maxilla (fig. 55, d) is very similar to the appendage of the first copepodid except that the
setae hâve lengthened appreciably.
The first leg (fig. 55, e) is biramous with bimcrous protopodite and rami. The trapézoïdal
intercoxal lamella is well developed. The coxopodite is unarmed and unornamented The basipodite
bears a relatively long seta on the latéral margin and an articulated spine at the distal médial corner.
The exopodite bears 1 spine at the distal latéral corner of the basal segment. The distal article bears
3 latéral spines, 1 terminal spine, 1 terminal seta and 2 médial setae. The basal segment of the endo¬
podite bears no armature. The terminal segment bears 1 latéral seta, 2 terminal setae and 1 médial seta.
It should be noted that the appendage has gained no element of armature at this stage from the condi¬
tion seen in the first copepodid. The setae ail are plumose, but there has been a shortening of the setae
of the endopodite.
The second leg (fig. 55, f) has a bimerous protopodite and rami. The coxopodite is unarmed.
There is a relatively long seta laterally on the basipodite. The basal segment of the exopodite bears
1 distal latéral spine. The terminal segment bears 2 latéral spines, 1 terminal spine, 1 terminal seta.
Source : Mt-JHN, Paris
ASr.IDICOLIDAE AND SUBFAMILIf-S
167
and 3 médial setae. Thus 1 médial seta has been added at this stage. The basal segment of the endo-
podite has no armature. The terminal segment has 1 distal latéral seta, 2 terminal setae and 2 distal
médial setae. Thus no element of armature has been added at this stage. The setae do not show
the pronounced shortening seen in the endopodite of the first leg.
The third leg (fig. 55, g) has a unimerous protopodite and rami. A subrectangular intercoxal
lamella connects the protopodites. A seta articulâtes on the distal latéral margin of the protopodite.
The exopodite bears 3 latéral spines, 1 terminal spine, 1 terminal seta and 2 médial setae. The endo¬
podite bears 1 distal latéral seta, 2 terminal setae and 2 distal médial setae.
At the posterior margin of the first segment of the urosome are found the rudimentary fourth
legs (fig. 55, h). Each consista of a bilobed pad with a single seta articulating on the latéral lobe. In
the late second copepodid one can discern the outline of the biramous leg of the subséquent stage deve-
loping within the cuticle, as figured.
The second urosomal segment and the anal somite hâve no ornamentation. The caudal rami
(fig. 55, i) are approximately as long as the anal somite. Each bears 6 setae, 1 of these just posterior
to the proximal third of the latéral margin ; 1 dorsally at the distal fifth ; and 4 apical. The médial
seta is greatly reduced over the condition of the previous copepodid and that next latéral is now the
longest, approximately twice as long as the ramus. The most latéral seta on the apex has been reduced
to a spiniform element.
The third copepodid (fig. 55, j) shows some modifications, with partial fusions of body segment
and a general shortening of ail the setae of the appendages. Because we had only a single specimen
we could not completely make out ail the appendages. The antennule (fig. 55, k) is obscurely penta-
merous. The basal portion shows indications by indentations in the cuticle that 2 segments, clearly
demarcated in the second copepodid, are incorporated by a considérable degree of fusion. The portion
of this basal complex corresponding to the first segment of the second copepodid bears 2 setae and a
setule. The distal portion of the complex bears 5 setae and 3 thick hook-like éléments. The remaining
formula for the antennule : Article III — 2 setae, 1 aesthete ; IV — 2 setae, 2 hook-like éléments, 1 aes-
thete ; V — 6 setae, 1 hook-like element, 1 aesthete.
The antenna (fig. 55, 1) is very similar to that of the second copepodid. It is trimerous ; the
2 basal articles and the apical armature of the terminal article are exactly as in the second copepodid.
One seta on the inner margin of the terminal article and 1 curved row of spinules on the face appear
to hâve been lost.
The mouthparts are not illustrated, but dissection showed that they were very similar to those
of the second copepodid.
The first leg (fig. 55, m) has a unimerous protopodite and bimerous rami. There is no artîcu-
lative line between the coxopodîte and basipodite, this havîng been lost in the transformation from the
second copepodid. There is a seta at the distal latéral corner and a spine on the distal médial corner
of the protopodite. The exopodite has 1 spine at the distal latéral corner of the basal article, 2 latéral
spines, 1 terminal spine, 2 terminal setae, and 1 médial setule on the distal article. Ail of these éléments
hâve been greatly reduced from the condition of the second copepodid, and 1 médial seta has been
lost. The basal article of the endopodite bears no armature ; the terminal article bears 1 distal latéral
setule, 2 terminal spines, and 2 distal medial setae. Although the number of éléments of this ramus
is the same as in the second copepodid, the setae hâve been greatly reduced in size and the 2 terminal
setae hâve been transformed into spines.
The protopodite of the third leg (fig. 55, o) is as in the second leg. The rami are now obscurely
bimerous rather than unimerous as in the previous stage. The basal article of the exopodite bears
a single distal latéral spine. The terminal article bears 2 latéral spines, 1 terminal spine, 1 terminal
setule, and 3 medial setules. One setule has been added to the medial margin. The setae of the ramus
hâve been greatly reduced from the condition of the second copepodid. The poorly defined basal
article of the endopodite bears no armature. The distal article bears 1 distal latéral setule, 2 terminal
spines and 2 distal medial setules. Numerically the éléments of armature correspond to the second-
copepodid but the setae are greatly reduced and the terminal éléments of the ramus bave been trans¬
formed from setae into spines.
Source : MNHN, Paris
168
PAUL L.
AND PATRICIA L. DUDLEY
The fourth legs (fig. 55, p) which were markedly rudimentary in the previous stage now consist
of a unimerous protopodite, with a latéral seta, and unimerous rami ; the exopodite bears 3 setiform
éléments and the endopodite 2.
The urosome consists of only 2 segments and the anal somite. The first segment bears 2 distal
setae, 1 on either side. Each is the rudiment of a fifth leg. The caudal rami (fig. 55, q) are approxi-
mately as long as the anal somite. Each hears 6 setae, 1 at the middle of the latéral margin, 1 on the
dorsal surface at the distal eighth, and 4 terminal setae. The medial terminal and the 2 latéral terminal
setae are very short ; the second medial seta is the longest but it is shorter than the ramus at this stage,
thus showing a great réduction in length from the condition of the second copepodid.
We did not collect fourth or fifth copepodids.
Development of Enterocola ferlilis
(Figures 56, 57)
The first nauplius is a lecithotrophic larva, with the basic characteristics of other non-feeding
nauplii. We hâve drawn the habitus (fig. 56, a) from an exuvia. The body is somewhat globose and
not markedly pyriform, as in the Notodelphyidae. There is a protrusion at the anterior end which
is possibly homologous with the rostrum. Three pairs of appendages, the antennules, antennae and
mandibles, articulate distally to the anterior third. There is a subcircular supraoral plate and the
cuticular remnant of the inner stomodaeal lining protrudes from the mouth. This latter is closed and
the gut is not patent at this stage. The posterior end of the exuvia has 2 sectors, a ventral truncated
abdominal end which bears 2 latéral setae, and the dorsal shield, which extends over the end of the
abdomen and which appears to be the precursor of the anal shield of the copepodids. The antennule
(fig. 56, b) is trimerous, uniramous. The basal “ article ” is simply a scleritic ring that articulâtes
the appendage with the body. It bears no armature or ornamentation. The second article bears a
single long seta near the distal margin on the ventral face. Proximal to this on the ventral face and
at the middle, set on a lobe, is a very small setule. The terminal article is slightly longer than the second
and shows a certain amount of taper. On the apex there is a long ventral seta accompanied by a
dorsally placed seta which is about 1/3 as long. Ail the setae are plumose. It should be notedthat
the apical aesthete présent in other subfamilies described here is missing, an interesting feature in con-
trast to the case in Enterocola kuiceps, described below.
The antenna (fig. 56, c) is biramous with bimerous protopodite, tetramerous exopodite and
unimerous endopodite. The basal and second articles of the protopodite lack armature and ornamen¬
tation. The second article shows some coalescence with the basal article of the exopodite. This basal
article of the exopodite is longer than the other articles combined. Each of the 3 basal articles of the
exopodite bears a seta at the distal medial corner. The terminal article is rounded apically, but with
a latéral and a medial emargination, in each of which articulâtes a short seta. There is a long terminal
seta on this article. The endopodite is subrectangular and reaches approximately to the end of the
second article of the exopodite. It bears a short seta in an emargination at the middle of the medial
margin and 2 long terminal setae. AH long setae of the appendage are coarsely plumose.
The mandible (fig. 56, d) is biramous with bimerous protopodite, tetramerous exopodite and
bimerous endopodite. The basal article of the protopodite bears no armature or ornamentation. The
basipodite has a minute point at the middle of the medial margin. The 3 basal articles of the exopo¬
dite bear single setae at the distal medial corners. The rounded apical article has a pointed unarti-
culated terminal process and a terminal articulated seta. The basal article of the endopodite has a
minute setule at the middle of the medial margin. The second article, broadening somewhat at the
apex, has a minute setule in an emargination at about the middle of the medial margin, and two sube-
quai long apical setae. Ail setae of the appendage are coarsely plumose.
Although our cultures were vigorous and we obtained many first and second copepodids, we
did not make a complété study of the detailed changes in the nauplii. The naupliar development
was very rapid, with about 30 hours elapsing from hatching to the first copepodid stage.
Source : Mt'IHN, Pans
Fie. 56. ErOerocola feriilU : fiwt nauplius (a-d) ; firet copepodid (e-n) : a, habitus, ventral ; b, antennule ; c, antenna ;
d, mandible ; e, habitus, latéral ; f, rostrum ; g, antennule ; h. antenna ; i, mandibular remuant ; j, maxillule ;
k, maxUla ; 1, first leg ; m, second leg ; n, end of uroeome and caudal rami, ventral. Soales for a, b c de)
m, n = 0.1 mm : for f, g, h, i, j, k = 0.05 mm.
Source ; MNHN, Paris
Fig. 57. — Enlerocola fertilis : second copepodid : a, habitus, latéral ; b, urosome, ventral ; c, rostnim, latéral ; d, anten-
nule ; e, antenna : f, en face view of posterior cephalosome : mand. = madible, Mxl = maxillule, Mx2 = maxilla,
p.o.p. SS postoral protubérance ; g, maxillule ; h, maxilla ; i, first leg ; g, second leg; k, third leg. Scalee for a, b,
i, j, k = 0.1 mm ; other scales = 0.05 mm.
Source : MNHN, Paris
ASCIDICOLIDAü: and SUUI'AMII.IICS
171
The ])ody of the first copepndid (fig. 56, e) is composed of the cephalosoino, metasome and uro-
some. The cephalosome includes the portion of the body anterior to the segment of the first legs.
The metasome has 2 leg-bearing segments. The urosome is composed of 2 segments and the anal
somite bearing the caudal rami. The first urosomal segment bears the rudimentary third legs.
Thcre is a rostrum (fig. 56, f) which is long and bulbous, rcaching beyond the bnses of the anten-
iiae. The antennule (fig. 56, g) is obscuroly tetramerous. The basal and terminal articles are set
ofl' Ly distinct articulations, but the second and third articles are dcHmited from each other only by
a cuticular indentation. The armature is as follows : Article I — 3 setae ; If — 2 setae and 1 setule ;
III — 2 setae ; IV — 7 setae, 1 vestigial seta, and 1 aesthete.
The antenna (fig. 56, h) is clearly biramous at the early copepodid stage, but very soonafter
tlic molt from the last nauplius, the internai tissue of the exopodite régresses and the cuticle crumples.
The tissue within the protopodite and the endopodite does not regress, and by the end of the stage,
the appendage assumes an obscurely trimerous form, with an attached remnant of the exopodite.
The 2 basal articles bear no armature or ornamentation. The iiarrower apical article bears 3 flexible
terminal setiform éléments.
The mandible (fig. 48, i) is regressing from immediately alter the completion of the molt. It
consists of a crumpled sac with some indication of biramous cojidition. Within the crumpled sac
there is developing a minute setigerous lobe which will be the only evidence of tins appendage at the
second copepodid stage.
The maxillule (fig. 56, j) is unequally bilobed, with a more latéral large lobe bearing 2 apical
setules perhaps represonting the palp at least apically. A more basal, minute lobe extends medially
with no articulation between the parts of the appendage.
The maxilla (fig. 56, k) consists of a low lobe with a single articulated apical spiniform projec¬
tion. There is no maxilliped.
The first leg (fig. 56, 1) is biramous with unimerous protopodite and rami. The lobed inter-
coxal lamclla is subquadrate and well-dcveloped. The protopodite has a small seta at the distal latéral
corner near the articulation of the exopodite. There is a flattened spiniform projection, obscurely
articulated on the distal médial margin at the articulation of the endopodite. The exopodite has
1 spine at the proximal latéral third, 1 spine at the distal latéral third, 1 terminal spine, 1 terminal
seta, and 2 distal médial setae. The endopodite bears 1 latéral seta in an emargination at the distal
third, 2 terminal setae, and 2 distal médial setae.
The second leg (fig. 56, m) has a unimerous protopoditc and rami. The intercoxal lamella is
subquadrate. A small seta articulâtes at the distal latéral corner of the protopodite. The exopodite
bears 1 latéral spine at the proximal third, and 2 distal latéral spines, the more distal much longer,
1 terminal spine, 1 terminal seta, and 2 médial setae. The endopodite bears 1 distal latéral seta, 2 ter¬
minal setae, 2 distal médial setae and 1 proximal mcdial seta. The third legs are rudimentary, with
the basal articulation with the body undeveloped. Each leg consists of 2 sacs which encapsulate
the developing rami. The exopodal portion has 3 spinelike projections and the endopodal portion
has 2 projections.
The caudal rami (fig. 56, n) are aljout 1/2 as long as the anal somite. There arc 6 setae on each
ramus, 1 at the proximal latéral third, 1 at the distal third on the dorsal surface, and 4 terminal. The
médial terminal seta is the longest, approximately 3 times as long as the caudal ramus.
The second copepodid (fig. 57, a) has a hody composed of the cephalosome, metasome of 3 leg-
bearing segments, and urosome of 2 segments and the anal somite bearing the caudal rami (fig. 57, b).
There is a rostrum (fig. 57, c) very long and almost cylindrical. The antennule (fig. 57, d) is
clearly 4-segmented. The armature is as follows : Article I — 5 long setae and 2 short setae ; II 3
setae ; III — 3 setae ; IV — 7 setae, 1 aesthete.
The antenna (fig. 57, e) is uniramous and obscurely trimerous. Two basal articles, which are
only obscurely articulated with each other, are iinarmed and unornamented. Tlie terminal article,
which is the longest, has a terminal armature of a very narrow curved articulated hook and 3 small
setae. Each of the articles is very much narrower than in the previous copepodid stage and there is
no evidence of the exopodite.
Source ; MNHN, Paris
4
172 PAUL I» ILLG AND PATRICIA L. DUDLEY
The labrum (fig. 57, f) is well differentiated at this stage, but is completely unornamented.
The madibular remnant (fig. 57, f) consists of a lobe with an apical seta and a subapical setule.
The maxillule (figs. 57, f, g) is a bilobed structure. In an en face view the more ventral lobe
terminâtes in 2 spiniform processes. The more dorsal lobe, somewhat longer at this stage, also terminâtes
in 2 spiniform processes, and there is an articulated setule on the ventral surface.
The maxilla (figs. 57, f, h) is subtriangular in outline with an expanded base and an apical lobe
somewhat demarcated and tapering acutely apically as a hook-like process. The apical lobe bears
2 articulated éléments, the more distal possibly the rudiment of the second hook-like process of later
stages. There is a large hemispherical bulge of the cuticle between the bases of the maxillae ; we hâve
termed it the postoral projection (fig. 57, f : p.o.p.).
The first leg (fig. 57, i) is biramous, with unimerous protopodite and bimerous rami. The
subquadrate intercoxal lamella is well developed. There is a small seta on the distal latéral margin
of the protopodite and a spine on the distal médial margin at the articulation of the endopodite. The
basal article of the exopodite bears 1 spine at the distal latéral corner. The second article bears 1 spine
at the distal latéral corner, a terminal spine, a terminal seta, and 2 médial setae. The basal article of
the endopodite bears no armature. The distal article bears a seta at the middle of the latéral mar¬
gin, 2 terminal setae and 2 distal médial setae. No new éléments of armature hâve been added to
either ramus.
The second leg (fig. 57, j) is biramous, with unimerous protopodite and bimerous rami. The
intercoxal lamella is subquadrate. The protopodite bears a short seta at the distal latéral corner near
the articulation of the exopodite. The basal article of the exopodite bears a distal latéral spine. The
terminal article bears 2 distal latéral spines, 1 terminal spine, 1 terminal seta, and 3 medial setae.
The basal article of the endopodite bears a distal medial seta. The terminal article bears 1 seta at
the middle of the latéral margin, 3 terminal setae, and 1 distal medial seta. Only 1 seta has been added
to the appendage, this to the medial margin of the terminal article of the exopodite.
The third legs (fig. 57, k) are articulated with the body for the first time. Each consists of
unimerous protopodite and rami. A subquadrate intercoxal lamella is well developed. The proto¬
podite bears a short seta at the distal latéral corner. The exopodite bears 2 distal latéral spines, 1 ter¬
minal spine, 1 terminal seta, and 2 medial setae. The endopodite bears a latéral seta at the distal third,
3 terminal setae, 1 distal medial seta, and 1 proximal medial seta.
The first segment of the urosome bears the rudimentary fourth legs. Each consists of a bilobed
pad, with a single seta articulating on the more latéral lobe.
The caudal rami (fig. 57, b) are slightly shorter than the anal somite and each bears 6 setae.
The latéral and dorsal setae are as in the first copepodid, the second medial seta of the 4 terminal
is now the longest, a little more than twice as long as the ramus.
We hâve no material available of the third, fourth and fifth copepodid stages.
Development of Enterocola laticeps
(Figure 58)
We hâve been able to obtain only the first nauplius stage of this species. The first nauplius
is a lecithotrophic larva, with basic characteristics of cyclopoids of non-feeding type (see Dudlek,
1966).
The antennule (fig. 58, a) is uniramous and functionally trimerous. The basal article lacks
armature and ornamentation, and, since in similar forms, it coalesces with the head in later develop¬
ment, it may be only a mechanically differentiated sclerotized ring. The much larger second article
bears a single seta, long and well developed, inserted on the ventral face at the distal margin. Proxi¬
mal to this on the ventral margin is a minute setule, inserted in an indentation at the distal fifth. At the
distal third there is a minute pointed projection which may be homologous to such a setule. Three
rows of spinules are about equally spaced on the distal half of the medial surface. The terminal article
is somewhat longer than the preceding and tapers alightly distally to a truncate apex. On the apical
Source : Mt'IHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
173
Fie. 58. — Enterocola laticeps : fir»t naupJius (a-d) ; third copepodid (e-n) : a, antennule ; b, anteona ; c, mandible ;
d, endopodite of mandible : e, babitus, latéral ; f, urosome, ventral ; g. antennule ; h, antenna ; i, maxiUule ;
j, maxilla ; k, first leg ; 1, second leg ; m, third leg ; n, caudal ramu». Scaieg for a, b, c, d = 0.05 mm ; other
gcales = 0.1 mm.
margin there is a long ventral seta, accompanied by a short aesthete and somewhat removed from
these a dorsal seta, about half as long as the ventral seta. Ail long setae of the appendage are coar-
sely plumose.
The antenna (fig. 58, b) is biramous with an obscurely bimerous protopodite, an obscurely
tetramerous exopodite and a unimerous endopodite. The basal article of the protopodite lacks arma-
Source ; MNHN, Paris
174
PAUL L.
AND PATRICIA L. DUDLEY
turc and ornamentation. The second article lias 2 rows of spinules on the surface and 2 pointed pro¬
jections on the médial margin distal to the midpoint. The basal article of the exopodite is rather
poorly articulated with the protopodite and is slightly longer than the combined lengths of the remaining
3 segments. It bears a long seta at its distal médial corner and 3 rows of spinules on the surface.
The second and third articles are wider than long and each bears a long seta at the distal médial corner.
The terminal article is rounded, with an emarginalion on its médial face ; a short seta is inserted in
this. Apically there are a long seta and a much shorter seta. The endopodite is subrectangular and
reaches only slightly beyond the distal end of the basal article of the exopodite. It bears a setulc in
an cmargination at the middle of the médial margin and 2 long terminal setac. Ail long setae of the
appendage bear coarse hairs.
The mandible (fig. 58, c) is biramous, with an obscurcly bimerous protopodite, obscurely tri-
mérous exopodite and bimerous endopodite. The basal article of the protopodite has no armature
or ornamentation. The second article has a single row of spinules. The basal article of the exopodite
is only faintly set oiï from the protopodite and its length about equals the combined lengths of the
other 2 article. The first and second articles each bear a single seta at the distal médial corner. The
terminal article is rounded with an emargination at the middle of the margin, in which is set a single
long seta. There is a single long apical seta. The endopodite (fig. 58, d) is long, subrectangular,
reaching to the end of the second segment of the exopodite. The basal article has a minute setule at
the middle of the médial margin. The second article has a minute setule at the middle of the médial
margin and 2 long setae on the truncate apex. AU long setae of lhe appendage bear coarse hairs.
Third Copepodid of Enterocola laticeps
(Figures 58, e-n)
From material taken from the host ascidian we hâve a specimen that wc assign with some mis-
giving to the third copepodid stage. The overall size and the advanced development of the 3 pairs
of legs présent would suggest a later stage. The segmentation of the urosome, the lack of setae on the
endopodite of the legs, and the fact that lhe species is a large one are indications that this could he
the third stage.
The habitus (fig. 58, e) is cruciform, the animal having assumed a pattern of considérable con-
formity to the adult structure. The overall length is 0.66 mm, measured from the apex of the head
to the end of a caudal ramus. The body divisions are cephalosome, bearing 4 pairs of appendages ;
a metasome, of 3 demarcated segments, each bearing a salient pair of legs ; and the obscurely S-seg-
mented urosome (fig. 58, f), bearing rudiments of fourth legs on the first segment and terminal caudal
rami.
The antennule (fig. 58, g) is unimerous, but a pattern of sclcrotization suggests 4 ingrédient
articles. There are no basal setae, and distally 11 setae are rather regularly distributed.
The antenna (fig. 58, h) is obscurely bimerous, the basal portion below a weakly suggested sub¬
division much larger than the terminal portion. There are 2 minute setules on the apex.
The labrum was not discerned in the dissection. The mandible is absent.
The maxillule (fig. 58, i) is bilobed, with unsegmented base and lobes. The base extends in
a médial lobe, torming a basal piece which is rather heavily sclerotized. The minor lobe is produced
into two subequal processes. One process terminâtes acutely, the other is larger, truncate with acutc
terminal corners. On the face there is a spinulose seta. The lobe of the basal piece is obscurely arti¬
culated. It is bilobed terminally, 1 lobe bearing 2 short setae, the other with 3 setac. There is an
additional very short seta at the middle of the ventral margin. A few spinules are found on the
surface.
The maxilla (fig. 58, j) is bimerous and, although resembling the appendage of the adult, is not
quile so modified and suggests the aiiatomical composition somewhat more strongly. The basal article
is long and bears a thuinb-like, spinulose, articulated endite distally. The terminal article is produ-
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
175
ced medially as 2 strong, pointed, spinulose processes, each with a proximal serrated margin. There
is a seta, with an expanded base, inserted on the distal margin.
The 3 pairs of legs ail hâve bimerous protopodites, obscurely articulated on the body and are
biramous. In the first leg (fig. 58, k) the endopodite is unimerous, approximately as long as the exopo-
dite. It terminâtes in 2 small, subequal, acute processes. The exopodite is bimerous, with the basal
article produced distally as a stout process. The distal article is subconical. The latéral margin
bears 2 minute, equispaced spinules.
The second leg (fig. 58, 1) has an obscurely bimerous endopodite, slightly shorter than the exo¬
podite. It terminâtes in 2 small, acute processes, one about twice as long as the other. The exopodite
resembles that of the first leg, except thaï the terminal article is slightly longer and lacks latéral spiniiles.
The third leg (fig. 58, m) has an obscurely bimerous endopodite, about 3/4 as long as the exo¬
podite. It terminâtes in 2 small, subequal, acute processes. The exopodite is bimerous, the terminal
article reduced to about half the length of the basal article. The basal article has a stout, long, seti-
form process at the latéral distal corner, about twice as big as the distal article. The protopodites and
endopodites of ail the legs hâve rows of ornamentating spinules.
The foiirth leg is a simple lobe on the posterior latéral margin of the first iirosomal segment
(fig. 58, f).
The caudal ramus (fig. 58, n) is flattened, subrectangular in outline, slightly longer than the anal
somite. There are 3 terminal lobes and an apical seta, which is about 1/3 as long as the ramus. There
is a short dorsal médial seta at about the distal fifth and a minute knob-Hke, articulated élément on
the latéral margin just proximal to the middlc. Rows of fine spinules are found on the surfaces of the
urosomal segments and the caudal ramus.
The absence of setae on the endopodites of the legs, although well-devclopcd ones occur on the
adult, furnish the strongest indication to us that this is a third copepodid.
Developmental Stages of Enleropsis capitulatus
(Figure 59)
The first nauplius is a lecithotrophic larva, with basic characteristics of cyclopoids of non-
feeding type (Dudley, 1966). We hâve found 5 naupliar stages and the usual 6 copepodids can be
assumed to occur from our sampling. It appears, however, that the mid-gut is well-formed when the
nauplius hatches. Yellow yolk is contained within the gut and large globules of purplish-red yolk
are associated with the mid-gut wall. It would seem that both the stomodaeum and proctodaeum
are patent at this stage. There is a very large red eye. There is no other significant pigmentation,
except some yellow globules.
The antennule (fig. 59, a) is uniramous, and functionally trimerous. The basal scleritic ring
lacks armature and ornamentation. The second article bears a single long seta at the distal margin
on the ventral face. Proximal to this on the ventral margin are 2 minute setules, set at the distal
thiid and fifth. Three rows of spinules are equally spaced on the médial surface. The terminal article
is about 1/3 longer than the second and shows littlc taper, but is rounded apically. On the apex there
is a long ventral seta, accompanied by a short aesthete and a dorsal seta about 1/3 as long. The
latter seta bears coarse hairs. Two rows of spinules, one at the proximal third and one the distal third
ornament the médial surface.
The antenna (fig. 59, b) is biramous, with obscurely bimerous protopodite, obscurely tetrame-
rous exopodite, and unimerous endopodite. The basal article of the protopodile lacks armature and
ornamentation. The second article has rows of spinules on the surface and shows a coalescence with
the basal article of the exopodite. On the latter there is a seta which represents the corresponding
article at the distal médial corner. The remaining articles of the exopodite are poorly articulated
with each other, indentations indicating the ingrédient éléments. The second and third articles each
has a long seta at the distal médial corner. The terminal article is rounded and has a short médial
seta and a long terminal seta. The endopodite is subrectangular and reaches to the end of the second
Source ; MNHN, Paris
Fig. 59. — Enleropais capilulalus : fînt nauplius (a-d) ; fourth Dauplius (e) ; fifth DaupJius (f) ; first copepodid (I, p, r) ;
E. roacoflenais : first copepodid (^k, m, o, q, s-u) : a, antennule ; b, antenne ; c, mandible ; d, subcuticular post*
mandibular appenda^s ; e, subcuticular postraandibular appendages and exuvial Unes ; f, subcuticular post-
mandibutar appendages and exuvial structure ; g, habitus, latéral ; h, rostrum, latéral ; i, antennule ; j, antenna ;
k, antenne, late stage ot first copepodid ; 1, antenna, late first copepodid ; m, mandible, early ; n, mandible, late
first copepodid ; o, maxillule ; p, maxillule ; q, maxilla ; r, maxilla ; s, first leg ; t, second leg ; u, urosome and
third legs. Legend : CR = caudal ramus ; ExLl = exuvial indication of leg 1 ; ExL2 = exuvial indication of
leg 2 ; ExMxl s exuvial indication of maxillule and maxilla ; Ex^Ll = exuvial indication ot protopodite of
leg 1 ; L1 = leg 1 ; L2 = leg 2 ; L3 = leg 3 ; Mxl = maxillule ; Mx2 = maxilla. Scales for g, s, t = 0.1 mm ;
other scales = 0.05 mm.
Source ; MtJHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
177
article of tte exopodite. It bears a seta in an cmargination at the rniddle of the médial margin and
2 long terminal setae. AU long setae of the appendage are coarsely plumose.
The mandible (fig. 59, c) is biramous, with obscurely bimerous protopodite, unimerous exopo-
ditc and bimerous endopodite. The basal article has a single row of spinules on the surface. This
article is fused with the exopodite. The latter, although unimerous has some indentations, possiblv
indicating a basic tetramerous condition. The ramus bears 3 long médial setae and 1 long terminal
seta. The endopodite is subrectangular and elongate, reaching to about the rniddle of the exopodite.
The basal article has a minute setule at the distal médial corner and a row of spinules. The second
article has a minute setule at the rniddle of the médial margin and 2 long setae on the truncate apex.
Ail long setae of the appendage bear coarse hairs.
Even at this first stage, the subcuticular indications of the postmandibular appendages are
well-formed (fig. 59, d). The maxillule (Mxl) and the maxilla (Mx2) are simple lobes ; there is no
maxilliped ; legs 1 and 2 (L1 and L2) hâve indications of the protopodite and rami, the exopodites with
3 pointed terminal projections and the endopodites with 2. The third leg (L3} is a simple lobe, with
a slightly undulate posterior margin. The second and third nauplü apparently do not differ mucb
from the condition described for the first nauplius. The legs and the gut are much the same. In
the exuviae of these latter second and third stages lines are présent for maxilla 1, maxilla 2, leg 1 and
leg 2. In the fourth nauplius the reddish-purple yolk has disappeared and is replaced by smaller
globules of yellowish-orange. In this nauplius the postmandibular appendages show some additions
(fig. 59, e). The maxillules (Mxl) and maxillae (Mx2) are much the same as before, in legs 1 and 2
(L1 and L2) the exopodites and the endopodites hâve each added 1 terminal projection. Ail such
éléments hâve elongated. The third legs (L3) show some posterior division into rami and the endopo¬
dite has 3 small pointed projections. Lines on exuviae indicate the margins of the maxillule (ExMxl)
and the maxilla and indicate the margins of the exopodites and endopodites of legs 1 and 2 (ExLl ;
ËxL2) and the endopodite of leg 3.
In the fifth nauplius (fig. 59, f) the condition of the exuvia, although changed some what from
the fourth, does not express the entire complément of developed subcuticular appendage rudiments.
The mouthparts are the same. The subcuticular pads of the first and second legs (Ll, L2) hâve very
well-developed protopodites and intercoxal lamellae. An additional terminal process has been added
to the endopodite. The third legs (L3) hâve added 3 terminal projections to the apex of the exopodite
in each. On the exuvia exopodites are indicated each by a line and a sac which encapsulâtes 3 or 4
internai éléments. The endopodites are indicated each by a line and 2 pointed sacs, one much larger
than the other. The larger sac encapsulâtes 3 internai éléments and the smaller 1. The caudal rami
(CR) are well indicated, with each a rectangular lobe with 2 latéral and 3 terminal setae. One of
the terminal setae on each ramus was already présent in the first nauplius.
The naupliar development in this species is very rapid, taking approximately 33 hours. In
the culture studied, hatching was observed at 10 a.m. on one day and by 10 a.m. the day following,
most nauplü were molting to the fifth stage and by 7 p.m. the first copepodid was seen.
In none of thé naupliar stages was any evidence of the maxilliped found.
In this species we hâve been able to study the first 3 copepodids, which were obtained by rearing
the nauplü. There were large numers of first copepodids, many second copepodids, but a single third
copepodid. The first copepodids were molting to second copepodids after about 24 hours. We did
not déterminé the duration of the second copepodid stage. Although the only other stage available
to us of this species is the adult, we feel we hâve been able to piece together the représentative life
history for the genus Enteropsis by adding in the fourth and fifth copepodids from E. superbus. Com¬
parative material, serving to corroborate our interprétations, is available to us also in the first and
second copepodids of E. roscoffensis which we were able to obtain through culturing methods. We
will describe the copepodids in the order of the succcessive stages, referring to each the available com¬
parative material.
In the first copepodid the body (fig. 59, g) is composed of the ccphalosome, metasome and uro-
some. The overall length is .434 mm {E. capiiulatus) or .392 mm [E. roscoffensis). By analogÿ
with the later developmental stages we consider the ccphalosome to include the portion of the body
Source : MNHN, Paris
178
PAUL L. ILLG AND PATRICIA L. DUDLEY
anterior to the segment of the first legs, although there is not an actual articulation so far. The remain-
der of the body is well articulated. The metasome has 2 leg*bearing segments, these legs functional.
The urosome consists of the segment bearing the rudimentary third legs, one legless segment and the
anal somite, with its terminal caudal rami.
There is a rostrum {fig. 59, h), bulbous in latéral outline, with several terminal protubérance.
The antennule (fig. 59, i) is 3*segmented, the terminal 2 articles about half as wide as the basal at its
greatest width. The armature is a follows : Article I — 3 setae, 1 setule, 1 spine ; II — 3 setae, 1 setule ;
III — 5 setae, 1 vestigial seta, 1 aesthete.
The antenna (figs. 59, j-1) at the early first copepodid stage is clearly biramous (fig. 59, j) but
very soon the internai tissue of the exopodite régresses and the cuticle crumples (figs. 59, k, 1). The
resuit is a uniramous appendage of 2 articles. The basal article, representing the protopodite, has
no armature or ornamentation. The terminal article, the endopodite, in E. roscojfensis (figs, 59, j, k)
has one terminal seta, which is slightly hooked. In E. capitulatus (fig. 59,1) the endopodite has 2 ter¬
minal hooked setae, one about twice as long as the other, and what appears to be a regressing seta.
The mandible (figs. 59, m, n) at the early first copepodid stage is biramous (E. roscojfensis
[fig. 59, m]) but very soon the tissue in both rami régresses. It would appear, however, that some
part of the exopodite will be retained, at least at the next stage. After the régression ail that remains
is a crumpled cuticular sac {E. capilulatus [fig. 59, n]).
The maxillule (figs. 59, o, p) in E. roscojfensis consists of a bilobed structure. The ventral,
more proximal lobe bears 3 short spinose articulated processes. The distal lobe, more medially placed,
bears 2 terminal setae, 1 twice as long as the other. Neither lobe is articulated. In E. capilulatus
(fig. 59, p) the appendage is similar in outline, but the ventral lobe appears to bear only 2 unarticulated
processes. The other lobe has 2 setae, but these are much longer and one is about 3/4 as long as the
other.
The maxilla (figs. 59, q, r) is a small domed projection, bearing an eccentrically placed setule ;
this is larger in E. roscojfensis (fig. 59, q) than in E. capilulatus (fig. 59, r). There is no maxilliped.
The first leg (fig. 59, s) is biramous, with unimerous protopodite and rami. The intercoxal
lamella is subquadrate and well developed. The protopodite has a small seta at the distal latéral
corner, near the articulations with the exopodite. The latter, flattened and about twice as long as
the endopodite, has 1 spine at the middle of the latéral margin, 1 spine at the distal latéral corner, 2 seti-
form éléments terminally and two setae distally on the médial margin. The endopodite has 1 latéral
seta, 2 terminal setae distally on the médial setae. The latéral terminal setiform élément of the exopo¬
dite would usually be replaced by a spine.
The second leg (fig. 59, t) is biramous, with unimerous protopodite and rami. The intercoxal
lamella is subquadrate. The armature of the protopodite and the exopodite is as in the first leg.
The endopodite is about 2/3 as long as the exopodite, has 1 distal latéral seta, 2 terminal setae, 2 distal
médial setae and 1 proximal médial seta.
The third legs (fig. 59, u) are non*functional, whith the basal articulation with the body unde-
veloped. Each leg consists of 2 sacs which encapsulate the developing rami. The endopodal portion
has 2 terminal spine-hke projections projections ; the exopodal portion has 2 spine-like projections
and an articulated seta. The projections encapsulate developing setae and spines. The second urosoraal
segment and the anal somite hâve no ornamentation. Each caudal ramus is about i/2 as long as the
anal somite. There are 5 setae on each caudal ramus, latthe proximal latéral third, 1 at the middle
of the dorsal surface and 3 terminal. The medial terminal seta is the longest and bears long coarse
hairs. Ail the other-setae are without hairs.
The second copepodid of E. roscojfensis (fig. 60) and E. capilulatus has a body composed of
cephalosome, somewhat more clearly delimited from the metasome than in the first copepodid, a
metasome of 3 leg-bearing segments and urosome of 2 segments and the anal somite with its terminal
caudal rami. The overall length is .392 mm în E. roscojfensis.
The rostrum (fig. 60, b) is long, basally articulated, the antennule (fig. 60, b) is 4-3egmented.
The armature is as follows : Article I — 5 setae, 2 spines ; II — 2 setae, 1 setule ; III — 3 setae ; IV —
7 setae, perhaps including an unrecognizable aesthete.
Source : Mt-JHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
179
The antenna (figs. 60, c, d) is uniramous and unimerous. There are 2 faint lines on the surface
at the proxinial third and distal third, but no actual articulations. The medial basal part of the appen-
dage has an indentation approximately at the position formerly occupied by the exopodite. There
is a single row of spinules on the surface of the appendage at the base of an apical elongate spinose
process.
The labrum (fig. 60, e) is a semicircular lobe, ornamented distaUy with fine hairs, and 2 setiform
processes project near the latéral extrêmes.
The mandibular remuant (figs. 60, f, h) consiste of a minute, cylindrical lobe, with 5 or 6 tiny
setac radiating from the tip. This lobe is inserted between the antenna and the maxillule, very close
to the base of the maxillule (fig. 60, h).
The maxillule (figs. 60, g-i) consists of a bilobed structure. The ventral more proximal lobe,
soinewhat smallcr proportionately than in the first copepodid, in some specimens is weakly articulated
at the base, in others is unarticulated (fig. 60, i). This lobe terminâtes in 2 spinose processes and there
is an additional third process laterally inserted on the lobe. The longer, more medial lobe is somewhat
fingershaped and bears 2 long terminal setae, 1 of these about 1 and 1/4 to 2 times longer than the
other.
The maxilla (figs. 52, j, k) is a low lobe with a single ventral spiniform, unarticulated projec¬
tion. A patch of spinules ornaments the lobe at the base of the projection. There is no maxilliped.
The first leg (fig. 60, 1) is biramous, with bimerous protopodite and rami. The subquadrate
intercoxal lamella is well developed. The coxopodite has neither armature nor ornamentation. The
basipodite has a small seta at the latéral margin and 2 rows of spinules at the surface near the articu¬
lation of the endopodite. The exopodite has 1 spine at the distal latéral corner of the basal article.
The second article has a spine at he distal latéral corner, 1 terminal spine and 1 terminal seta and 2
distal medial setae. The spines are very fiat, with broad serrated marginal flanges. The basal
article of the endopodite has no armature but has a row of spinules on its distal margin. The
second segment has 1 latéral seta, 2 terminal setae and 2 distal medial setae. This is the same complé¬
ment of armature as found in the first copepodid. In the later period of this stage one can see the
developing rami of the third copepodid under the cuticle (fig. 60, m).
The second leg (fig. 60, n) is biramous, with bimerous protopodite and rami. The intercoxal
lamella is subquadrate. The armature, segmentation and ornamentation are exactly as in the first
leg except that there is an additional seta on the medial distal corner of the basal article of the endo¬
podite.
In the late second copepodid one can see the leg of the third copepodid developing under the
cuticle (fig. 60, o).
The third leg (fig. 60, p) now metasomal and functional at this stage, is biramous, with bimerous
protopodite and unimerous rami. The intercoxal lamella is subquadrate, well developed. The coxo¬
podite has no armature or ornamentation. The basipodite has a short seta on the latéral margin.
The exopodite has 1 minute spinule at the middle of the latéral margin, a distal latéral spine, 1 terminal
spine and 1 terminal seta and 2 distal medial setae. The spines are greatly flattened and hâve broad
serrate flanges marginally. The endopodite has 1 distal latéral seta, 2 terminal setae, 2 distal medial
setae and 1 proximal medial seta. In the late second copepodid one can see the leg of the third cope¬
podid developing under the cuticle (fig. 60, q).
The first segment of the urosome bears the rudimentary fourth legs (fig. 60, r). Each consists
of a bilobed pad, with a single seta articulating on the face of the medial lobe. The second urosomal
segment has no armature or ornamentation. The unornamented anal somite bears only the caudal
rami. Each ramus is about 2/3 as long as the anal somite (fig. 60, s). There are 5 setae, 1 at the proxi¬
mal latéral third, 1 at the distal third of the dorsal surface and 2 setae, and 1 setule terminally. The
medial terminal seta is the longest. The 2 long terminal setae and the latéral seta bear coarse hair.
The remaining setae are unornamented.
The third copepodid of E. capituUitus (fig. 61, a) shows a drastic change of body configuration,
12*
Source ; Mh}HN, Paris
Fig. 60. — EtUeropai» roacoffenaia from Styela : second copepodid ; a, habitus, latéral ; b, rostrum and antennuie ; c,
antenna ; d, antenna -, e, labrum ; f, mandibular remuant ; g, mandibular remuant and maxillule ; fa, mandibular
remnant and maxillule ; i, maxillule ; j, maxilla ; k, maxilla ; 1, first leg ; m, outline of rami of iirst leg of late
second copepodid, showing developing leg ot tfaird copepodid ; n, second leg ; o, outline of endopodite of late second
copepodid showing developing endopodite of second leg of tfaird cope podid ; p, third leg; q, outline of rami of leg
of late second copepodid sfaowing developing leg of tfaird copepodid ; r, fourth leg : s, caudal ramus. Scales for a,
n, P « 0.1 mm ; other scales = 0.05 mm.
Source : MhNN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
181
Fie. 61. — Ejiteropais capitulalus : third copepodid (a-f) ; Enteropsia auperbua : fourth copepodid (g-p) : a, habit, laté¬
ral ; b, antennule ; c, antenna ; d, base and distal process of maxillule ; e, first leg ; f, caudal ramua ; g, habitus,
latéral ; h, antennule ; i, antenna ; j, maxillule ; k, maxilla ; 1, maxilia ; m, flrst leg ; n, second leg ; o, third leg ;
p, rudiment of fourth leg. Scale for g = 0.2 mm ; other scales = O.t mm.
no longer typically cyclopoid, but approaching the definitive eruciform habitus. There are slill only
3 pairs of legs well-formed. The segmentation is now much obscured, with indentations rather than
articulations spaced along the body. Because we had only a single spécimen we could not make out
ali the appendages. We describe what we could discern satisfactorily.
Source : MNHN, Paris
182
PAÜL L. ILLG AND PATRICIA h. DUDLEY
The antennule (fig. 61, b) is obscurely bimerous, with 4 terminal setae.
The antenna (fig. 61, c) is bimerous, with the much larger basal article ornamented with a row
of spinules. The terminal article tapers to a sharp point, evidently due to a coalescence of a termina)
seta. There is a subterminal setule, and a general ornamentation of spinules.
Only one lobe of the maxillule was made out (fig. 61, d). This is set on a base and ends in a
setiform process and an accompanying articulated seta.
The first leg (fig. 61, e) is biramous, with obscurely bimerous protopodite and unimerous rami.
The articulations of the rami with the protopodite are also obscure. The articles of the protopodite
hâve no armature but there is a general covering of spinules. The exopodite has 1 terminal, articulated
seta and a médial terminal spiniform projection. The endopodite ends in 2 spiniform projections.
There is no ornamentation on the rami. The présent form of the appendage was apparent wilhin
the cuticle of the previous copepodid stage when we observed this specimen in life (fig. 60, m).
The second and third legs are relatively similar to the first but probably hâve additional pro*
cesses, as seen within the cuticle of the preceding stage (figs. 60, o, q).
There are clear indications of 3 urosomal components, 2 segments and the anal somite bearing
the caudal rami. Each caudal ramus (fig. 61, f) consists of a truncate cône, with a single articulated
spiniform projection continuing its taper.
We cannot support our concept of the life history within the genus by a continuons sequence
of stages molted one into the next, but by comparison with related notodelphyids and from the anato-
mical graduations we are relatively sure the copepodids collected from the host represent the penul-
timate and the antepenultimate stages. As cyclopoids they should thus be the female fourth and
fifth copepodids.
As in the third copepodid of E. capitulatus the body of the female fourth copepodid of E. super¬
bus is eruciform (fig. 61, g), with the segmentation obscured. The overall length is 0.80 mm. There
are still only 3 pairs of legs developed, with indications of the fourth. The composition of the urosome
remains 2 segments and the anal somite. There is a general body covering of fine hairs and the ventral
surface of the urosome is covered with broken rows of spinules.
The antennule (fig. 61, b) is unimerous. There are 4 conspicuous setae and several setules,
the exact number obscured by the spinulose ornamentation.
The antenna (fig. 61, i) is bimerous, heavily sclerotized. The basal article is ornamented with
patches of spinules. The distal article tapers to a spiniform process. At the base of this there is an
articulated setule.
There is no mandible. The maxillule (fig. 61, j) is bilobed. A small subquadrangular lobe is
well articulated basally and bears 2 terminal setae. The main portion of the appendage is extended
as a long, finger-formed lobe, obscurely articulated on the base. It bears 2 subequal terminal arti¬
culated setae.
The maxilla (figs. 61, k, 1) is bimerous and heavily sclerotized. It has assumed essentially the
definitive form for the genus at this stage. The basal article has no armature or ornamentation but
there is a triangular projection at the distal médial margin. The terminal article forms a complicated
articulation with the base. It is so arranged as to direct a pointed beak-like process medially. Set
in a subcircular unsclerotized space on the ventral face is a stout seta. There is no maxilliped.
The first three legs are similar (figs. 61, m-o). Each of these legs is biramous, with poorly
articulated protopodites, of obscurely indicated components, and with the rami coalesced with the base.
The protopodite portion is ornamented with a general covering of hairs and rows of spinules. The
endopodite is a very short lobe ending in 2 spiniform projections. The exopodite ends in 1 major
spiniform projection surrounded basally by several shorter spiniform projections.
The fourth legs are represented only by a pair of lobes (fig. 61, p). The caudal rami are not
actually articulated on the anal somite and they diverge markedly (fig. 62, a). They are proportio-
nately long at this stage, each approximately equal in length to that portion of the urosome corres-
ponding to the anal somite. At a slight démarcation each continues as an equally long spiniform
process. The rami are covered with hairs and the spiniform projection has a short latéral spinuli-
form ornamentation along the proximal half of its length.
Source : Paris
ASCIDICOLIDAE AND SUBFAMII.IES
183
Fie. 62. — Enteropsis superbus : fourth copepodid (a) ; female fifth copepodid (b-h) ; immature adult female (i-j) :
a, caudal rami ; b, habitus, latéral -, c, antennule ; d, antenna ; e, maxillule ; f, maxilla ; g, flrst leg ; h, caudal
rami ; i, habitus, latéral ; j, caudal rami. Scales for b, i = 0.2 mm ; other scales = 0.1 mm.
No males were discovered by us, but it is very probable that Haligryps aculealus Aurivillius
is the male fourth copepodid of Enteropsis sphinx, as discussed above under that species.
The eruciform body of the female fifth copepodid of E. superbus (fig. 62, b) strongly resembles
the adult. The overall length is 1.54 mm. There are 4 pairs of legs. The composition of the urosome
appears to be 1 segment and the anal somite with its caudal rami. There is a general body covering
of fine hairs and the ventral surface of the urosome is ornamented with broken rows of spinules.
Source : MNHN, Paris
184
PAUL L. ILLG
PATRICIA L. DUDI-KY
The antennule (fig. 62, c) is uiiimerous. There are 5 setac and 2 obvions setules. There is
no ornamentation.
The antenna (fig. 62, d) is bimerous, heavily sclerotized. It is as strongly dcvcloped as in the
adult and has not changed in composition from the preceding copepodid.
There is no mandible.
The labrum is a simple, sclerotized lobe. The maxilhile (fig. 62, e) is bilobed. The longer lobe
has a clear articulation on only one surface, the shorter is clearly articulated. The latter is flattened
and terminâtes in 3 unarticulated, flattened processes, each triangular in outline. Two of these are
subequal, the third is much shorter. Spinules ornament the ventral surface. The longer lobe is sligh-
tly tapered, truncate, and bears 2 articulated, subequal, terminal setae. There are 2 subapical spi'
nules.
The maxilla (fig, 62, f) is like that of the fourth copepodid and the adult. There is no maxil-
liped.
The four pairs of legs (first leg, fig. 62, g) are much alike and just as in the adult. Each leg
is uniramous, bimerous. There is a general covering of fine hairs. The proximal article has heavy
sclerotizations. On the anterior surface there is a single short setule emerging from a pit in the sclero-
tization and a row of spinules ornaments the distal margin. The distal article has a complicated apex.
There is an apical médial tuberculated lobe, possibly a remnant of the endopodite. Beside this is a
Socket which received an articulated, heavily sclerotized terminal spine. The latter is possibly a
remnant of the exopodite.
There is no sign of fifth or sixth leg at this stage.
The caudal rami are not articulated on the anal somite and they do not diverge as markedly
as in the preceding copepodid stage. They are still relatively long, each just slightly shorter lhan
that portion of the urosome coiresponding to the anal somite. Beyond a slight démarcation at mid-
length each ramus continues as a tapered spiniform process. Each ramus is rather sparsely coated
with fine hairs.
The smallest adult of E. superbus obtaincd was 1.5 mm long, (fig. 62, i) almost the same size
as the fifth copepodid. Its general body aspect and the appendages are exactly those of the larger
specimens. Because of its smaller size, the caudal rami appear more salient than in large females
(fig. 62, j). Since the largest specimen found measured 8.2 mm, it is apparent that a great increase
in size can take place in this stadium without molting.
Developmental stages of a species of Bolryllophilus were dealt with in a study of development
of notodelphyids (Dudley, 1966). The first nauplius has the foUowing features. The antennule
is trimerous, the basal article is unarmed, the second has only 1 ventral seta, the terminal article has
2 apical setae and an aesthete. The antenna consiste of a bimerous protopodite, unimerous endopodite
and tetramerous exopodite. The protopodite has no setae. The endopodite has 1 medial seta and
2 terminal setae. Each of the 3 basal articles of the exopodite has 1 seta and the terminal article has
3 setae. The mandible consiste of a bimerous protopodite, bimerous endopodite and tetramerous
exopodite. The protopodite has no setae. The endopodite is about 2/3 as long as the exopodite
and bears 2 setules and 2 setae. Each article of the exopodite bears a single seta.
The copepodid of Botryllophilus features the emergence of the maxilliped of the adult and ail
the other head appendages are présent (Dudley, 1966, p. 160). The characters of these appendages
form highly distinctive features of the lineage of Botryllophilus and close relatives among the ascidi-
colid copepods.
Canu’s description and figures of features of the nauplius and the copepodids of “ Aplostoma ”
brevicauda Canu (1892, p. 88*90) hâve been confirmed (Dudley, 1966, p. 157, 160). In the first
nauplius the antennule is trimerous, the basal article unarmed, the second with 2 setae, the terminal
article with 2 apical setae. There is no terminal aesthete. The antenna has a bimerous protopodite,
unimerous endopodite and tetramerous exopodite. The protopodite lacks setae. The endopodite
has 1 medial setule, 2 terminal setae. Each of the 3 basal articles of the exopodite has 1 seta, the ter¬
minal article has 3 setae. The mandible has a bimerous protopodite, bimerous endopodite and tetra¬
merous exopodite. The protopodite lacks setae. The endopodite has 1 setule and 2 setae. The
Source : MNHN, Paris
ASCIDICOLIDAE AND SUBFAMILIES
185
endopodite is about 2/3 as long as the exopodite. Each of the articles of the exopodite bears 1 seta.
In the copepodid the haplostomin aiso shows the well developed maxilliped characteristic of
Botryüophilus. However, other of the mouthparts may show great réduction. The features of deve¬
lopment and the developing and adult appendages hâve been worked out by Ooishi (to be published).
OoiSHi & Illc (1977) invoked these developmental features to interpret the anatomies in the haplos¬
tomin généra. Added to the information on Botryllophilus from the work of Dudley (1966) and the
details compiled above, we hâve the basis for our interprétation of the anatomical details of the appen¬
dages in the ascidicolid subfamilies.
Source : MNHN, Paris
186
PAUL ILLG AND PATRICIA L. DUDLEY
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Distribué le 13 juin 1980
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