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Qy
Cifiol
Robert H. Palmer Estate
f
Bulletiii of the Miuenm of Comparative Zodkgy
AT HARVARD COLLEGE.
Vol. LI. No. 7.
*
Y
.t
THE CIDARIDAE.
X^
\^'
»^3
Bt Hubert Ltman Clark.
'/
With Eleven Platbs.
CAMBRIDGE, MASS., U. S. A. t
PRINTED FOR THE MUSEUM.
Dbcbmber, 1907.
^
No. 7. — The Cidaridae. By Hubert Ltman Claek.
Introduction.
The opening years of the present century have witnessed the pablica-
tion of an unusaal number of quarto volumes dealing with the morphol-
ogy and classification of the Echini. In each of these the Cidaridae
receive considerable attention, and many genera of that family, new
either in name or in contents, are proposed. As the dififerent writers
reveal wide divergence of opinion as to the relative importance of the
characters on which the classification of the Echini is based, the arrange-
ment of the Cidaridae differs to an unusual degree in these several re-
ports. Mortensen (:03)^ practically rejects previous classifications and
the principles on which they are based, and, ignoring the fossil forms,
to which his method is not applicable, recognizes thirteen genera and a
subgenus, defined wholly in terms of the pedicellariae, the spicules of the
pedicels, and occasionally the spines. It is only fair to state, however,
that the writer says frankly, these features are not " sufi&cient for definitive
diagnoses." He includes in his classificntion 42 species, and lists 12
others which he is unable to place satisfactorily because of lack of infor-
mation about the pedicellariae. Very soon after this volume appeared,
de Meijere's (:04)* valuable report on the "Siboga" Echini was pub-
lished. Unwilling to accept Mortensen's genera unreservedly, the writer
adopts the clumsy and unsatisfactory method of recognizing only a single
genus, Cidaris, and using Mortensen's names for subgenera. Later in
the same year Agassiz (:04)' in his report on the Panamic deep-sea
Echini, points out the weaknesses of Mortensen's method and the unsatis-
factory nature of his results, and emphasizes anew the great morpholog-
ical significance of the test (including the abactinal system). Two years
^ The Danish Ingolf-Expedition, 4, 1. Echinoidea. Part 1. Th. Mortensen.
Translated by Torben Lundbeck. 193 pp., 21 pis. Copenhagen, 1903.
3 Die Echinoidea der Siboga-Expedition. J. C. H. de Meijere. 252 pp., 23 pis.
Leiden, 1904.
' The Panamic Deep Sea Echini. Alexander Agassiz. Mem. Mus. Comp.
Zool., 31, 243 pp., 112 pis. '1904.
166 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY.
later Doderlein ( : 06),^ iu an effort to avoid some of the difficulties of
Mortensen's system, and yet to retain the valuable results of his work,
offers a classification of the recent Gidaridae, consisting of ten genera and
five subgenera, defined chiefly in terms of the pedicellariae. This classi-
fication, however, is quite different from any of its predecessors because,
while Ddderlein attempts to apply rigidly the recent International Code
of zoological nomenclature, his interpretation of certain perplexing cases
is quite different from either Mortensen's or Agassiz's. Finally Agassiz
and Clark ( : 07) ^ reject the proposed innovations of both Mortensen
and Ddderlein and offer considerable evidence in support of their view
that the pedicellariae of the Gidaridae are as unreliable for generic char-
acters as are the spines.
It is perfectly obvious, therefore, that the classification of the Gidaridae
is at the present time in a state of great confusion, and that some effort
should be made to reduce it to order and place it on a permanent basis.
Thanks to the great kindness of Mr. Agassiz, a very unusual amount of
material, both recent and fossil, has been accessible to me during the past
two years, and I have endeavored to find and formulate a natural arrange-
ment of the Gidaridae. Needless to say, Mr. Agassiz is not responsible
in any way for statements made or opinions expressed in the following
pages, but whatever value my results may have are due to his constant
sympathy and encouragement, and I wish here, in this inadequate way,
to express my thanks to him. I have also to thank Dr. Richard Rath-
bun for the privilege of examining the collection of Gidaridae in the
United States National Museum, and this proved to be of added interest
because it has recently been studied by Dr. Mortensen, who, in many
cases, left labels in his own hand, showing the views he held as to the
identification of the specimens. As my point of view differs fundamen-
tally from his, I desire to do him full justice, and the examination of a
collection, a large part of which has been named by him, was therefore
of special importance to me. Finally I may add that in the prepara-
tion of this report I have personally handled not less than 3,100 speci-
mens, representing 48 of the 60 recent species which appear to me to be
valid, and all of the 15 recent genera herein recognized.
^ Die Echinoiden der deutschen Tief see-Ezpeditioii. Ludwig Doderlein. 290 pp.,
42 pis. Jena, 1906.
^ Hawaiian and other Pacific Echini. The Gidaridae. Alexander Agassiz and
Hubert Lyman Glark. Mem. Mus. Comp. Zool., 34, 42 pp., 44 pis. 1907.
CLARE: THE GIDABIDAB. 167
Historical summary.
The first writer to use the name Cidarls for a genus of Echini was
Klein (1734), who, however, included all of the regular sea-urchins under
that name. Linn^ (1758) used the same name for a species of Echinus,
but Leske (1778) was the first writer subsequent to Klein who recog-
nized Cidaris as a genus. Only one of the 28 species which he includes
in the genus belongs in the family Cidaridae as understood to-day, and to
that one he gave the name jpapillcUa, Kow it is clear from both text
and figures that Leske intended to include under the name " Cidaris
jpapillata " all those regular Echini with the conspicuous interambulacral
tubercles of the Cidaridae. His ** species " is therefore a composite group,
including not only the now well-known European Dorocidaris papUlcUa,
but also Phyllacanthus tmpericUis and several species of the restricted
genus Cidaris, one of which appears to have been tribulaides Lamarck.
The next writer to deal with the classification of the Echini was Lamarck
('16)9 and he clearly indicates and defines the group which we now
call the Cidaridae. He called them " Turbans," under his genus Cidarites.
So far as the Cidaridae are concerned the name Cidarites is equivalent to
Leske's Cidaris papillata and is obviously a synonym of Cidaris. It
cannot be used, therefore, at the present time for any genus of animals.
Lamarck listed eleven species of " Turbans/' all but one of which were
recognized and described by Alexander Agassiz in 1872, in his classic
^'Eevision of the Echini." No attempt to subdivide the genus Cidaris
was made until 1835, when Brandt established the genus Phyllacanthus
for a supposedly new species, dvhia. He divided Lamarck's Cidarites
into two sections, A (including the species not in B and for which he
selected and named tribuloides Lam. as the type species) and B, Phyllar
canthus, with dttbia for the type, and including also imperialism hystrix^
geranioidesy and pistillaris. Later investigation made it plain that of
these four only imperidlis and pistillaris are congeneric with dvMa, and
the other two were therefore returned to Cidaris. In 1872 A. Agassiz
showed, however, that Lamarck's baculosa, verticillaia, and anmUi/era
had important features in common with dubia a,nd.imperialis and accord-
ingly placed them in Phyllacanthus. When Agassiz and Desor ('46)
considered the Cidaridae, they neglected Phyllacanthus, but established
Goniocidaris with geranioides for the type, and with it associated a " new "
species quoyi, which subsequently proved to be synonymous with
Lamarck's tuharia. In 1854 Desor suggested as genera of fossil Cida-
ridae, Bhabdocidaris, Diplocidaris, Porocidaris, and Leiocidaris, and in
168 bulletin: museum of comparative zoology.
1858 he described the fossil Eocidaris. The same year (1858)
Quenstedt named Polycidaris and Leptocidaris for fossil forms. la
1862 Cotteau described the remarkable fossil Orthocidaris, and the fol-
lowing year the equally interesting fossil Temnocidaris. In 1863,
A. Agassiz suggested the name Stephanocidaris for Liamarck's bispinosay
and Prionocidaris for pistiRaris, At the same time he proposed Chon-
drocidaris as a new genus for a notable species from the Hawaiian Islands,
and Gymnocidaris for metularia Lam. and a supposedly new species,
minor. He also proposed Orthocidaris and Temnocidaris as new genera
of recent Cidaridae, but later (1869) withdrew them as preoccupied by
Cotteau's fossil forms. At this later date he suggested Dorocidaris for a
new species, ahyssicolay associating with it affims Phil, and^^^^^'^o^a Leske.
With the last Lamarck's hystrix is synonymous, and consequently, as a
result of these various changes, there remained in Lamarck's genus
"Cidarites: Turbans" only the well-known West Indian species,
tribtdoides.
In the "Eevision of the Echini" (1872) A. Agassiz recognized only
six genera of the recent Cidaridae, as follows : —
Cidaris Klein, with 3 species. (Including Gymnocidaris A. Ag.)
Dorocidaris A. Agassiz, with 1 species. (Including Orthocidaris A. Ag.)
Fhyllacanthos Brandt, with 6 species. (Including Prionocidaris A. Ag., and
Chondrocidaris A. Ag.)
Stephanocidaris A. Agassiz, with 1 species.
Porocidaris Desor, with 1 species.
Goniocidaris Desor, with 3 species. (Including Temnocidaris A. Ag.)
This classification has been maintained by Agassiz ever since, without
any changes other than the addition of ten more species (1881, 1883,
1898) and the unique genus Centrocidaris (1904).
In 1877 Studer described Schleinitzia as a recent genus allied to
Phyllacanthus. In 1883 Pomel divided the "Cidarides" into three
subfamilies, the Gidariens, Goniocidariens, and Bhabdocidarieus. The
first contains four genera, including of Agassiz's six only Cidaris, which
is divided into five sections (subgenera 1 ) ; the second subfamily con-
tains four genera also, including Dorocidaris and Goniocidaris of Agassiz's
list ; the third contains seven genera, including the remaining three of
Agassiz, though Stephanocidaris is considered only a subgenus (?) of Phyl-
lacanthus. Although Pomel thus recognizes fifteen genera and six sub-
genera (1), his classification of the recent forms is essentially identical
with that of A. Agassiz. The new genera which he proposes are Tylo-
cidaris, Stereocidaris, Typocidaris, and Pleurocidaris, ail for fossil
CLARK: THE CIDARIDABL 169
forms. His proposed subgenera of Cidaris are, Plegiocidaris, Para-
cidaris, Procidaris, Poly cidaris, and Eucidaris. In 1884 Zittel proposed
Anaulocidaris for a fossil cidaroid, and in 1885 Ddderlein used the name
Discocidaris for some recent Japanese species. In 1887 Doderlein pub-
lished a classification of the Cidaridae, including the fossil as well as th«
recent forms. Of the 22 genera which he recognizes, 15 include only
fossil species. He rejects Stephanocidaris altogether, and uses Desor's
name Leiocidaris for Phyllacanthus. For some inexplicable reason he
considers Paroddaris sharreri A. Ag. as a living representative of
PomeFs genus Pleurocidaris. To another of Pomel's genera, Stereo-
cidaris, he assigns three recent Japanese species which he describes.
He proposes four new genera of fossil cidaroids, but only gives names to
three : Mikrocidaris, Triadocidai*is, and Miocidaris. In 1889 Duncan's
** Revision of the Genera ... of the Echinoidea " appeared, with a classi-
fication of the Cidaridae, which at first sight seems unique, but on exam-
ination proves to be novel only in the rank assigned to the different
groups. The writer divides the family into two sections, of which the
first contains four genera and one subgenus, and the second contains
two genera. For recent forms only the genus Cidaris, with a sub-
genus Goniocidaris, is allowed, but the heterogeneous nature of such a
genus is so far acknowledged that it is divided into seven '* divisions," of
which five contain the recent species. These five " divisions ^ with the
subgenus Goniocidaris correspond in name and contents to the genera
maintained by A. Agassiz. In 1902 Lambert proposed for certain fossil
and recent Cidaridae previously referred to Stereocidaris, the name
Phalacrocidaris, and in 1903 he suggested for some fossil species allied
to Phyllacanthus, the name Aulacocidaris.
In 1903 Mortensen entirely rearranged the recent species of the
family, uniting or separating them according to resemblances or differ--
ences in the lai^e globiferous pedicellariae. In this way he makes thir-
teen genera and a subgenus, and although he uses the names of the six
genera of A. Agassiz, the grouping of the species is wholly different from
that writer's. Mortensen's classification is as follows : —
Dorocidaris A. Ag. (emend. )i 4 species.
Tretocidaris, g. u., 3 species.
Stepbanocidaris A. Ag. (emend.), 3 species.
Schizocidaris, g. n., 1 species.
Cidaris Klein (emend.), 8 species.
Chondrocidaris A. Ag., 1 species.
Acanthocidaris, g. n., 1 species.
170 BULLETIN: MUSEUM OP COMPABATIVE ZOOLOGY.
Stereocidaris Fomel, 10 species.
Gbniocidaris Desor, 4 species and sal^nus Discocidaris Doderleiu, 3 species.
Fetalocidaris, g. n., 1 impedes.
Fhyllacanthas Brandt (emend.), 3 species.
Histocidaris, g. n*, 1 species.
Porocidaris Desor, 1 species and 1 variety*
Genus undetermined, 1% species.
Total, 56 species and 1 variety.
Of these 56 species, seven, aad the one variety^ are described for the first
time, but only one of them is figured. Unfortunat-ely Mortensen was
handicapped by lack of material and the apparent necessity of not de-
nuding even in part the specimens which were available, and as a conse-
quence his descriptions are, with one exception, incomplete, and in several
cases quite inadequate. Good photographs of his types would be a very
great help in recognising these supposedly new species.
In 1906 D(kierlein presents his classification of the recent Cidaridae,
the result of more than twenty years' study of the family. It is radically
difibrent from his earlier (1887) arrangement, not merely because no
reference is made to fossil forms, but because he endeavors to make use
of Mortensen's principles, which his own observations often contradict ^
and his judgment not infrequently condemns.^ This latest ariangement
of the family is as follows : -^
Cidaris Leske (syn. Dorocidaris A. Ag.), 4 species.
Tretocidaris Mortensen, 3 species.
Cidarites Lamarck (syn. Cidaris emend. Mortensen).
Subgenus Dorocidaris A. Ag., 4 species.
Gymnocidaris A. Ag., 3 species and 1 variety.
Stephanocidaris A. Ag., 5 species and 7 varieties.
Chondrocidaris A. Ag., 1 species.
Goniocidaris L. Agassiz et Desor.
Subgenus Goniocidaris s. str., 6 species.
Discocidaris Dod., 6 species.
Stereocidaris Fomel, li species.
Acauthocidaris Mortensen, 1 species.
Fhyllacanthas Brandt, 1 species and 5 varieties.
Histocidaris Mortensen, 2 species.
Forocidaris Desor, 1 species and 1 variety.
Genus undetermined, 6 species.
Total : 10 genera, 5 subgenera, 57 species, and 13 varieties.
^ Compare page 102, line 24, with page 106, lines 84-86 and page 109, lines 20-21.
* See p. 93 et uq.
CLARK: THE CIDARIDAE. 171
In 1907 A. Agassiz and Clark published descriptions and numerous
figures of nine new species of Cidaridae and instituted two new genera,
Anomocidaris and Aporocidaris. They also furnished much additional
information concerning Stephanocidaris, Ceutrocidaris, and Acanthoci-
daris and in regard to diversity of form in the pediceUariae of the
group.
Fundamental Principles for a Natural Classification.
Before attempting to set forth a revised classification of the Cidaridae,
if it is hoped to have it stable and generally acceptable, one ought to
make plain the principles on which it is based. These principles must
take into account not only the characters afforded by the specimens
themselves and the proper estimation of the relative value of these, but
also the selection of names for the genera and species held to be valid.
Fortunately there is coming to be more and more general agreement
among zoologists as to the principles which should govern in the selec-
tion of names, and the very general acceptance of the International Code
of Nomenclature, at least in its essentials, indicates clearly the approach
of the time when nomenclature will be fixed. In the following pages
adherence has been given to the rules of the International Code, but
whenever there has been room for difference of opinion as to the appli-
cation of those rules, that course has been followed which would cause
the least possible change from currently accepted names. Consequently
there are few changes from the names established or indorsed by A.
Agassiz in the "Kevision of the Echini" and almost universally used in
the last quarter of the nineteenth century. Unfortunately there is no
code by which can be determined the relative importance of the various
characters which distinguish the different species and genera of Echini.
Here each writer is thrown upon his own resources, and his proposed
classification will stand or fall according to the judgment he displays
in selecting stable and significant characters. The^ fiindamental diffi-
c ulty wit h the classification of Mortensen is that it is based almost
wholly upon the characters of the pediceUariae alone, and the history of
zoology shows again and again that a classification based on a single char-
acter, however suggestive it may be, is never reliable. The characters
afforded by the pediceUariae are important, but those organs are, like all
calcareous formations among echinoderms, liable to great diversity. It is
of no special importance in this connection whether the pediceUariae are
modified spines or not, the only point being whether, like the spines.
N
172 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
they show great individual variability. The evidence offered by A.
Agassiz and Clark (: 07) cannot be ignored or denied, and we are there-
fore forced to conclude that neither spines nor pedicellariae can be de-
pended on to furnish unvarying characters/^ On the other hand, Duncan
errs in placing his reliance almost exclusively on the test ^d in neglect-
ing the characters afforded by the spines and pedicellariae^ The classi-
fication used by A. Agassiz and the first one proposed by Ddderlein
(*87) show a judicious balancing of the various characters, and un-
doubtedly must serve as the basis for the natural classification we are
.seeking. (Ddderlein's latest arrangement of the Cidaridae does not ap-
peal to me as being well-balanced, for many excellent characters afforded
s by the test and spines are neglected or given little weight, while the in-
teresting diversities of the pedicellariae are permitted to outweigh all
N else.)( It seems to me there can be little question, either on a priori
grounds or as a result of observation, that the characters afforded by the
test are the most important in determining relationships among the
Cidaridae, and that those of the corona appear to be more reliable than
those of the abactinal system and actinostome.^ The size of the two
latter as compared with each other and with the size of the test are
useful factors in many cases, but there is considerable individual di-
versity in these proportions. This is true also of the arrangement of
the plates of the abactinal system, the position, form, and size of which
nevertheless often furnish characters of very great weight. The pri-
mary spines reveal obvious and tempting features, but these must be used
with caution, they are generally so variable., ' Curiously enough, how-
ever, in certain cases a character afforded by the primaries is very constant,
even though in nearly related species the same character may be very
variable:: The pedicellariae well repay careful examination and often re-
veal interesting and constant peculiarities, but, as has already been em-
phasized, they, like the spines, are subject to great individual diversity.
Indeed, it seems to be true that a spectes which has very variable spines
is likely to have equally variable pedicellariae. ^ The secondary, and even
the miliary, spines sometimes show characters of real value, although in
certain cases they are as variable as the primaries. The calcareous par-
ticles in the tube-feet seem to be so uniform in the family but so variable,
within these limits, in the individual that they afford no real help in
classification.
In the classification set forth in the following pages I have attempted
to place the proper value on each of the features of Cidaridean anatomy
mentioned above, and I have also taken into account geographical and
CLARK: THE CIDARIDAB. 173
bathymetrioal distribution. Even the suggestions of size, color, habitat,
and habits have not been ignored in the effort to learn the real interrela-
tionships of the species. At the suggestion of Mr. Agassiz, I have in-
cluded the genera of fossil Gidaridae, as well as the recent forms, in
order that the result may be as useful to palaeontologists as to z65logists,
and I have endeavored to give special consideration and due weight to
those characters upon which palaeontologists are obliged to rely I am
forced to the conclusion, however, that in most cases little value attaches
to the presence or absence of crenulation on the tubercles, to the straight
ness or sinuosity of the ambulaci-aj^pr to the amount of confluence of the
areolae. While these features are frequently very obvious in fossils, ex-
perience with large series of specimens shows that they are very variable
in individuals of the same species, and the most striking differences may
be due to the age or condition of the specimen. Far be it from me to
claim that the genera which I have adopted are all of equal value or that
they ought to be adopted as herein defined by all future writera. The
genera Phyllacanthus and Stereocidaris are notably unsatisfactory, and it
is quite likely that they will be entirely rearranged in the light of further
knowledge. Perhaps the same is true of Goniocidaris. But it is hoped
that the classification and nomenclature set forth in the following pages
may be a real step towards the ideal which we seek.
The Genera.
In attempting to apply the principles outlined above, it will be con-
venient to begin with those genera which are accepted by A. Agassiz,
D5derlein, Mortensen, and Pomel, and virtually by Duncan also. These
genera are : —
Cidaris Leske. Porocidaris Desor.
Goniocidaris L. Agassiz et Desor. Phyllacanthus Brandt.
Doderlein (:06) has reached the very disturbing conclusion that
papillata is the type of Cidaris, and that consequently Dorocidaris
A. Ag. is a synonym of Cidaris Leske. Acting on this belief, he has
introduced Lamarck's name Cidarites for Cidaris as commonly used, and
divides it into three subgenera, to one of which he applies the name
Dorocidaris A. Ag. In doing this, Doderlein overlooks the very im-
portant fact that Leske's Cidaris papillata is a composite group which
was first broken up by Lamarck. It includes at least three species, —
irnperialisy which Brandt removed to Phyllacanthus; pajnllcUa, which
174 BULLETIN: MUSEUM OP COMPAUATIVB ZOOLOGY.
A. Agassiz removed to Dorocidaris ; and tribuloides (or possibly mdidaria ;
it matters little which), which remains thus as the type of Cidaris.
Moreover Brandt, who was the first writer to subdivide Cidaris, dis-
tinctly states that tribiUoides is the type of Cidaris s. str., and as
** first reviser " he undoubtedly had the right to select the type. There
is therefore no need of upsetting a number of familiar names and caus-
ing considerable confusion by insisting on papilkUa as the type of
Cidaris. Indeed, if we are to discuss this question, imperialia has a
better claim than papillata to be the type of Cidaris, for it is undoubt-
edly the first species Leske names, though he has it confused with
papillata under the varietal name major. In resurrecting Lamarck's
name Cidarites, which is clearly a substitute for, and synonym of, Cidaris,
DoJerlein violates the old principle " once a synonym, always a syno-
nym," and certainly if Dorocidaris A. Ag. is a synonym of Cidaris
Leske, as Doderlein says, it cannot be used for a subgenus of Cidarites.
It is surprising that so good a zoologist as Doderlein should have com-
mitted two such errors. Since Doderlein's Cidarites equals Cidaris
Mortensen and his "Cidaris" is equivalent to Dorocidaris A Ag.,
the latter can be added to our list of accepted genera, which will also
include several genera of recent Cidaridae adopted by Mortensen,
Doderlein, and Agassiz and Clark, as follows : —
Dorocidaris A. Agassiz. Stereocidaris Pomel.
Cliondrocidaris A. Agassiz. Acanthocidaris Mortensen*
We may also add five genera of fossil Cidaridae, accepted by Pomel,
Doderlein, and Duncan, regarding which there can be little question : —
Orthocidaris Cotteau. Poljcidaris Quenstedt.
Temaocidaris Cotteau. Diplocidaris Desor.
Tetracidaris Cotteau.
The following genera are fully described and figured by A. Agassiz or
by A. Agassiz and Clark, and their validity is not likely to be questioned,
with the possible exception of Staph anocidaris, which some zoologists
may not wish to separate from Phyllacanthus. So far as the evidence
goes, however, it is fully entitled to recognition.
Stephanocidaris A. Agassiz. Aporocidaris A. Agassiz and Clark.
Centrocidaris A. Agassiz. Auomocidaris A. Agassiz and Clark.
There still remain no less than 21 genera and several subgenera of
Cidaridae which have been proposed and are entitled to consideration.
CLARK: THE CIDARIDAE. 175
To these I have given special attention, but the great majority do not
seem to me to be based .on sufficiently reliable or tangible characters to
warrant their recognition. The following list includes them all, with
my opinion as to the proper status of each ; those which appear to me
to be worthy of use are indicated by black-faced type.
Khabdocidaris Desor : not distinguishable from PhyUacanthus.
«/ Leiocidaris Desor : " " « " ^
Eocidaris Desor : not distinguishable from Cidaris, or else from Archaeocidaris,
according to what species is considered the type. It is true that the first
species mentioned by Desor (keyserlingi) does not agree with the diagnosis
of the genus, but since Doderlein ('87) has definitely selected that species as
the genotype, Eocidaris becomes a synonym of Cidaris.
Leptocidaris Quenstedt : very probably not one of the Cidaridae.'
Gymnocidaris A. Ag. : not distinguishable from Cidaris.
Prionocidaris A. Ag. : " " ** Phyllacanthus.
Schleinitzia Studer :
Tylocidaris Pomel : apparently a valid genus, though allied to Cidaris.
Typocidaris Pomel : not clearly distinguishable, and too near Cidaris and Doro-
cidaris.
Pleurocidaris Pomel : not distinguishable from Phyllacanthus.
Plesriocidaris
Paracidaris
Pomel : hopelessly indistinguishable.
Doderlein: not distinguishable from each other and too near
Cidaris and Dorocidaris.
Procidaris
Eucidaris
Anaulocidaris Zittel : not distinguishable from Cidaris.
Discocidaris Doderlein : not " " Goniocidaris.
Mikrocidaris
Triadocidaris
Miocidaris
Phalacrocidaris * Lambert : ?
Aulacocidaris * Lambert : ?
Tretocidaxis Mortensen : see below.
Schizocidaris Mortensen : not worthy of separation from Goniocidaris.
Petalocidaris Mortensen: "
Histocidaris Mortensen : " " " " " Porocidaris.
^ I have been unable to see the original descriptions or any figures of these two
genera, as the papers in which they are published are not to be found in either
Cambridge or Boston. But Aulacocidaris (Lambert, 1903; Bull Soc. Hist. Nat.
Savoie, (2) VIII, p. 222) is evidently closely related to Phyllacanthus and is proba-
bly not distinguishable, while Phalacrocidaris (Lambert, 1902; Mem. Soc. Geol.
France, Pal. IX, fasc. Ill, Mem. 24, p. 27) is based on Doderlein's living species of
Stereocidaris from Japan, but includes a number of fossil forms. As Stereocidaris
is itself only distinguishable with great difficulty, it is very unlikely that Phalcuro-
cidaris is tenable.
176 BULLETIN : MQSEUM OF COMPAKATIVE ZOOLOGT.
The genus Dorocidaria is difficult to separate, on the one hand, from
Cidaris, and on the other from Stereocidaris, but is particularly close to
the latter, and it is almost impossible to draw a sharp line between
them. Moreover, it contains a rather heterogeneous lot of species.
One of these, D. micans Mortensen^ seems to be quite unique, and I think
it may well be made the type of a new genus for which I would suggest
the name Calooidaris. The remaining species fall naturally into three
groups, distinguished from each other by their abactinal systems, spines,
pedicellariae, and distribution. I see no objection to recognizing these
group^i as genera, and such a course has some obvious advantages. A
typical Dorocidaris such as papillata has the abactinal system irregularly
angular and often indistinctly defined, and the globiferous pedicellariae
have a conspicuous end-tooth on each valve. But other species have
the abactinal system circular or pentagonal and sharply defined, and
some of the globiferous pedicellariae are often more or less like those
of Cidaris. To this group Z>. hartletti A. Ag.. belongs, and as Morten-
sen has made that species the type of a new genus, Tretocidaris, that
name must attach itself to this section of Dorocidaris, even though
few of the species have the remarkable pedicellariae which Morteusen
considers the distinguishing character of the genus. Finally, a group
of three small species, characterized by their thickened secondaries,
globiferous pedicellariae without end-tooth on the valves, sparsely
tubercled abactinal system, and antarctic or subantarctic distribution,
may be conveniently designated as Austarocidaris. The table on the
opposite page gives the genera adopted in the present paper, with their
authors, the year in which they were proposed, and the type-species of
each. The number of recent species in each, which seem to me valid, is
also indicated.
The number of fossil specimens to which specific names have been
given is in tha vicinity of 200 ; of these, D5derlein lists 135, but there
is reason to believe that many of these represent different ages or indi-
vidual forms of single species, and it is not unfair to assume that the
number of extinct species actually known to science does not exceed
the number pi species now living. The following key will bring out the
^ obvious if not the most important characters by which the 21 genera
here recognized may be distinguished. It is hoped that such a key may
be of use to palaeontologists as well as to zoologists. The dimensions
are given in millimeters, and the horizontal diameter of the denuded test
(al>breviated for convenience to ** h. d.")jjbaken at the ambitus, is used
as the unit for determining the relative proportions of the various
I
CLARK: THE CIDABIDAE.
177
Geniu.
Cidaris . . .
Phyllacanthufl .
Goniocidaris
Diplocidaris . .
Porocidaris . .
Polycidaris . .
Orthocidaris
Temnocidaris .
Stephanocidaris
Chondrocidaris
Dorocidaris . .
Tetracidaris
Tjlocidaris . .
Stereocidaris .
Tretocidaris . .
Acanthocidaris
Centrocidaris .
Anomocidaris .
Aporocidaris
Calocidaris . .
Austrocidaris .
Author.
Tear.
Leske . .
• •
1778
Brandt .
• •
1836
L. Agassiz
et
Desor .
• •
1846
Desor . .
• •
1854
Desor . .
• •
1864
Quenstedt
1868
Cotteau .
1862
Cotteau .
1862
A. Agassiz
1868
A. Agassiz
1863
A. Agassiz
1869
Cotteau .
1872
Pomel .
1883
Pomel .
1883
Mortensen
1903
Mortensen
1003
A. Agassiz
1904
A. Agassiz
and
Clark .
1907
A. Agassiz
and
Clark .
1907
gen. noy.
1907
gen. noy.
1907
Type-spMiea.
tribuloides Lamarck .
imperialis Lamarck . . .
geranioides Lamarck . . .
gigantea Agassiz ....
yeronensis Desor, but of re
cent species, purpurata Wyv
Thomson ....
multiceps Quenstedt .
inermis A. Gras . . .
magnifica Cotteau . .
bispinosa Lamarck . .
gigantea A. Agassiz .
abyssicola A. Agassiz
reynesi Cotteau . . .
gibberula L. Agassiz et Desor
cretosa Man tell, but of recent
species, grandis Doderlein
bartletti A. Agassiz . . .
curyatispinis Bell ....
doederleini A. Agassiz . .
japonica Doderlein
milleri A. Agassiz . .
micans Mortensen . .
canaliculata A Agassiz
Number
of Recent
Species.
3
5
7
6
3
1
5
9
9
3
1
8
1
3
21 genera and 60 recent species.
parts. The other abbreviations used are self-explanatory. The " vertical
diameter '* means the vertical distance from the margin of the abactinal
system, at the end of an ambulacrum, to the lowest part (usually several
millimeters distant from the edge of the actinostome) of the same am-
bulacrum, measured with a pair of dividers. When the measurement
from the centre of the abactinal system is normally very different from
this, special reference is made to the fact. In all cases maximum
measurements are used for comparison ; thus, when it is said that the
''abactinal system equals .40 h. d./' what is meant is that the greatest
diameter of the abactinal system (it is not always circular) equals .40
of the greatest diameter of the test. '' Primary spines about equal to
h. d." means that the longest primary is about equal to the greatest di-
ameter of its own test. The relative position of the pores of a pair is
indicated as " horizontal " or " oblique," according to whether a line
drawn outward from tbe tubercle on the margin of the median am-
bulacral area, at right angles to that margin, passes above both pores or
VOL. LI. — NO. 7 12
178 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
through the outer pore of the pair. Uuless otherwise noted, the colora
given are those of dried Museum specimens.
In using this, and all other keys given, it should be constantly borne
in mind that the younger the individual, the less will it show generic
and specific characters ; in proportions, number of coronal plates, and of
secondary and miliary spines, arrangement of the abactinal system, form
of the primary spines, and color, the young are often quite different
from the adults. They can only be identified with certainty on com-
parison with other specimens, old, young, and intermediate, and usually,
for very young specimens, it is necessary to know the place and means
of collection. On the other hand, unusually large specimens often have
the abactinal system and actinostome relatively smaller than in speci-
mens of more moderate size. Variations of five per cent or more, on
either side of any mean given, may therefore be expected. The keys
are all based on supposedly normal, mature specimens, the age being
estimated by the presence and size of the genital openings, the appearance
of the primary spines and abactinal system, and to some extent by the
size. Although the radial plates of the abactinal system are not con-
nected with any sort of light-detecting or visual organs, they have been
so generally called *' ocular " (ocellar in German and ocellaires in French)
plates that the name is here retained, as preferable to the alternative
term "radial," which Duncan uses, but which is not really quite so
distinctive.
Key to the Genera.
Genen marked with an * have no living lepreoentatiTea.
Pores horizontal or nearly so, distant (space between the two of a pair
evidently exceeding diameter of a pore) ; surface of interval flat, or
with a groove connecting pores, never elevated. (Individuals in
which this feature is obscure are characterized by stout or more
or less thorny spines, 1.5-2 6 h. d. [if less than 1.6 h. d., coronal plates
very few, 6 or 6], and unsunken and, even actinally, quite distinct
! areolae.) Jlecent species exclusively Indo-Pacific«.
With pores in 4 more or less regular vertical series in each poriferous
' zone.
With 4 vertical series of coronal plates in each interradius from ac-
tinostome to ambitus *Tetraa'dar{s l^^
With only 2 series of coronal plates in each interradius . . . .fDiplocidariB /^i .
With pores in only 2 vertical series in each poriferous zone.
Ambulacral and interambulacral plates with more or less numerous,
nearly circular pits, irregularly scattered *TemnQcidaris fU
I £ 3
CLARK : THE CIDARIDAE. 179
Ambulacral and interambolacral plates without such pits.
Abactinal system of numerous thin plates, with yery large anal sys-
' tern around which ocular and genital plates form a single narrow
ring ; genitals, except madrepore, much wider than high, often
twice as wide ; oculars nearly as high ; collar of primaries spotted
with white ; lowest actinal primaries with very wide collar and
11 short thick cap of outer layer of spine, flattened curved, and
somewhat serrate at tip, when fully developed . . . Stephanoctdaris /^2L '
I Abactinal system not as above; collar of primaries not white-
spotted ; actinal primaries not provided with a distinct cap.
Median interambulacral area less than .30 of interambulacrum
PhyUacanthusJSi^-
Median interambulacral area more than .30 of interambulacrum, \
densely covered with minute tubercles Chondrocidaris / Y ''
Pores nearer together, usually more or less oblique, often separated by an
elevation and never yoked together by a groove.
All primary tubercles large, smooth, and imperforate ..... *Tyhcidaris I h\' • ^ '
Primary tubercles, at least at ambitus, perforate. j.
Ambulacra more than half as wide as interambulacra .... Centvcidari* ^ / ^ t
Ambulacra not half so wide as interambulacra, usually mnch less. '' ' ' . '~ *- •
Coronal plates with areolae so small their diameter is less than one-
quarter horizontal length of plate and only about one-half verti-
cal height *Orthocidari8lCO
Coronal plates with areolae which occupy a large proportion of plate.
Ambulacra broad, .35-. 45 of interambulacra, with median area
correspondingly wide, sometimes sunken and more or less
bare; median space of interambulacra, especially along verti-
cal, and inner portion of horizontal sutures, sunken deeper than
areolae, especially at angles, and more or less bare ; in some
species, however, miliary tubercles cover so much of inner half
of each coronal plate that parts of vertical suture are concealed
and only short, bare, horizontal furrows are visible, and even
these may be only faintly indicated. Coronal plates numer-
ous in proportion to h. d., 6-11. Primaries always rough and
more or less thorny or prickly, often flaring at tip . . GonioctdarisjO S -
Ambulacra less than .85 of interambulacra, or, if more than that,
primaries not thorny.
Coronal plates numerous and narrow, 9-15, with areolae merg-
ing into each other throughout the whole series . . *Polycidaris Jif^-
Coronal plates rarely more than 9, areolae at ambitus and abac-
tinally never merged together.
Primary spines long, 2-3 h. d.,not at all thorny or prickly, broad
and somewhat depressed at base, tapering much but gradu-
ally, often slightly curved, and with aconspicuous light-col-
ored or spotted collar, one-fifth or more of the length ^cantAoacfons 2 ^ '^ .
Primary spines very diverse, but never as above.
Only tridentate or, more rarely, bidentate, pedicellariae
present, but these abundant and often very large
(2-6 mm. high) Pwocidaria *- ^ 7
• . .
1 2^^'
180 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGT*
GlobiferouB pedicellariae present, but often onlj small ones.
Abactinal system rery large (.60-.70 h. d.); ambulacral
plates lew, generally less than 80; poriferous zones
not at all sunken; secondary and miliary spines
alike, cylindrical and more or less club^haped; no
tridentate pedicellariae present AjwrpcidariM 2, -
Abactinal system less than .60 h. d. ; ambulacral plates
more than 40 (except, of course, in young indi-
Tiduals).
Abactinal surface conspicuously bare, with no primary
spines or well-developed tubercles or areolae much
above ambitus; no tridentate pedicellariae pres-
ent Anomocidaris 2. .
Abactinal surface not so conspicuously bare ; at least two
primary spines well above ambitus in each interradius.
Areolae little or not at all sunkeii ; actinostome gen-
erally larger than abactinal system, which is usu-
ally .40-.46 h. d. ; median ambulacral area with
only a single marginal series of tubercles, though
there are usually other smaller, scattered tubercles
between, and these may form 1-6 vertical series. .
Primaries .65-1.60 h. d. but commonly about equal
to li. d., rather stout, usually blunt; secondaries
broad, flat, and truncate Cidaris \ * j
Areolae more or less deeply sunken ; actinostome usu- ' -
ally smaller than abactinal system; median am-
bulacral area usually with a double marginal series
of tubercles, inner much smaller. Primaries 1-8
h. d. ; secondaries diverse.
8mall (25-40 mm. h. d.) ; abactinal system with few,
generally less than 200, tubercles ; secondaries,
especially ambulacral, rounded, thickened, and
more or less club-shaped ; no tridentate pedicel-
lariae ; large globiferous pedicellariae with no
end-tooth on the valves. Subantarctic, north to
about 35° S Austrocidans T / ^
Larger (30-70 mm ) ; abactinal system with more
numerous tubercles; secondaries flat and thin,
and usually narrow. Tridentate pedicellariae
usually present and large globiferous, often with
an end-tooth on the valves. Northern hemi-
sphere, seldom south of the equator.
Abactinal system sharply defined, more or less dis-
tinctly circular or pentagonal in outline ; ocular
plates with outer margin convex or straight,
little notched by ambulacra. Some or all of
large globiferous pedicellariae, if not like small
ones, have curved valves, large terminal open-
ing, and no end-tooth, as in Cidaris . . Tretocidaris Z^co
I ? 5 u r :
GLABK: the CIDA.RIDAE. 181
Abactinal system not verj sharply defined, rather
irreguUr in outlinei with re-entering angles,
between ocular and genital plates; oculars
with more or less concave outer margin or
deeply notched by ambulacra. Large glob-
iferous pedicellariae never as in Cidaris.
Abactinal system thick and solid, more or less
elevated; genital and ocular plates with
more or less convex surfaces, thickly and
uniformly covered with tubercles of approxi-
mately equal size ; ambulacral secondaries
usually larger than those on genital, ocular,
and uppermost coronal plates and often
conspicuously contrasted with tbem. Cor-
onal plates few, 4-7, rarely 8 or 9 ; upper-
most 1 or 2 or even 8 without primary
spines. Primaries never smooth, but pro-
vided with longitudinal rows of granules, or
with ridges, 1 or more of which may be de-
rated to form conspicuous, though delicate,
buttress-like ** wings " along basal half of
spine; if these buttress-like *' wings" are
not present, terminal portion of spine often
more or less fluted and flaring. Globiferoos
pedicellariae, both large and small, com-
monly lack conspicuous end-tooth . . Stereocidaris (\ / 9
Abactinal system flat, usually not uniformly: ' '-^^ z^' ■.•,
covered witli tubercles, some of which are ' ' '
also larger than others ; ambulacral second-
aries not noticeably contrasted with others
abactinally. Coronal plates 6-8, rarely 9,
all (except usually uppermost 1, or rarelj
2) with primary spines. Primaries some-
times perfectly smooth, never with " wings,"
and seldom with flaring tip. Globiferous
pedicellariae, both large and small, com-
monly with conspicuous end-tooth.
Median ambulacral area .65 of ambulacrum
in width; primaries shining as though
polished, white more or less shaded with
greenish or pink, or both Calocidaris
'•'• .
/ /
Median ambulacral area less than .50 of< am-
bulacrum; primaries never shining as
though polished Dorocidaris '/ f' /
^ 7 '^
The above key gives little clue to the relationships of the genera with
each other, and a natural arrangement must necessarily be largely a
matter of speculation. There can be little question that Cidaris is
182
BULLETIN : MUSEUM OF COMPA&ATIVE ZOOLOGY.
nearest to the ancestral form and the centre from which the different
genera have come. Whether Tylocidaris represents a more primitive
type, because of its imperforate tubercles* is an open question. The
other genera fall rather naturally into three groups, which correspond
to the three '* sous-tribus ** of Pomel, but the lines between these groups
are not clear enough to warrant any recognition of subfamilies. The
following table indicates these three groups, and in the succeeding pages
the genera will be taken up in the order here given^ which indicates
roughly their possible relationships.
Phyllacanthas.
Chondrocidaris.
Diplocidaris.
Tetracidaris.
Stephanocidaris.
Temnocidaris.
Tylocidaiis.
Cidaris.
Goniocidaris.
Folycidaris.
Ortliocidaris.
Dorocidaris.
Tretocidaris.
Calocidaris.
Austrocidaris.
Centrocidaris.
Aporocidaris.
Stereocidaris.
Anomocidaris.
Acantliocidaris.
Porocidaris.
Diafimoses of the Genera, and the Recent Species.
In view of the large number of recent Cidaridae described since the
publication of A. Agassiz's ** Challenger " Echini, a complete revision
of the family will not be without value, so, to the extended diagnoses of
the genera here accepted, artificial keys to the recent species contained
in each are added, with a few remarks concerning each one, and a refer-
ence to a good figure when one has been published. Three apparently
new species, represented in the Museum of Comparative Zoology by
several specimens each, are also described and figured. Ko attempt at a
synonymy is made, since the " Ee vision of the Echini " gives all that is
needed in that line for the species long enough known to have been
burdened with many names. Eeferences to published figures are given
for every species which has ever been figured, and photographs are
added of all species which have never been figured hitherto, except only
Dorocidaris nuda, of which no specimen has been available.
CLARK: THE CIDARIDAE. * 183
TYLOCIDARIS.
Tylocidaris Pomei; 1883, Class. Meth. Gen. Ech., p. 109.
Plate 1064, flgrfl. 1-7, PaL Franc. Terr. Cx«t«, 7, Cotteau, 1862.
Test small or of moderate size, much as in Dorocidaris ; coronal plates 5-8 ;
areolae distinctly sunken, sometimes large, and tending to merge together yertically ;
primary tubercles large, smooth, and imperforate ; median interambulacrai and am-
bulacral areas and poriferous zones as in Cidaris or Dorocidaris ; pores large, close
together, slightly oblique. Abactinal system of moderate size, about .4&-.60 h. d.
Actinostome somewhat smaller than abactinal system. Primary spines very stout,
club- or acorn-shaped. Secondaries and pedicellariae 1
It is difficult to know how much weight can wisely be laid on the absence of
perforations in the tubercles, but it is a character never shown in perfect tubercles
of living Cidaridae. On the whole, the combination of imperforate tubercles with
the curious short, stout spines makes the genus easy to recognize. Doderlein
('87) lists four species,^ from the Cretaceous of Europe.
CIDARIS.
Cidari$ Leske, 1778. Add. Nat. Disp. Ech., p. 17.
Test moderately high ; vertical diameter usually about .60 h. d. (ranges from
.60-.75) ; thick and solid (in mettdaria, thickness of an ambulacral plate at ambitus
is about .56 of its liorizontal length) ; coronal plates &-9 (sometimes 10, very rarely
11); areolae not sunken but tending to merge together actinally; median inter-
ambulacrai area little or not at all sunken, more or less uniformly tuberculated ;
sutural lines often not visible at all ; ambulacra .20-.35 of interambulacra in width ;
poriferous zones little sunken ; median ambulacral area with a single conspicuous
marginal series of tubercles and 1-3 (rarely none, or in large specimens 4 or 6)
irregular vertical series of much smaller ones between ; sutural lines more or less
obscured and not conspicuously sunken ; pores oblique, with distance between two
of same pair about equal to diameter of a pore and with surface of interval more
or less elevated. Abactinal system .30-.50 h. d. Actinostome .40-.55 h. d., usually
larger than abactinal system, sometimes half as large again. Primary spines about
equal to h. d. (range from .65-1.60 h. d.), stout, cylindrical or terete, usually blunt,
slightly rough but not thorny, covered with longitudinal series of granules which
are usually low and rounded but may be conspicuous and sharp ; actinai primaries-
not peculiar, little or not at all flattened; ends rounded and generally fluted.
Secondary spines flat, truncate, rather broad and not tapering towards tip, which
may indeed be widened. Pedicellariae of 3 kinds present as a rule, but tridentate
may be wanting, or rarely large globiferous ones fail ; latter have curved valves,
large terminal opening, and no end-tooth.
This genus is one of the most easily recognized of the family, although some of
the individuals with long spines approach quite nearly in appearance to Tretoci"
184 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
darit afimis. Indeed it is possible that some of the specimens of C. iribuloides
with long, tapering spines, which have been collected in the West Indies, are
really hybrids between that species and affiiUs, but there is no proof that this is
the case. There are only 3 valid recent species of Gidaris, and they are quite
sharply distinct from each other. The form which Doderlein ('87) described
under the name galapagensU is not constantly distinct from tkouarsii and must be
referred to that species. All of the living species are littoral forms, and rarely
occur at a greater depth than 50 fths., but are found along nearly all tropical
and subtropical coasts, ^umerous fossil species from Tertiary, Cretaceous,
Jurassic, Triassic, and even Permian strata have been named. The following key
tothe recent species is based on the examination of 845 specimens representing
all three.
Key to the Species.
Small, h. d. rarely exceeding 30 mm. ; median areas .45-.60 of ambulacral
width, usuaJly bare and often sunken ; abactinal system .45-.50 h. d. ;
genital plates always clearly in contact with each other; coronal
plates 6 or 6, rarely 7 metularia
Medium to large, h. d. 30-70 mm.; median areas seldom more than .40 of
ambulacral width, always provided with miliary tubercles ; abactinal
system usually less than .45 h. d. ; some or all of genital plates sepa-
rated in mature specimens ; coronal plates 7-10, rarely 11.
Median interambulacral area more than .10 h. d. ; abactinal system
usually over .40 h. d. ; small spines olive, fawn-color, or red-
brown, with tips usually darker tribtdoides
Median interambulacral area less than .10 h. d. ; abactinal system
usually less than .40 h. d. ; small spines dark red-brown, purple,
or nearly black, with tips not noticeably darker thouarsii
Cidaris metularia.
Cidarites metularia Lamarck, 1816, Anim. s. Vert;, 3, p. 66.
Cidaris metularia Blainville, 1830, Zoophytes : Diet. Sci. Nat., 9, p. 212.
Plat« Ig, fig. 1, Bev. Bch., A. Agrassiz, 1873.
Although having a far more extensive range than either of the others, this species
shows much less diversity in the length and form of the primary spines ; they are
generally about .80 h. d. and are rarely if ever 1.20 h. d. The stalks of the large
globiferous pedicellariae have a well-developed " limb.** The colors are generally
brighter than in the lai^er species, and the cross-banding of the primaries is
usually very distinct; some Hawaiian specimens are very red, more or less
marked with yellowish or reddish white. The geographical range is from Cape
of Good Hope, northward on the east coast of Africa into the Red Sea (including
Madagascar, Mauritius, Bourbon, and the Seychelles), thence eastward along the
CLARK: THE CIDARIDAE. 185
sonthem coast of Asia with the adjoining islands, through the East Indian archi-
pelego and out into the Pacific, as far as the Solomon, Fiji, and Hawaiian Islands.
Curiously enough, tnetularia does not seem to reach either Japan (except the Liu-
kiu Islands) or Australia. The only difference that can be detected between
Mauritian and Hawaiian specimens is that, in the latter, the median ambulao-
ral area is somewhat broader and flatter, but the difference is very slight and
inconstant.
Cidaris tribuloides.
Cidarites tribuloides Lamarck, 1816, Anim. s. Vert., 3, p. 66.
Cidaris tribrdaides Agassiz, 1835, Prodrome, p. 188.
Plate Id, Plate 2, figs. 1-3, Ber. Bcb., A. Agassiz, 187S.
Little need be said further in regard to this well-known species, save that the
primary spines are frequently cross-banded, especially in young specimens, and in
old specimens are almost always more or less encrusted with colonies of Bryozoa,
and similar foreign material. The relative length and thickness of the primaries
differ to a remarkable degree in specimens from different localities. The general
appearance of specimens from the Cape Verde Islands is thus strikingly different
from that of the ordinary West Indian form. On the other hand, many of the
specimens dredged in the West Indies, by the " Blake," have the primaries so long
and slender that there is a noticeable superficial resemblance to Tretocidaris affinis,
Conuecting forms between the extremes are, however, common. The stalks of the
large globiferous pedicellariae have no " limb." The geographical range is con»
fined to the Atlantic Ocean, from the Bermudas and Azores on the north to Brazil,
the Gape Verde Islands, and Cape Palmas on the south. In the Museum are
several old tests without spines, which are almost certainly this species, labelled
" Mer Rouge," but a mistake in labels is always possible, and these have doubtless
been mixed at some time with West Indian specimens. There is also a very small
(5 mm. h. d.) but perfect specimen from "Sl^ 26' S. andeS** 5' W., 57fths.,"
collected by the " Hassler." If there has been no mistake, this would indicate a
remarkable southern range. Small specimens .from Ascension Island, Atlantic
Ocean, in the collection of the National Museum, like those collected by the
" Challenger " at Bahia, and Fernando Noronha, Brazil, have verticillate swellings
on the primaries, but are not otherwise peculiar. .
Cidaris thouarsii.
Cidaris thouarsii Agassiz and Desor, 1846, Cat. Rais. Ann. Sci. Nat. (3) 6, p. 326.
Plate 10, Jap. Seelgel, Doderleln, 1887.
This is the well-known substitute for tribuloides on the west coast of America.
It is easily distinguished from that species by the color and other characters men-
tioned above. Its range is comparatively limited, however, as it is not known from
south of the equator (save in the Galapagos) nor from north of the Gulf of Cali-
186 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
fomia. After a careful comparison of numerous excellent specimens from Mexico,
Panama, and the Galapagos, it is clear that there is no constant character by which
C* galapageiuU Doderlciu ('B7) can be distinguished from thonanii. Specimens
from the Galapagos usually have the short and very stout spines figured, by Dd-
derlein, and apparently do not have tridentate pedicellariae, but some Galapagos
specimens have long, slender, tapering spines, while some from the coast of Mexico
have spines like those of most Galapagos specimens; and individuals from
Panama occasionally kck the tridentate pedicellariae. DOderlein's present opinion
(1906) seams to be that galafttgemU should be regarded as a variety of ihouarsii^
PHYLLACANTHU8.
PhyllacanthuM Brandt, 1836, Prodrome, p. 267.
Test much as in Cidaris but thinner ; thickness of an ambulacral plate only .30-
.40 of its boricontal length ; coronal plates vary greatly in different species, ranging
from 6 to 11 ; areolae not at all sunken and usually quite distinct even near actino-
atome ; median interambulacral area not deeply sunken, though it may be bare and
sutural lines distinct; ambulacra .20-.40 of interambulacra in width; poriferous
zones little sunken ; median ambulacral area generally with a double series of
marginal tubercles (inner much smaller than outer) and 1-4 additional, more or
less regular, vertical series between ; but when ambulacra are very narrow, median
area may be as in Cidaris, and wlien very broad, median area may be bare and
without additional tubercles; pores nearly or quite horizontal; distance between
two usually much greater than diameter of pore ; surface of interval flat or hori-
zontally grooved, 80 that pores are connected by a furrow. Abactinal system much
as in Cidaris. Actinostome varies greatly in different species. Primary spines
exceedingly variable, usually 1.6-3 h. d. and quite stout; actinal primaries either
as in Cidaris or somewhat flattened, thick and truncated at tip, slightly curved and
somewhat serrate. Secondary spines flat, but length and breadth very variable.
Liarge globiferous pedicellariae variable in form and often entirely lacking, but
tridentate and small globiferous pedicellariae are generally present.
Large specimens of this genus are easily recognized, but small ones are often
pnzzling. In very young specimens the pores are arranged much as in Cidaris,
and this condition has not wholly disappeared in specimens 20 mm. in diameter ;
in thomasii even the largest specimens do not have the interval between the pores
perfectly flat. On the whole the genus is difficult to characterize properly and the
recent species are not well defined. But the combination of characters mentioned
in the key to genera is unlike that of any other cidaroid, and with proper care a
specimen of Phyllacanthus over 30 mm. h. d. should be recognized without great
difficulty. No other genus, however, shows so great diversity in the length and
form of the spines, and, as might be supposed, the pedicellariae are also very varia-
ble. There seem to be only five valid species in this genus, but it must be con-
fessed that the confusion of hacnlosa-^iih annulifera^ and the latter with Stepha-
nocidaris bispinosa, has led to a most unfortunate situation, and there can be no
doubt that a careful revision of the.genus based upon abundant material from the
CLARK: THE CIDARIDAE. 187
Ked Sea, Mauritius, the East Indies, and Australia is sadly needed. In the light
of such material I believe that additional species will be recognized, and it is quite
possible that the genus will need to be divided. For the present, however, I see
no better course than to let the genus stand as it is. It seems to be generally
agreed that Studer's ('80) Schleiniizia crennlaris is a Phyllacanthus, probably
annulifera ; while the observations of Doderlein ('87 and : 03) and de Meijere
(: 04) show that Fh. duhia Brandt ('35) and parvi^pina Woods ('80) are appar-
ently synonyms of imperialis. The species designated awttralis by Ramsey ('85)
is apparently baculosa and Rhabdocidaris recens Troschel is clearly annulifera.
All of the recent species are littoral and are confined t(J the Indo-Pacific region,
but many extinct species have been described from Tertiary, Cretaceous, and
Jurassic stmta of Europe and America. The following key to the living species
is based on the examination of only 118 specimens, but each of the live species is
represented by at least four examples.
Key to the Species.
Ambulacra very broad, .40 interambulacra or more; median area broad,
sunken and bare ; median interambulacral area also sunken and bare ;
primaries seldom exceed h. d., provided with several whorls of vertical
plate-like projections or fiat, blunt tliorns verticHlcUa
Ambulacra less than .40 interambulacra; median ambulacral and inter-
ambulacral areas not conspicuously sunken and bare.
Collar of primary spines without spots or longitudinal lines of deep red
or purple.
Coronal plates 6-6 (rarely 7) ; abactinal system small (.30-.40 h. d.) ;
actinostome large (.60-.55 h. d.) imperialis
Coronal plates 6-9 (rarely 10) ; abactinal system nearly equals or often
exceeds .actinostome.
Primary spines stout 1.5-2.6 h. d., terete, slightly swollen above
collar, smooth, or with granules arranged in longitudinal series, '
becoming ridges near tip; no conspicuous thorns or pro-
jections thomasii
Primary spines not as above, sometimes flattened at base, usually
with conspicuous thorns ; collar smooth, reddish or purplish,
unspotted annulifera
Collar of primary spines with noticeable spots of purple or deep red,
arranged in longitudinal rows and sometimes merged into lines . . baculosa
Phyllacanthus verticillata.
Cidarites verticillata Lamarck, 1816, Anim. s. Vert., 3, p. 66.
Phyllacanthus verticillata A. Agassiz, 1872, Rev. Ech.. pt. 2, p. 151.
Plate If, fig:. 3, BeT. Ech., A. Agassiz, 1873.
This well-known and unmistakable species reaches a diameter of 35-40 mm.
The general coloration is dark brown and green, with the shades lighter in young
188 BULLETIN : MUSEUM OF COMPABATIVE ZOOLOGY.
individuals. It ranges throughout the East Indian region, north to Anima
Oshima in the Liu-kiu Ishtnds and southward along the east coast of Australia ;
it has been reported from as far west as Mauritius and Zanzibar, and as far east
as the Fiji, Samoan, and Hawaiian Islands. Its occurrence in the latter group
seems doubtful, as it was not represented in the very extensive collections made
by the ^ Albatross " in 1903. Although ordinarily a littoral form, a specimen
firom a depth of 647 ftbs. is reported by de Meijere (:04).
Phyllacanthus ixnperialis.
Cidarites imperialU Lamarck, 1816, Anim. 8. Vert., 3, p. 64.
Phyllacanthus imperialU Brandt, 1836, Prodrome, p. 268.
Plate If, fist. 9» 6, 7, Bev. Bch., A. ApiMis, 1873. Plate 58, figs. 3» 4, Semon'a
gesam. Bch., Duderleln, 1903.
This is another well-known species, dark brown or purple in color, and of large
size (up to 75 mm. h. d). Some or all of the primary spines frequently have
two or more narrow rings of light color near the distal end. The geographical
range of this species is from the Red Sea and Zanzibar to and throughout tbe
East Indies and along the east coast of Australia. I am in doubt as to whether
the varieties recognized by Dbderlein are really sufficiently constant to be worthy
of names.
Phyllacanthus thomasii.
Phyllacanthus Thomasii A. Agassiz and Clark, 1907, Haw. Pac. Ech. : Cid., p. 16.
Plates SI7-80, Haw. Pac Kch. Cid., A. AgtMslm and Clark, 1907.
This handsome species reaches as large a size as the preceding, and the long,
tapering, stout spines give it a very characteristic appearance. In the largest
specimens tbe small spines and test are dark reddish-brown, but in specimens
•SCf-.^O mm. h. d., the ambulacra and their spines are very pale brown, in sharp
contrast to the interambulacra and abactinal system. At all ages the primary
spines are salmon-colored, thickly spotted with white, and having a brown collar,
but in old specimens they are more or less encrusted with foreign material which
conceals the true color, and the collar is much wider and darker than in the
young. This species is known only from the vicinity of the Hawaiian Islands.
Phyllacanthus annulif era.
Cidarites annulifera Lamarck, 1816, Anim. s. Vert, 3, p. 67.
Phyllacanthus annulifera A. Agassiz, 1872, Rev. Ech., pt. 1, p. 160.
Plate 58, figfs. 5-11, Semon's se«am. Ech., Duderlein, 1903.
This, species has been so persistently confused, on the one hand with the much
rarer Stephanocidaris bispinosa (q. v.), and on the other with an East Indian
variety of the much commoner Ph* baculosa, that the limits of its geographical
claek: the cidaeidab. 189
range are really unknown. There appears to be a variety of baeulosa common in
the East Indies, in which the primaries are cross-banded as in this species, and
this form has been confused with annuli/era. Now if de Loriol ('73) and Mor-
tensen ( : 03) were correct, it would be clear that Lamarck's annulifera is this
variety of baculosa, and in that case the present species should be called lutkeni^
as de Loriol clearly figures and describes it under that name. Mortensen says
he has examined Lamarck's type and it is baeulosa, but A. Agassiz examined all
of Lamarck's types some forty years ago and satisfied himself that the present
species is Lamarck's annulifera. In a disagreement such as this it is obvious
that the earlier investigation is the one least liable to error, for there had been
considerably less time for a chance confusion of labels or specimens. Both
de Loriol and Mortensen apparently overlook the fact that A. Agassiz examined
Lamarck's types in Paris and that there has never been the slightest reason for
supposing that he made any mistake, in associating Lamarck's name with this
species. Until it can be shown that such a mistake was made, the name it has
borne so long should be retained for this species. So far as we now know, it is
an Australian and East Indian form, and does not occur in the Red Sea or along
the African coast. The Museum of Comparative Zoology has two fine specimens
from the Gulf of Siam, received from the Copenhagen Museum. They were
collected by Mortensen, and labelled by him " Stephanocidaris bispinosa." The
species is apparently nearly as variable as baculosa, both in coloration and in
the form of the primary spines ; in some cases the secondaries are green and
the primaries cross-banded with purple and green, but in other specimens the
secondaries are pale brown and the primaries are dull with less distinct markings.
The secondaries usually (perhaps always?) have a median longitudinal stripe,
darker than the ground color. The primaries are frequently flattened and wid-
ened at the base, tapering to the tip and quite thorny, much as in Stephenocida-
ris, but they are often nearly cylindrical with few thorns. I am not satisfied
that the varieties recognized by Doderlein are sufliciently constant to warrant
their recognition by name.
Phyllaoanthus baculosa.
Cidarites bcundosa Lamarck, 1816, Anim. s. Vert., 3, p. 65.
Phyllacanthui baculosa A. Agassiz, 1872, Rev. Ech., pt 1., p. 160.
Flato If, figs. 4, 5, Bev. Sksh., A. Agrassis, 1873. Plate 60, fiirs. 1-^, Semon's
sesam. Ech., Doderlein, 1903.
Common, variable, and widely distributed as is this much-discussed and per-
plexing species, its true characters and the limits of their variability are still little
understood. It seems useless in the present state of our knowledge to attempt
to recognize varieties, and we can only say that with all the diversity of coloration
and of primary spines, the deep red or purple spots on the collar of the primaries
is an obvious character almost always present. It is true de Loriol ('83) and de
Meijere (:04) have described specimens with a narrow unspotted collar, but it is
190 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
quite possible that these are not really baculosa. It is interesting to note that the
purple spots on the collar may merge together, not only longitudinally so as to
form parallel vertical lines, but also diagoually, so that the collar appears check-
ered with light-colored, diamond-shaped spots. These spots are occasioaally
rounded, and then the color shows some resemblance to that of the primaries of
Stephanocidaris. Further evidence of the close relationship existing between that
genus and baculota is found in the abactiual system of the latter, where some or
all of the ocular plates may be broadly in contact with the anal system. The
coronal plates are 8-10 or even 11 in the krgest specimens (64 mm. h. d.)» and
the color is brownish-red or purplish, but is quite variable. The geographical
range appears to coincide with that of imperials. A remarkably handsome spine
of a Phyllacanthus, quite unlike any of baculosa which I have seen, in the Mnseum
collection from '* He Bourbon," inclines me to Mortenscn's (:03) yiew that the
identity of baculosa and pMUaris is still open to question. If it is not doubtful,
this species ought to be called by the latter name, as it has precedence in La-
marck's work. Doderlein (:06), on the strength of Loven's ('87J descrip-
tion and figure, adopts the Linnean specific name cidarts for this species, quite
overlooking Loven's own statement (p. 146) : " Be that as it may, the species
name : Cidaris L., left to its fate by the author himself, is to be laid aside as
without validity, though of some historical interest." In the collection of the
United States National Museum there is a notably fine specimen (No. 14,032)
from the Boniu Islands, labelled " annulifera " ; the secondaries are very long,
with a deep brown longitudinal stripe, and the collar of the primaries has some
indistinct white spots as well as the characteristic deep purplish-red dots. It is
quite possible this is an undescribed species. In the same collection is a large
series of specimens from Aden (No. 21,459), which have been labelled by Dr.
Mortensen " Cidaris metularia " ; the primaries are remarkably short and stout,
much as in Cidaris, and as Mortensen did not clean an ambulacrum, it is not
strange that he failed to see the very characteristic poriferous zones. But it is
hard to understand how he overlooked the conspicuous purple spots on the collar
of the primaries.
CHONDROCIDARIS.
Chondfocidaris A. Agassiz, 1863, Bull. M. C. Z., 1, p. 18.
Test much as in Phyllacanthus, but densely covered with minute tubercles bearing
miliary spines and small globiferous pedicellarlae ; median interanibulacral area
very broad, generally .36-.40 of interambulaerum, nearly flat; ambulacra narrow,
only .20-.25 of interambulacra ; median ambulacral area covered with about eight
vertical series of tubercles, of which the marginal ones are slightly larger ; pores
horizontal, widely separated, connected by a groove. Abactinal system .35-. 40 h. d.,
with ocular plates entirely excluded from anal system ; genitals broadly in contact.
Actinostome about equal to abactinal system. Primary spines stout, nearly cylin-
drical, sometimes slightly tapering, about equal to, or somewhat exceeding h. d.,
provided with stout, blunt, thorny projections, and often near the tip with longi-
CLARK: THE CIDARIDAK 191
tudinal lamellae. Secondary spines few, flat, and blunt, confined to scrobicular
circles .and margins of ambulacra; latter very slender. Large globiferous pedi-
cellariae usually wanting ; tridentate infrequent, with slender straight valves ; Bmall
globiferous abundant, on very short stalks, with prominent end-tooth on valves.
This is a monotypic genus, closely related to the preceding but easQy distin-
guished at a glance by the peculiarly bare appearance of both ^ambulacra and
interambulacra.
Chondrocidaris gigaoitea.
Chondroddaris gigantea A. Agassiz, 1863, Bull. M. C. Z., 1, p. 18.
Plate la. Rev. Kch., A. Agassiz, 1873.
This species is of special interest because of its huge size (up to 95 mm. h. d.),
its remarkable primary spines, and its very broad median interambulacral areas
densely covered with minute miliaries. The color is brown of some shade, the
countless miliaries with a distinctly greenish-yellow cast. It is a curious fact
that really young specimens of gigantea have not yet been taken, none in the
collections of either the National Museum or the Museum of Comparative Zool-
ogy being less than 75 mm. h. d., and de Loriol's ('83) specimen, the smallest
yet recorded, was more than half that size. Most of the known specimens are
from the Hawaiian Islands, but it is also reported from Lifu, Loyalty Islands
(Bell, '99), and Mauritius (de Loriol, *83). The latter is remarkable for having
only 5 coronal plates, while Hawaiian specimens have 8-10. The record of this
species from the Lepar Islands, given by Sluiter ('95), is said by de Meijere
(:04) to rest only on spines of " C, (Stephanoeidaris) bispinosaJ*
DIPLOCIDARIS.
Diplocidaris Desor, 1854, Syn. Ech. foss., p. 45.
Plate 1, fig. 6, Syn. Kch. foss., Desor, 1854.
Test much as in Phyllacanthus ; coronal plates 7-8 ; areolae little or not at all
sunken, sometimes merging together actinally ; median interambulacral area not
sunken or bare, but with few, scattered tubercles ; ambulacra narrow, less than .25
of interambulacra in width; poriferous zones more or less sunken; median am-
bulacral area narrow, with usually only a single marginal row of tubercles, the
intervening bare space sometimes conspicuous ; pores nearly horizontal and widely
separated, in vertically very narrow plates, which are so crowded that they have
the appearance of having slipped on each other laterally, so that the pores are
apparently in 4 vertical series in each zone. Abactinal system small, with large,
usually angular, genital and small ocular plates. Actinostome larger thnn abacti-
nal system. Primary spines very stout, with longitudinal series of low tubercles
which tend to merge into ridges near the tip. Secondaries and pedicellariae ?
This genus is very diflferent from any living Cidaridae in the arrangement of the
pores, but in all other respects it is strikingly like Phyllacanthus, especially some
192 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
specimens of Ph. imperalis. The crowding of the pores is very similar to wliat
occurs in Asterias and other starfishes, where the ambulacral plates are so crushed
together that a straight, single row of pores is forced into such a zigzag arrang^e-
ment that it has the appearance of two parallel series. There is no reason to con-
sider the arrangement in Diplocidaris as anything other than a highly specialized
condition. It seems strange that it is not found in any living species of Cidaridae.
Doderlein ('87) lists 5 species of this genus, all from the Jurassic strata of
Europe.
TETBACIDABIS.
Tetracidaris Cotteau, 1872, Rev. et Mag. ZooL (2), 23, p. 44&
Plate 29, fifftf. 7-11, Bev. et Mag. ZooL {2), S3, Cottean, 1878.
Test larg^, circular at ambitus, somewhat depressed ; coronal plates very numer-
ous (16 in each complete vertical series), arranged in 4 series in each interradius
from actinostome to above ambitus and thence in a double series to the abactinal
system ; areolae somewhat sunken ; median interambulacral areas narrow and with
few miliaries ; ambulacra narrow, only about .20 of interambulacra in width ; porif-
erous zones little sunken ; median ambulacral area nearly bare, with a marginal
series of tubercles and a few scattered miliaries ; pores nearly horizontal, widely
separated, and crowded into a double series in each zone, much as in Diplocidaris.
Abactinal system '* large." Actinostome ? Primary spines rather slender, nearly
cylindrical, somewhat ridged. Secondaries and pedicellariae ?
In the arrangement of the pores this species is intermediate between Diplo-
cidaris and Phyllacanthus, but it is not in any sense a connecting link between
these genera. It may be regarded as a specialized offshoot of the Diplocidaris
branch. Duncan ('89) thinks it may be related to Astropyga, and there is some
reason for thinking it is not genetically connected with the Cidaridae at all. Only
one species is known, reynesi, from the European Cretaceous strata.
STBPHANOCIDARIS.
Stephanocidaris A. Agassiz, 1863, Bull. M. C. Z., 1, p. 18.
Test, ambulacra, interambulacra, and relative proportions as in Phyllacanthus,
but coronal plates 6-8; abactinal system .40'.45 b. d. and actinostome either larger
or smaller; anal system ^large and made up of numerous plates (in a specimen
42 mm. h. d. there are over 50 anal plates, and in a young specimen 12 mm.
h. d. there are 25) ; all plates of abactinal system relatively thin ; genital plates
much wider than high, except madrepore, which is much larger than others;
ocular plates wide and high, 4-sided, outer side convex, inner usually correspond-
ingly concave; genitals and oculars together form a ring around anal system of
CLARK : THE CIDARIDAE. 193
nearly uniform width except where madrepore juts in.^ Primary spines some-
what flattened near base, cons picBQUsI y t^nmy; collar wide, greenish, reddish
or dark with conspicuous white spots; in young specimens tliese white spots
project as granules, but in mature specimens, coUaf is smooth; actinal prim-
aries slightly curved, with a very wide collar, often more than half their length,
and provided with a distinct cap of outer layer of spine ; this cap is truncate, thick,
and somewhat serrate. Large globif erous pedicellarlae are wanting in all available
specimens.
Although there can be no doubt of the close relationship between this genus
and Fhyllacanthus, the discovery of a new species of Stephanocidaris in the
Hawaiian Islands, of which numerous specimens are available for study, shows
how clearly justified A. Agassiz ('63) was in making Cidaritet bispinota Lamarck
the type of a separate genus. The characters shown by the primary spines are
exhibited in specimens only 12 mm. h. d., and even in these specimens the genital
plates are widely separated ; it is not, however, until a diameter of over 20 mm. has
been reached that the remarkable character of the abactinal system becomes ap-
parent ^be three species here recognized are confined to the central and eastern
portions of the Indo-Pacific region. The following key is based on the examination
of 106 specimens of the first and third species.
1 It is worth noting that in a young Stephanocidaris 6 mm. in diameter, the
ocular plates are all excluded from the periproct, except that of the left posterior
ambulacrum, which barely touches an anal plate ; in a specimen 7 mm. in diam-
eter, the left posterior ocular is clearly in contact with the anal system and the
right posterior ocular barely touches it ; in a specimen 12 mm. in diameter, the two
posterior, and the left anterior oculars are all clearly in contact with, while the
odd anterior ocular barely touches, the periproct ; in another specimen of the same
size, all the oculars except the right anterior are clearly included ; in a specimen
14 mm. in diameter, and in all larger ones, all the oculars are broadly in contact with
the anal system. It seems to be true, therefore, of Stephanocidaris that the oculars
of the bivium come into contact witli the periproct before those of the trivium do
and of the latter the right anterior ocular is the last to enter. Examination of a
series of young Cidaris tribtUoides shows that the same course is followed in that
species, except that in one specimen the odd anterior ocular was excluded, while
the right anterior was no longer so. These facts are strikingly in accord with the
condition often found in Tretocidaris and always in Acanthocidaris, where the
right anterior ocular is the only one excluded. And I way add that in Arbacia
nigra and spattdigera, in adult specimens of which the posterior oculars, and often
the left anterior, are in contact with the anal system, the same course of entrance
of the oculars is followed ; and while I have found a very few specimens in which
the odd anterior ocular is also insert, I have yet to find an Arbacia in which
the right anterior ocular is not excluded. The reason for this condition is not
clear.
VOL. LI. — NO. 7 13
194 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
Key to the
Primary spines not red ; interarabalacral secondaries whitish with a longi-
tadinal green stripe.
Primaries stout, less than 2 h. d. ; ambulacral secondaries dark green ;
abactinal system larger than actinostome bispinosa
Primaries slender, 2-3 h. d. ; ambulacral secondaries like those of inter-
ambulacra ; abactinal system smaller than actinostome .... glandtdosa
Primary spines red (in young, sometimes greenish) with more or less in-
distinct cross-bands of white ; secondary spines reddish or brownish,
not longitudinally striped ; abactinal system not larger than actino-
stome hawaiienM
Btephanocidaris bispinosa.
Cidariten bispinosa Lamarck, 1816, Anim. s. Vert., 3, p. 67.
Stepkanocidaris bispinosa A. Agassiz, 1872, Rev. £ch., pt. 1, p. 160.
Plate If, fiff, 1, Bev. Kch., A. AgassiXy 1873.
It would be amusing were it not irritating to note bow entirely recent writers
have ignored Agassiz's ('73) description and figure of this beautiful and appar-
ently very rare species. The trouble appears to date from de Loriol's ('73) fig-
ure, which is certainly not bispinosa, but is probably P. annuHfera, in one of its
various color phases ; his figure of lutkeui is certainly annulifera, while bis figure
of annulifera appears to be daculosa. Koehler ('95) evidently refers to the same
form of Maculosa under the name annulifera^ while his Stepkanocidaris bispinosa
is probably true annulifera. Bedford (1900) has apparently identified correctly
his specimens of annulifera, so far as can be determined from his figures. D6-
derlein ('-OS) and Mortensen (^04) entirely ignore Agassiz's description, or else
intimate that the description is inadequate because it fails to apply to tlieir speci-
mens. As a matter of fact, it seems clear that neither of them lias seen a speci-
men of the real bispinosa, but since they call the variety of baculosa with banded
primaries annulifera, they are obliged to do something with their specimens of real
annulifera, and so they suppose them to be St, bispinosa, Agassiz's description
and figures to the contrary notwithstanding ! Their lead is sonSewhat reluctantly
followed by de Meijere (*04), who is unwilling to ignore Agassiz's statements ;
but he, too, records Ph. annulifera as St, bispinosa. This species reaches a
diameter of over 50 mm. Authentic specimens are known only from Australia
and Malacca.
Stephanocidaris glandulosa.
Cidaris (Cidaris) glandtdosa de Meijere, 1904, Siboga-Exp. Ech., p. 13.
Plate 1, figs. 6, 6, Slboga-Bxp. Ceh., de Meijere, 1904.
Among the interesting Echini collected by the " Siboga," in the Dutch East
Indies, were 14 small (7-25 mm. h. d.) specimens, taken at depths of 38-51 fths..
CLARE: THE CIDARIDAE. 195
which de Meijere described as Cidaru glandulosa. There can be no question of
their close relationship to St. dispinosay and it is quite possible, as de Meijere (p. 5)
himself suggests, that they are tbe young of that species. Besides the characters
already mentioned in tbe key, these specimens were remarkable for the number
of large globiferous pedicellariae, like those of P. baculasa^ which they bore.
Stephanocidaris hawaiiensis.
Stephanoddaris hawaiiensis A. Agassiz and Clark, 1907, Haw. Pac. Ech.
Cid., p. 18.
Plates 24 and 25, Haw. Pac. Bcli. Cld., A. AgMfAm and Clark, 1907.
A large series of specimens of this handsome species was collected among the
Hawaiian Islands by the " Albatross/' at depths of 20-320 fths. It is a typical
member of the genus, and is not at all likely to be confused with any other species.
The largest specimens are 34-42 mm. h. d. and have primaries 90-105 mm. long.
TBMNOCIDARIS.
Temnoddaria Cotteau, 1863, Pal. Fran9. Terr. Cr€t., 7, p. 366.
Plates 1065-1087 bis, Pal. Franc Terr. CMt., 7> Cottoan, 1803.
Test large, much like Phyllacanthus ; coronal plates 0-8 ; areolae very distinctly
sunken ; median interambulacral area broad, well covered with miliary tubercles,
with more or less horizontal, narrow grooves and deep, circular pits ; ambulacra
narrow, .20-.25 of interambulacra ; poriferous zones considerably sunken ; median
ambulacral area with numerous tubercles, often arranged in horizontal series, and
with a few deep, circular pits ; pores widely separated, more or less nearly horizon*
tal and connected as in Phyllacanthus. Abactinal system apparently larger than
actinostome. Primary spines stout, as in Phyllacanthus. Secondaries and pedi-
cellariae ? •
If Duncan C89) is correct in his surmise that the pits are of post-mortem
origin, Temnocidaris becomes of course a synonym of Phyllacanthus. Until this
can be demonstrated, however, the genus is entitled to recognition. The three
species which have been named are all from the Cretaceous.
QONIOOIDARIS.
Gonioddaris Agassiz et Desor, 1846. Cat. Bais. Ann. ScL Nat. (3), 6, p. 337.
Test moderately high, .50-70 h. d., but not especially thick or solid ; coronal
plates numerous in proportion to h. d., 6-11; areolae somewhat deeply sunken;
median interambulacral areas deeply and distinctly sunken (deeper than areolae,
especially at angles), and usually bare along vertical suture, often with short,
bare, lateral depressions along inner end of horizontal sutures ; in some cases, how-
ever, yertical suture nearly concealed and only lateral furrows conspicuous ; in still
196 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
other cases, even lateral furrows onlj faintly indicated; ambulacra broad, .3&-.45
of iuterambulacra ; poriferous zones more or less sunken ; median area much broader
than a poriferous zone, usually sunken and often bare along middle ; each ambu-
lacral plate bears a single secondary tubercle, a little above inner pore, and iu
addition 1-8 miliary tubercles, between which more or less space is left bare (amount
of bare space varies greatly in different species ; in tubariaf entire median ambulac'
ral area is sunken and bare save for marginal tubercles, while in mikado scarcely
any bare spaces are visible; other species clearly connect these two extremes);
pores oblique or rarely horizontal; distance between two less than diameter of
pore ; surface of interval elevated or roughened. Abactinal system variable, rang-
ing from less than .40 to over .50 h. d. Actinostome about equal to abactinnl system
or smaller. Primary spines very variable, .76-2.00 h. d., always rough, and thorny
or prickly ; tips of some usually more or less expanded into a large and conspicuous
crown, cup, or even plate, which is often only of a little greater diameter than thick-
est part of spine, but may become as much as .60 h. d. ; actinal primaries variable,
rough or serrate, usually somewhat flattened ; secondaries thick, of moderate length,
more or less flattened, rounded at the end. Tridentate pedicellariae wanting, and
IftTgCf globiferous ones with no end-tooth on the valves.
The typical examples of this genus, such as tubaria, are very easily recognized,
but it is less easy to place such species v&florigera and mikado. Nevertheless the
genus is vefy generally accepted and seems to be a natural group. Mortensen
(: 03) has made two new genera (Fetalocidaris and Schizocidaris) and a new sub-
genus (Discocidaris) out of the species here included in Goniocidaris, but none of
these rest on anything better than some trifling pecuh'arity in the large globiferous
pedicellariae. Whether we are to find the origin of Goniocidaris in such a form
as Phyllacanthus verticillata may be open to question, but the median ambulacral
and interambulacral areas of that species could easily be transformed into those of
G* tubaria^ while perfectly horizontal pores are found in Q. biserialis. There can
be little question, in any case, that the three southern species are closely related
to each other, and the same is true of the Japanese forms, while fiorigera seems
to be structurally, as well as geographically, intermediate. The genus is appar-
ently recent and confined to the southern and western Pacific Ocean. The follow-
ing key is based on the examination of 133 specimens, including all of the species,
except forigera.
Key to the Species.
Each coronal plate with but few (30-70) secondary and miliary tubercles,
median interambulacral area conspicuously bare and often sunken ;
median ambulacral area commonly without miliary tubercles, except
near margin, so that it is often bare and usually much sunken.
Small (20 mm. or less h. d.) ; coronal plates, 6-8 ; abactinal system about
.60 h. d. and actinostome nearly equal ; some primaries taper to a
point, while in many specimens, others, abactinal ones, are abruptly
and enormously expanded at tip into a plate, diameter of which may
be .60 h. d. ; primaries usually more or less covered, at least near
base, with a coat of woolly, calcareous hairs clypeata
CLARK: THE CIDAEIDAE. 197
Moderate or large (20-50 mm. h.d.); coronal plates 6-11; abactinal
system generally about .40 h. d., aud only partially covered with
miliary tubercles of various sizes; primaries seldom pointed and
with no covering of woolly calcareous hairs.
Abactinal system equal to, or larger than, actinostome; coronal plates
with tubercles near vertical suture much smaller than those next
to areolae tubaria
Abactinal system smaller than actinostome; coronal plates with
tubercles rather large and of nearly uniform size umlnraculum
Each coronal plate with numerous miliary tubercles, so that median inter-
ambulacral area is usually covered by them, except on sutures ; if bare
sunken areas are conspicuous at all, it is only on inner half of hori-
zontal sutures ; median ambulacral area with numerous miliary tuber*
cles, tending to cover it, so that it is never wholly sunken and bare.
Large (25-60 mm. h. d.) ; abactinal system almost uniformly covered with
small tubercles ; miliary tubercles on ambulacra, in horizontal series
with deep furrows between geranioides
Small (15-35 mm. h.d.); abactinal system not uniformly covered with
small tubercles ; miliary tubercles in median ambulacral area never
conspicuous, but often filling up entire space.
Lower edge of ambulacral plates occupied by minute tubercles, leav-
ing distinct bare spaces forming small, rectangular pits, which
alternate with each other ; primaries white or whitish in contrast
with reddish-yellow secondaries florigera
No definite arrangement of tubercles on ambulacra clear, and no dis-
tinct bare pits; primaries not "whitish in contrast with" darker
secondaries.
Test high, .G0-.70 h. d. ; abactinal system much less than vertical
diameter ; primaries more or less covered with calcareous hairs
and usually with a conspicuous, flat, horizontal plate just above
collar mikado
Test low, .50-.00 h. d. ; abactinal system nearly or quite equals ver-
tical diameter; primaries with relatively few, long and stout
thomSf but otherwise smooth biserialis
Qoniocidaris clypeata.
Goniocidaris clypeata Doderlein, 1885, Arch. Naturg., 51 Jhrg., 1, p.. 82.
Plate 6, Plate 4, figs. 9^-20, Jap. Seeigel, DUderlein, 1887.
This is one of the interesting species discovered by Doderlein in Japan, and
will be easily recognized from his excellent figures and description. The prevail-
ing color is whitish, pinkish, or brown of some shade. The material collected by
the " Albatross " shows beyond question that the little cidaroid described by
Doderlein ('87) as Porocidaris gracilis is a small example of this species, in
which the spines with enormously expanded tips are wanting. The " Siboga "
cidaroid called C. hirsuiispinus by de Meijere (: 04) is also evidently a young
example of this species ; the secondaries of clypeata are frequently exactly like
de Meijere's figure. Except this "Siboga" specimen, clypeata is known ouly
from the vicinity of Japan.
198 BULLTTIH: MUSEUIC OF OOKPARATIYB ZOSlOOT.
Goniocidaiis tnbaria.
CidariteM tMbaria Lftmarck, 1810, Anini. b. Yert , 3, p. 67.
GmwddariM tubaria Lfitken, 1864, Bid. Kand. Kch,, p 1S7.
Plate le, fls. ft. Plato 11.
Of this wdl-known species, notbtug farther need be said than that it seems to
be perfecUj distinct from geranioidei, although the ookir (light yellowish, red, or
deep browniab-red) is the same as that of many specimens of the latter. The
geographical range of this species is Tasmania and northward along the east coast
of Australia ; a specimen labelled ** Goniocidaris geranioides ? £ast India " is in
the collection of the M. G. Z.
Ooniooidaris tunbraoolnxn.
' Gonwcidaris umbracMlum Button, 1878, Trans. N. Z. Inst, U, p. 306.
Plate 10, fig^ 3 and 4.
This is the New Zealand representative of the preceding species, and so far as
can be determined from the three specimens at hand, is weU entitled to speci6c
rank. The bright green color of the test and the burger number of coronal plates
(10, as against 8 in tubaria of the same size) are good characters in addition to
those given in the key.
(Soniocidaris fferanioides.
CidartteM geranioides Lamarck, 1816, Anim. a. Vert., 3, p. 56.
Goniocidca-iB geranioides Agassiz et Desor, 1816, Cat Rais. Ann. ScL Nat (3), 6,
p. 387.
Plato Ig, flgs. 3, 4, ReT. Ifich., A. Afl^aaoix, 1873.
Although this species is quite similar to tubaria in general appearance, the
differences between them seem very constant; in addition to those mentioned
above may be added the frequently darker color (nearly black) and the much less
thorny spines o( geranioides. The geographical range appears to be the same.
Qoniocidaria floriKera.
Goniocidaris Jlorigera A. Agassiz, 1881, Challenger Echini, p. 46.
Plato 1, fiss. 7-20» Cballenicer Kcli., A. AgasslB, 1881.
This '' Challenger " species from the £ast Indies shows the same extraordinary
variety in its primary spines which is seen in elypeataj and it would be surprising
if the pedicellariae were not also variable. As I have no greater confidence in
the characters furnished by pedicellariae than I have in those which spines afford,
I can find no good reason for recognizing the genera and species based on the
** Challenger" material, which Mortensen (: 03) proposes : — Discocidaris serrata^
Schizocidaris assimilis, and Petalocidaris Jlorigera. Certainly if they are to be
CLARK: THE CIDARIDAE. 199
accepted, more adequate descriptions are necessary, and the differences between
the three species made more tangible. That C. fmbriata de Meijere (: 04) is iden*
tical Yfi^florigera seems to me practically certain.
Goniooidaris mikado.
Diseocidaris (Cidaris) mikado Doderlein, 1885, Arch. Naturg., 51 Jlirg., 1, p. 80.
Goniocidaris mikado Doderlein, 1887, Jap. Seeigel, p. 15.
Plate 7, Jap. Seeig<el, DiSderleIn, 1887.
This is another of the Japanese echinoids, which Doderlein's excellent work
has given us. Although undoubtedly nearly related to the preceding species and
to cfypeata, it is perfectly distinct and easily recognized. The minute, often
nearly spherical, secondary spines are very characteristic. The color is almost
cream-white, with a purplish tint abactiually and ou the primaries. Specimens
have as yet been taken only in the vicinity of Japan.
Qoniooidaris biserialis.
Stephanocidaris biserialis DUderleln, 1B85, Arcli. Naturg., 51 Jhrg., 1, p. 79.
Goniocidaris biserialis Doderlein, 1887, Jap. Seeigel, p. 10.
Plate 5, Jap. Seeigel, Doderlein, 1887.
This species is quite unlike the preceding in its general appearance, but re-
sembles it in the obliteration of the bare depressed areas on ambulacra and in-
terambulacra which characterize the typical members of the genus. The color of
.biserialis is quite variable, ranging from dull brownish-yellow, with more or less
of a green tint, to yellow, olive-green, or brownish-red. It is known only from
the vicinity of Japan.
POLYCIDARIS.
Pdycidaris Quenstedt, 1858, Der Jura, p. 644.
Plate 79, fkg, 69, Der Jara, Qaenstedt, 1858.
Test of moderate size, circular at ambitus, flattened ; coronal plates numerous
(O7I5) ; areolae somewhat deeply sunken, merging together throughout the entire
vertical series, even at ambitus; median interambulacral areas more or less bare
and depressed ; ambulacra narrow, .15-.22 of interambulacra, straight ; poriferous
zones little sunken ; median ambulacral area with only a single marginal series of
small tubercles; pores oblique, near together, separated by a slight elevation.
Abactinal system ? Actinostome ? Spines and pedicellariae ?
Doderlein ('87) appears to think this genus is near Dorocidaris, but to me it
is clear that its relationships are with Goniocidaris. Except for the narrow am-
bulacra and the merged areolae, P. nonarius is strikingly like 0. umbraculum.
Doderlein lists 5 species, all from the Jurassic strata of Europe.
200 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY.
OBTHOCIDABIS.
OrthocidarU Cotteau, 1862, Pal. Fran9. Terr. Cr€t., 7, p. 964.
Plate 1088, F1k»> 1-6, Pal. Fran^ Terr. Cr^t., 7, Cotteao. 1863.
Test of moderate size, circular at ambitusi very little flattened, so that it is
subspheroidal ; coronal plates numerous (14 or 15) ; areolae verj small, scarcely at
all sunken, their diameter less than one-fourth the horizontal diameter of plate at
ambitus, and little more than one-half its vertical height ; median interambulacral
area very broad, covered with miliaries and not sunken ; ambulacra narrow, .23
of interaoibulacra, straight ; poriferous zones very narrow, not sunken ; median
ambulacral area with about 4 vertical series of tubercles ; pores oblique, separated
by a low elevation. Abactinal system very small, about .25 h. d. Actinostome larger
than abactinal system, subpentagonal, about .88 h.d. Spines and pedicellariae ?
This is certainly a most un-cidaroid appearing sea-urchin, the straight, narrow
ambulacra, the numerous small and nearly uniform miliaries, and the remarkably
small areolae and primary tubercles are so unlike the Cidaridae, and yet if the
areolae were sufficiently enlarged to merge together vertically, the resemblance to
Pofyridaris muUiceps would be quite stiikiog. Only one species has been named,
inermis, from th^retaceous of Europe.
TRBTOCIDARIS.
Tretocidaris Mortensen, 1008, Ingolf-Ezp. Ech., p. 16.
Test moderately high but very variable (.45-.85 h. d.) ; coronal plates, 4r-S; are-
olae little or moderately sunken, tending to merge together actinally; median
interambulacral area more or less depressed, bare or covered with small tubercles,
sutural lines usually quite distinct; ambulacra .2(>-.d7 of interambnlacra in width;
poriferous zones more or less deeply sunken ; median ambulacral area with a double
series of tubercles along margin, inner much smaller ; intervening space may be
bare, or more or less covered with scattered tubercles ; pores as in Cidaris. Abac-
tinal system .40-.55 h. d., sharply defined, circular, or pentagonal ; ocular plates
with convex or straight outer margin, little or not at all notched by ambulacra;
•miliary tubercles covering abactinal system more or less variable in size and some-
what irregularly scattered, leaving bare spaces here and there, especially along
margins of genital plates. Actinostome, .37-.50 h. d., generally smaller than abac-
tinal system. Primary spines 1-8 h.d., usually more or less cylindrical or terete,
rarely with large and conspicuous thorns, but usually covered with longitudinal
series of granules, which may be very low so that spine is nearly smooth or only
granular, or may project sharply so that spine is prickly, or may be elevated and
merged together so that spine is longitudinally ribbed ; actinal primaries equally
diverse ; secondaries flat and not peculiar. Large globiferous pedicellariae some-
times wanting, sometimes as in Cidaris, sometimes with a ver^' small opening and
a powerful end-tooth on valves, and sometimes like small ones, which have a rather
large opening and usually an end-tooth.
This genus was established by Mortensen for three recent species {bartlettu
annulalay spinosa) hitherto placed in Dorocidans but whose pedicellariae, he
CLARK: THE CIDARIDAE.
201
found, were very diflFerent from those of D. papillata. While the pedicellariae of
bartletti are much too variable to be used as the basis for a genus, the abactiual
system of that species is so noticeably and constantly different from papillata
that I think the genus Tretocidaris may well be recognized. There are eight
other species of Dorocidaris which fall iuto the same group. It is a D)uch more
natural and better differentiated genus than Stereocidaris, which has been quite
generally recognized in the last decade. X^^ species of Tretocidaris are widely
distributed, occurring in the North Atlantic, the Caribbean and Mediterranean
Seas, the Gulf of Panama, northward along the Mexican coast, among the
Hawaiian Islands, along the Japanese coast, southward into the East Indies and
as far west as Ceylon. I have not attempted to determine whether any extinct
species are to be referred to this genus or not The following key is based on the
examination of 938 specimens, representing all of the species recognized except
iiara.
Key to the Species.
Test very high, .76-.86 h. d. ; ambulacra very broad, .83-.37 of interam-
bulacra, with median line bare ; primaries with 8 longitudinal ridges
(not notched or granular), pale pink at base, olive-green near tip . . tiara
Test more flattened, generally less than .70 h.d. ; ambulacra generally less
thaji .83 of interambulacra.
Median ambulacral and interambulacral areas bare along vertical sutural
line; coronal plates 6-8.
Test moderately flattened or high, .6O-.70 h. d. ; actinostome moderate,
.86-.45 h. d. ; median interambulacral area .26 or more of inter-
ambulacrum in width, with several series of miliary tubercles
on each coronal plate between scrobicular circle and vertical
suture; abactinal system fairly well covered with tubercles;
primaries 1.25-2.50 h. d. ; West Indian.
Abactinal system large, .45-.55 h. d. ; areolae at least actinally well-
sunken; primaries seldom cross-banded, usually terete, with
longitudinal series of numerous minute prickles but never
thorny affinis
Abactinal system small, .40-.45 h. d. ; areolae very shallow ; prima-
ries prettily cross-banded with reddish (or purplish) and yellow-
ish (or greenish), sometimes cylindrical, often terete, frequently
flaring at tip, not uncommonly flattened at base, with longitud-
inal series of rather coarse teeth and often more or less thorny . harikiti
Test much flattened, .45-.55 h. d. ; actinostome large (.40-.50 h. d.) ;
median interambulacral area .20 of interambulacrum, with only
1 or 2 incomplete series of miliary tubercles on inner end of coro-
nal plates; genital and ocular plates with margins free from
miliaries; Eastern Pacific.
Primaries reddish, very slender, 1-1.50 h. d. ; thickness of spine about
6 or 7% of length; covered with 14-15 longitudinal series of
low, rounded granules ; collar and secondaries dark, uniform,
brownish-red ; no tridentate pedicellariae panamensU
202 BULLKTIN: MUSEUM OF COMPARATIYS ZOdLOOT.
Primaries greenish, often cross-banded with darker, stoat, about
equal h. d. ; thickness 8-12 % of length ; coTered with 12-13
longitudinal series of coarse, sometimes sharp granules; collar
light reddish or whitish ; secondaries greenish, with a broad
longitudinal stripe of brownish- or purplishnred at tip ; triden-
tate pedioellariae common dnbia
Median ambulacral and interambuiacral areas not at all bare.
Coronal plates 4 or 6, rarely 6 even in large specimens; primaries
slightly swollen near base, terete, almost smooth ; large globifer^
ous pedicellariae wanting calaeaMika
Coronal plates 6-8, rarely 6 CTen in small specimens ; primaries not as
above.
AreoUe very small, those on largest coronal plate only .80-.06 of
length of pUte; abactlnal system .40 h. d. and actlnostome
.86 h. d perpUxa
Areolae moderate or large, those on largest coronal plate .70-.75 of
length of plate ; abactinal system about .60 h. d. and actino-
stome about .46 h.d.
Primary spines somewhat flattened, at least near base, with about
10 longitudinal series of coarse, sharp granules wliich usually
become fused near tip Into ridges; in oM specimens these
ridges may occupy entire length of spine, no separate gran-
ules being visible, while in other cases granules may be con-
spicuous as sharp prickles almost entire length of spine;
primaries white or whitish, spotted or banded with brownish-
red or purple ; collar very narrow bracteata
Primary spines terete, with 12-16 longitudinal series of fine, sharp
granules which do not lose their individuality entirely, even
near tip of old spines ; unicolor, white or pale yellowish ; collar
of moderate width rvtm
Tretocidaris tiara.
Doroeidarii tiara Anderson, 1894, Joum. Asiat. See. Bengal, 63, p. 188.
Plate 6, llgs. ff, ffa, IlL InTestigator Zo6T Boh., Aloook and Anderson, 1805.
This is one of the species collected by the " Investigator,'* the real position of
which is somewhat doubtful, although the figures given in " Illustrations . . .
Zoology . . . Investigator " (1895, pt 2, pUte 6, fi^. 2 and 2a) indicate its
position in Tretocidaris. The test is extraordinarily high, even though the
measurements given by Anderson represent some other method of estimating
the height of the test than that which is here used. There are several reasons
why tiara is not synonymous with St. indica DoJerlem, as has been suggested,
but it is still more incredible that it should be T, bracteata^ as Mortensea (: 03,
p. 173) asserts, unless Anderson's description and figures are to be entirely
ignored. Either Morteiisen has not seen a specimen of bracteata, or else Iiis
supposed specimen of tiara is not tiara at all. Anderson's figures and descrip-
tion are remarkably clear and complete, and unusually satisfactory, although he
CLAEK: THE CIDARIDAE. 203
fails to mention the pedicellariae. The test of tiara is chestnut-brown, green
abactiuall J, especially towards the anus ; the secondaries are olive-green with a
darker longitudinal band. The largest specimen was 42 mm. h. d. The only
recorded locality for tiara is off Colombo, Ceylon, in 142*400 fths.
Tretocidaris affinia
Cidaris affinis Philippi, 1845, Arch. Natarg., U Jhrg., 1, p. 861.
Plate 1» fig. 5, Bev. Boh*, A. Agassis, 1879. Plate 1, fig. 1, Insolf-Ezp. Ech**
Mortenien, 1903.
This well-known species has been confused with Dorocidaris papillata so long
that it may be hard to believe it is really quite different We are indebted to
Mortenseu (:03) for showing its right to specific rank (although he makes no
reference to the abactinal system !), but we cannot follow him in placing it in the
genus Cidaris. Mediterranean and West Indian specimens appear to be alike in
all particulars ; Mortensen says the ti-identate pedicellariae were wanting in his
Mediterranean specimens, but those in the collection of the M. C. Z. from Cape
Sagras and from the Mediterranean have them normally developed. Mortensen
says the spines are 1-1.5 h. d., but our large series of specimens show a much
greater range, 1.25-2.40 h. d. The largest specimen is 38 mm. h. d. The color
is variable, but the small spines of the test are more or less greenish, tipped with
dark red, while the entire abactinal system (or at least the sutural lines) and the
bare areas on ambulacra and interambulacra are dark red ; the primaries are dull
grayish, more or less pink or white near base, and with a greenish or brownish
collar. In West Indian specimens the color is often very light, the secondaries
and test being nearly cream-color with the former tipped with reddish. In other
West Indian specimens the color is sometimes nearly slate-color, with little trace
of reddish. This species ranges throughout the North Atlantic eastward into the
Mediterranean, and southwestward to Barbados and the Gulf of Mexico, down
to a depth of 500 fths.
Tretocidaris bartletti.
Dorocidaris BartleUi A. Agassiz, 1880, Bull. M. C. Z., 8, 2, p. 69.
Tretocidaris bartletti Mortensen, 1903, Ingolf-Exp. Ech., p. 16.
Plates 8 and 9. Also Plate 9, figs. 16-97, Blake Ech., A. Agasslx, 1883.
In his original description Agassiz called attention to the resemblance between
the primary spines of this species and of Stephanocidaris. Young specimens of
bartletti^ for this reason, show quite a striking resemblance to young specimens of
that genus, but a careful examination shows important differences in the primaries
as well as in the test. In spite of the very great diversity exhibited in both its
spines and its pedicellariae, there can be no question as to the real relationship of
this species. Mortensen (: 03) names two closely allied, supposedly new species,
which he found in the British Museum ; one, annulata, I am unable to distinguish
204 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGY.
from bartUtth for no characters are g^veii which do not occur in some specimens
of that species ; the other, itpinosa, may prove to be a Talid species, but its affinities
cannot be determined from the publisiied description. The largest specimen of
bartletti in the collection of the Musenm of Comparative Zoology is 49 mm. b. d. ;
another (Plates 8 and 9), not quite so large (47 mm. h. d ), has. the longest spines
93 mm., nearly all cylindrical and not at all thorny. The test of these specimens
is brown, varying from fawn-color to deep red-brown, or even deep red on the
abactinal system. In the National Museum there is a magnificent specimen of
bartletti 68 mm. in diameter. This species is known only from the West ladies
in 88-397 fths.
Tretocidaris pancunensia
Dorocidaria panameruis A. Agassis, 1898, Bull. M. C. Z., 33, p. 78.
Plates 1, 9, Fan. Deep Sea Ech., A. AffaMis, 1904.
This handsome species is the Paci6c representative of T. affinis^ but is quite ob-
viously distinct. The tridentate pedicellariae are wanting in all of the thirteen
specimens examined, of which the largest is 35 mm. h. d. The geograpliical range
oi panamensis seems to be limited to the west coast of Central America and around
Cocos Island, in 66-112 fthms.
Tretocidaris dubia, sp. nov.
Plate 6, figs. 3 and 4.
Test somewhat flattened ; vertical diameter about .62 h. d. ; coronal plates 6 ;
areolae distinct and not very deeply sunken; median interarobulacral area not
sunken, very sparsely covered with tubercles, only 6 or 7 on each coronal plate in
addition to the scrobicular circle ; ambulacra wide, nearly .40 of interambulacra ;
poriferous zones broad and little sunken ; median ambulacral area with a double
series of rather large tubercles on each margin, with space between perfectly bare ;
pores slightly oblique, rather large. Abactinal system .4&-.60 h. d., nearly circular,
and clearly defined, elevated at centre, very sparsely covered with small secondarj
spines ; genital plates rather large, higher than wide, with pores near outer edge ;
ocular plates more or less triangular, one (right anterior) or more excluded from
anal system, which is about one-half of abactinal system and has an outer series of
7-10 rather large plates and 9-12 smaller ones at centre ; all plates of abactinal
system carry a few rather coarse tubercles of nearly uniform size ; each genital
plate has 14-20 ± such tubercles and each ocular, 8-12 :J:. Actinostome slightly
smaller than abactinal system, not at all sunken, closely covered with stout plates,
3 or 4 in each interambulacrum and about 8 or 9 pairs in each ambulacrum.
Primary spines short, about equal to h. d., nearly cylindrical, seldom tapering, but
often truncate or slightly flaring at tip, covered with 12-13 low, longitudinal series
of coarse, sometimes sharp granules; actinal primaries much as in Cidaris and
usually longitudinally ridged at tip ; secondaries long and narrow, flat and slightly
widened at tip. Pedicellariae not peculiar; large and small globiferous, as in
panamensis ; tridentate much as in affinis. General color of test decidedly greenish,
especially abactinally, but anal system reddish-brown; miliary and secondary
CLARK: THE OIDARIDAE. 205
spines whitish, longitudinally striated with deep reddish-purple; on secoDdaries,
striations merge to form a broad stripe at tip of spine ; primary spines dull grayish,
sometimes indistinctly cross-banded with brown; collar flesh-color or whitish.
Largest specimen 25 mm. h. d. ; vertical diameter, 13 mm. ; abactinal system, 12 mm.;
actinostome, 11 mm. ; longest primary, 25 mm., a trifle mure than 2 mm. thick at
base.
That this species is closely related to panamensis seems clear, but that it is quite
distiuct is certainly indicated by the available material. None of the specimens of
either are in any way intermt diate. Both species were taken by the " Albatross *'
at Station 3378, in 112 fathoms off Galera Point, Cape San Francisco, Ecuador,
but only panamensis was found near Cocos Island, and only dubia at Station 3397>
in 83 fatiioms off Galera Point. Possibly dubia is a more southern species ; at
any rate, it is known only from the coast of Ecuador.
Tretocidaris calacajitha.
Doi'ocidaris calacantha A. Agassiz and Clark, 1907, Haw Pac. Ech. Cid., p. 11.
Plates 13, 14, 34, 36, Haw. Pac. Ech. Cid., A. Agassiz and Clark, 1907.
This very distinct species reaches a size of 43 mm. h. d., with spines 81 mm. long.
It is very pale brown with a greenish cast, especially on the abactinal system ; the
secondaries each have a broad green stripe ; the primaries are very faintly banded
with brown and at the base are finely spotted with white. This is one of tlie
species found by the " Albatross " at the Hawaiian Islands, where it is not rare
in 127-198 fths.
Tretocidaris perpleza, sp. nov.
Plate 6, figs 1 and 2 ; and Plate 7, figs. 1-4.
Test somewhat flattened ; vertical diameter, about .55 h. d. ; coronal plates 7 or
8 ; areolae small, only .60-.65 of horizontal length of plate, distinct and not very
deeply sunken; median interambulacral area very fully covered witii tubercles,
smallest next to vertical suture, which is quite distinct; ambulacra about one-third
of interambulacra in width; poriferous zones, broad and little sunken; median
ambulacral area with a double series of tubercles on each margin, inner much
smaller, and between these, 3-6 irregular series of small tubercles which sometimes,
but not always, conceal vertical suture ; pores nearly horizorttal, large, their hori-
zontal diameter much exceeding vertical. Abactinal system about .40 h. d., nearly
circular and clearly defined, flat and quite thickly covered with small secondary
spines ; genital plates rather large, nearly square or somewhat pentagonal, with
pores near outer edge ; ocular plates more or less triangular, with apex truncated,
when in contact with anal system, either wholly excluded, or some, or all except
right anterior one, in contact with a large anal plate : anal system about one-half
of abactinal, witii an external series of 10-12 large plates and 12-15 smaller ones at
centre ; except along margins all plates of abactinal system covered with rather
coarse tubercles of nearly uniform size ; each genital plate has 50-80 ± such tuber-
cles and each ocular 20-35 ±. Actmostome small, only about .35 h. d., not at all
206 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
sunken, closely covered with itout plates, 4 in each interambulacrum and about
10 pairs in each ambulacrum. Primary spines short, about equal to h. d., nearly
cylindrical, seldom tapering, but often flattened and widened at tip, covered with
14-24 longitudinal series of coarse, sharp granules; actinal primaries much as in
Cidarls and nearly smooth; secondaries, long and narrow, but rather thick and
often with a deep longitud.nal furrow on outer surface at tip, whicli is thus crescent-
shaped in cross-section. Pedicellariae not peculiar ; no large globif erous ones were
found, but small globlferous and tridentate, like those of dubia, are frequent.
General color of test decidedly greenish, especially abactinally; miliary spines
greenish ; secondary spines greenish with a broad longitudinal stripe of deep
reddish-purple; primary spines dull grayish with a bright olive-g^en base and
collar. Largest specimen, 60 mm. h. d. ; vertical diameter, 27 mm.; abactinal
system, 20 mm. ; actinostome, 18 mm. ; longest spine, 40 mm., 3 mm. thick at base,
6 mm. wide at tip.
lu some ways this species is much like dubia, but aside from the differences in
the tuberculatiou of the test, the small areolae, abactinal system and actinostome,
and the short piimaries with olive-greeu collar and conspicuously flattened tips,
are very characterisiic oiperplexa. The resemblance between the two species in
the color of the secondary spines is quite noticeable. Two of the five known
specimens of ihis species were collected by tlie " Albatross " in the Gulf of Cali-
fornia on a bottom of coarse sand, in 36-39 fathoms. The other three are said to
have been picked up on the shore of Clarion Island, the westernmost of the E^villa
Gigedo Islands.
Tretocidaxis bracteata.
Dorocidaris bracteata A. Agassiz, 1879, Proc. Amer. Acad., 14, p. 197.
Plate 10, flss. 1 and S.
This is apparently the East Indian representative of bartletH, though it is a
smaller species and obviously quite dififerent. Mortensen (: 03), on the supposed
characters of the large globiferous pedicellariae, places bracteata in Stephano-
cidaris, but as we have already seen, he probably did not have a specimen of that
genus for comparison. Moreover, the pedicellaria which he figures as a " large
globiferous " of bracteata is exactly like the smatl, globiferous pedicellariae of this
species, while the large globiferous pedicellariae of this species are actually like
those of Cidaris. However, these large ones are very infrequent and may be want-
ing, while the small ones are often very large, and it is apparently one of these
latter that Mortensen has figured as the characteristic pedicellaria of Stephanoci-
daris ! It seems to me that this serves as an illustration of the danger of relying on
the pedicellariae. This species is relatively small, the lai^est specimen being only
29 mm. h. d. The secondaries are pale purple or rose, with or without yellowish
tips, or flesh-colored with a longitudinal rosy stripe ; in old specimens those of
the ambulacra may be darker tlian those of tlie interambulacra, and thus noticeably
contrasted with them, and the abactinal system is dark brownish-red ; the prima-
ries always show more or less clearly the dark markings, which are usually pur-
CLARK: THE CIDAEIDAE. 207
plish, but may be reddish or greenish. Originally discovered by the " Challenger "
near Ambqiiia, this species has since been taken only by the ** Albatross " in
8agami Bay, Japan. Ita bathynietric range is 15-114 fms.
Tretocidaris reini.
Cidaris (Dorocidaris) reini Doderlein, 1887, Jap. Seeigel, p. 7.
Plate 4, flgi. 1-7, Jap. Seelflpel, Ddderlein,1887. Plate 1, tkgB^ 2, 3» Sibosa-Exp. Beh.,
de MeUere, 1904.
Although this species is closely related to the preceding, the material at hand
supports Doderlein's opiuion that lus Japanese specimens were a new species;
curiously enough, however, he makes no reference whateyer to bracteata! The
primary spines of the two species are quite distinct, as already shown ; the ocular
plates of reini are narrower and higher than in bracteata and more broadly in
contact with the anal system, and the difference in color is very marked ; when
reini is not uniformly yellowish with dull white spines, the uppermost coronal
plates, the iuterambulacral miliary spines, the genital plates and the anal system
are deep reddish, while the ocular plates and ambulacra with all their spines are
pale yellowish in marked contrast, just the opposite of the coloration in bracteata;
the primaries of reini ai-e apparently not banded or spotted in adults, but if
de Meijere's identification of his small £ast Indian specimens is correct, the young
must be very much like those of bracteata. In size and in the pcdicellariae, the
two species agree well; the largest reini reported is 34 mm. h. d. Excepting the
four young Cidaroids taken by the **Siboga" near the Kei Islands and Timor
which de Meijere refers to this species, but which might just as naturally be called
bracteata, reini has not been taken yet anywhere but in Sagami Bay and Ka-
goshima Gulf, Japan, in 83-158 fths.
DOROCIDARIS.
Dorocidaris A. Agassiz, 1869, Bull. M. C. Z., 1, p. 254.
Test much as in Tretocidaris, but ranging up to only .70-.75 li. d. Abactinal
system very different, its outline not often sliarply defined and rather irregular,
with re-entering angles between genital and ocular plates ; latter more or less pen-
tagonal and deeply notclied by ambulacra. Primary spines cylindrical, at least near
base, or terete, sometimes smooth, but usually with longitudinal series of granules,
or ridges, never ** winged " however, and generally not flaring at tip. Globiferous
pedicellariae, botli large and small, with a conspicuous end-tooth on the valves ;
tridentate pedicellariae usually present.
Although this genus is quite easily distinguished from the preceding, the line
of division between it and Stereocidaris is exceedingly hard to draw, and it is an
open question whether there is sufficient ground for keeping them separate. As
small genera are more convenient and wieldy, however, we may retain the division
recognizing that the line is a very arbitrary one. As here used, Dorocidaris in-
cludes five species, which are found only in the Atlantic Ocean and almost entirely
208 bulletin: museum of comparative zoology.
north of the equator. Numerous fossil Cidaridae from Tertiary, Cretaceous,
Jurassic, and possibly even Triassic strata are to be referred to either this genus
;or the preceding. The foUowiug key is based on the examination of 536 speci-
mens, representing all of the liviug species, except nuda.
Key to the Species.
Primary spines more or less white and smooth, rarely conspicuously gran-
ular, prickly, or ridged, and neither during iior conspicuously flattened
at tip ; median ambulacral area less than .60 of ambulacrum and almost
wholly covered with small tubercles (xbifssicola
Primary spines more or less prickly, granular, or ridged.
Each coronal plate with only a few tubercles on iniier half (generally
less than 25, not counting scrobicular circle) ; sutural line of ambu-
lacra usually distinctly visible; each ambulacral pUte with I or 2,
seldom 3, tubercles ; primaries more or less cylindrical, often flar-
ing at tip, and never conspicuously flattened there ; median inter-
ambulacral area less than .25 of interambulacrum in width ; sutural
line usually quite distinctly sunken and bare.
Whole abactinal surface well covered with light-colored secondary
and miliary spines papillata
Whole abactinal surface appearing noticeably bare from small number
of secondary and miliary spines present ; test light-colored, but all
spines reddish-brown or purple nuda
Each coronal plate with numerous (more than 30) tubercles on inner half ;
sutural line of ambulacra often not visible, each plate with 2-5
tubercles.
Median interambulacral area less than .25 of interambulacrum ; sutural
line quite distinct; abactinal system with numerous tubercles
(genital plate with 110±; ocular with 3Q±>; primaries often
. flattened near tip, sometimes greatly expanded into broad flat
fans hUxkei
Median interambulacral area often more than .25 of interambulacrum ;
sutural line well concealed by tubercles ; abactinal system with
rather few, large tubercles (genital plate with 55 db; ocular with
20 db); primaries terete, covered with sharp granules and never
either conspicuously flattened or flaring at tip rugosa
Dorocidaris abyssicola.
Dorocidaris abyssicola A. Agassiz, 1809, Bull. M. C. Z., 1, p. 253.
Plate 1, figrs. 1-4, Rev. Ech., A. Agassiz, 1873.
This species seems to be quite distinct from papillata, and while it is occa-
sionally much like blakei or rugosa in certain features of the test, the primaries
commonly distinguish it from either of them at a glance. In addition to the
characters given in the key may be meutioued the folio wiug: the abactinal sys-
tem is. very large (.48-.55 h. d.), while the actinostome is relatively quite small
(.35-. 15 h. d. but ouly .70-.80 of the abactinal system) ; the test is usually under
CLA.RK: THE OIDARIDAE. 209
.60 h. d. in vertical diameter, aad it, as well as the secondaries, is pale brown or
yellowish; the ahactiual system is sometimes quite red; the uppermost coronal
plates do not carry primaries, and eyeu the second ones may lack a well-developed
spine; the primaries are usually about 1.25 h. d. and never exceed 2 h. d. The
diameter of the test is usually about 25 or 30 mm. but is sometimes 35 or 40, and
the largest specimen is 68 mm. h. d. Tliis species ranges from St. Lucia north-
ward to the coast of South Carolina and the region south of Martha's Vineyard
at depths of 100-200 fths.
Dorocidaxis papillata.
Cidaris papillata Leske, 1778, Add. Nat. Dig. Ech. Klein, p. 61 (partim).
Dorocidaria vapillata A. Agassiz, 1869, Bull. M. C. Z., 1, p. 254.
late lb, Rev. Boh., A. Affauis, 1879.
Nothing more need be said of this well-known species than that it does not
seem to occur in the western part of the Atlantic, but is apparently confined to the
northern and eastern parts of that ocean and to the Mediterranean Sea. The
bathymetric range is from a few fathoms down to about one thousand. Mor-
tensen's (: 03, p. 170) assurance that the ** Challenger'* specimen from St. Paul's
Kock is resillj papillata is important in this connection, but I think it possible that
the individual may prove to be rugom! In size papillata reaches a diameter of
58 mm., while in color it is quite variable, ranging from grayish-white to reddisb-
yellow, becoming biick-red on the abactinal system, with dull grayish or yellowish
primaries.
Dorocidaria nuda.
Dorocidaris nnda Mortensen, 1903, Ingolf-Exp. Ech., p. 171.
This species is apparently distinct from all the other members of the genus, but
its real relationships can oniy be determined when it is more fully described.
Possibly it is not so closely allied to papillata as I have assumed. The size is not
mentioned, but the test is white and the spines purple or reddish-brown. It has
been taken only in the Gulf of Guinea and near the Cape Verde Islands, in 53-
250 fths.
Dorocidaris blakei. ^
Dorocidaris Blakei A. Agassiz, 1878, BhII, M. C. Z., 5, p. 185.
Plate 4, Bull. M. C. Z., 5, 9, A. Asassiz, 1878. Plate 1, Blake Ech., A. Agrassis,
1883.
This is one of the most interesting discoveries of the " Blake," and specimens
with fully developed primaries are indeed unique. The color is grayish with more
or less of a yellow-brown tinge to the test. The largest specimen is 37 mm. h. d.
with spines 76 mm. long. Specimens in which there are none of the conspicu-
ously flattened primaries are easily recognized by the large abactinal system,
VOL. LI. — wo. 7 14
210 bulletin: museum of comparative zoology.
.45'.55 h*d., almost uniformly covered with small tubercles; the narrow porifer-
ous zones, about .20 of ambulacra, and the numerous small tubercles on the
interambulacra. This species ranges from Havana to Barbados in 197-450 fths.
Dorocidaris rugosa» sp. nor.
Plates 4 and 5. Plate 7, fiffs. 5-^8.
Test rather high, Tertical diameter about .60 h. d. ; coronsl plates 7; areolae
deeply sunken and distinct ; median interambulacra! area rery fully covered with
tubercles, smallest next to rertical snture, which is quite distinct ; ambulacra less
than one-third of interambulacra in width ; poriferous zones narrow and deeply
sunken ; median ambulacral area with a double series of marginal tubercles, inner
much smaller, and between these some small scattered tubercles tend to conceal
vertical suture ] pores oblique, small. Abactinal system about .46-50 li. d., irregular
in outline, stout and heavy somewhat as in Stereocidaris, covered with rather
coarse tubercles ; genital plates somewhat pentagonal, with lateral margins con-
cave, and pores not far from centre ; ocular plates more or less pentagonal, usually
wholly excluded from anal system, but posterior ones sometimes in contact with
anal plates» more or less notched on outer edge by ambulacra ; anal system not
quite one-half of abactinal, with an external series of 10-12 large plates and 12-15
smaller ones at centre ; except along margins all plates of abactinal system covered
with rather coarse tubercles of nearly uniform size; each genital plate has 60^-60 db
such tubercles and each ocular plate 20-30 db> Actinostoroe small, about .40 h. d.,
not at all sunken, closely covered with stout plates, 6 in each interambulacrum
and about 10-12 pairs in each ambulacrum. Primary spines long, 2-2.5 h. d.,
terete, usually swollen just above collar, and thence tapering to tip, covered with
12-16 longitudinal series of conspicuous sharp granules ; actinal primaries slightly
flattened, a little curved and somewhat serrate ; secondaries not peculiar, of mod-
erate length and width, flat, blunt, or truncate at tip. Pedicellariae as in papillata.
General color of test yellowish or brownish, more or less rose-red or brick-red,
abactinally ; secondaries and miliaries same as test ; primaries whitish or grayish,
abactinal ones sometimes bright rose ; neck smooth, polished, white, brownish, or
pink ; collar narrow, pale brownish or rarely lighter than neck. Largest specimen
in the Museum of Comparative Zoology, 40 ram. h. d. ; vertical diameter, 24 mm. ;
abactinal system, 20 mm. ; actinostome, 17 mm. ; longest primary, 80 ram., 5 mm.
thick near base, somewhat more than 1 mm. thick at tip. In the National Museum
is a fine specimen 60 ram. h. d.
This species is clearly the representative of papillata in the western Atlantic,
but may be readily distinguished from tiiat species by the broader and more com-
pletely covered median iuterambulacral area, the much more fully tubercled
median ambulacral area, the more uniformly tubercled abactinal system, and the
terete and very prickly primary spines. The distribution of rugosa is only
imperfectly known; the specimens I have examined are from stations between
32® N. lat. (off Savannah, Ga.) and Barbados and St. Vincent, in 164-337
fathoms. There are 8 specimens in the collection of the U. 8. National Mu-
seum, several of which have been labelled by Mortensen. One (No. 21,444)
is labelled " Stereocidaris ingolfiana," which is a very natural mistake, as small
CLARK: THE GlDAfilDAE. 211
specimens of the two species are very difficult to distinguish. The others are
labelled " Dorocidans papillata/' which is what one would naturally call them, if
rugosa is not tcT be recognized as valid.
__ •
CAIjOCIDABIS, gen. noT. (Greek, iraX^f, beautiful, + cidaris).
Test large and rather high; coronal plates 7 or 8; areolae distinct and con-
siderably sunken, the most actinal tending to merge together vertically ; median
interambulacral area not at all sunken, covered with numerous miliaries and with
more or less horizontal grooves or narrow furrows, such as occur in Temnocidaris;
ambulacra about .25 of interambulacra in width ; poriferous zones scarcely at all
sunken ; median ambulacral area very wide, about .65 of ambulacrum, with very
few tubercles aside from the customary double marginal series; pores oblique,
large and close together. Abactinal system not quite .60 h. d., of very irregular
outline; ocular plates deeply notched by ambulacra. Actinostome very small,
only about .65 of abactinal system. Primary spines 3 h. d., cylindrical, white,
smooth, and polished like porcelain, more or less tinged with pink and green;
actinal primaries flat and longitudinally fluted, but not notched or serrate. Sec-
ondaries flat and tapering, many bluntly pointed. Pedicellariae as in Dorocidaris.
Although in many respects like Dorocidaris, the very broad and nearly bare
median ambulacral areas, the remarkable color, and especially the smooth, pol-
ished primaries, mark this genus at a glance. The largest primaries are all broken
in the specimen in the Musum of Comparative Zoology, so that their length is
not shown in the figure given. But a specimen in the U. S. National Museum,
which is the most beautiful echinoid I have ever seen, is nearly perfect. The
primaries are 160 mm. long, rather more than 3 times the diameter of the test,
and scarcely taper at all, but are cylindrical throughout their entire length. The
genus is monotypic and very few specimens are known. The above desorij^
tion is based on a specimen 61 mm. h. d., from near Barbados, but two other
specimens in the U. S. National Museum have been examined.
Calocidaris micans.
Dorocidaris micans Mortensen, 1903, Ingolf-Ezp. £ch., p. 23.
Plates.
This is easily the handsomest, as well as one of the largest, of the West Indian
cidaroids. It reaches a diameter of more than 60 mm. The test is white, and
the secondaries nearly so, but the abactinal system and adjoining coronal plates
are pale green ; the primaries when dry are shining white, with a pink base and
occasional faint, in*egular marks of the same color ; they look as though artificially
polished. In alcoholic specimens the spines have a greenish shade and the pink
is deeper. The only known specimens of this beantiful species were taken by
the " Albatross " off the northwestern coast of Cuba in 205 fths., and by the
" Blake" off Barbados in 125 fths.
212 BULLETIN : MUSEUM OF COMPABATIYE ZOOLOGT.
AUSTBOCID ABIS9 gen. dot. (Latin au$ter, the sonth wind, + cidaris).
Test flattened, .60-.60 h. d., bat otherwise much as in Dorocidaris ; abactinal
system much more sparsely corered with miliaries ; secondaries more or less nearly
cylindrical and thickened at tip; primaries generally short, often less than h. d.,
and usually smooth (in IndiTiduals where primaries are long and rough, second-
aries are nearly fiat, so that resemblance to Dorocidaris is marked). Tridentate
pediceUariae wanting and globiferous pedicellariae with no end- tooth on Talres ;
eggs and young carried by female {mortensenif).
Were it not for their geographical isolation it would hardly be worth while to
attempt the separation of these three small species from Dorocidaris, but as they
haye the above given peculiarities in common and are probably more nearly
related to each other than to any other forms, it is convenient to keep them apart.
They are confined to the southern parts of the Atlantic and Indian oceans, their
known range extending from 75° W. to 90^ E. longitude and from about 35 ^
to nearly 70° S* latitude. The following key is based on the examination of 70
specimens of nutrix and canaliculata.
Key to the Speoies.
Actinal primaries not conspicuously flat, trowel-shaped, and entire.
Median ambulacral and interambulaoral areas bare and more or less
deeply sunken ; interambulaoral area usually with a conspicuously
deep vertical furrow ; vertical diameter about .65-.60 h. d. ; abacti-
nal system and actinostome rather small, .35-.40 h. d., about equal,
or former smaller canaUcdata
Median ambulacral and interambulacral areas little bare, and not at all
, « sunken ; vertical diameter about .45-.65 h. d. ; abactinal system and
actinostome large, about .60 h. d., about equal or former larger . . nutrix
Actinal primaries conspicuously flat, trowel-shaped, and entire ; primaries
long mortenseni
Austrocidaj*is oanalioulata.
Temnocidaris canalicuUUa A. Agassiz, 1863, Bull. M. C. Z., 1, p. 18.
Plate 1} gf lis. It, Bev. Bch., A. Agassis, 1873. Plate ff, figs. 1-S» Challenger
Eoh., A. Agaisls, 1881.
Some of the differences between this species and the next have already been set
forth by Mortensen ( : 03), but he has entirely ignored the more impoHant differ-
ences in the test and abactinal system. Moreover he has himself been led astray
by the remarkable diversity which this species exhibits in its color, spines, and
pedicellariae, and has described as a new species of Stereocidaris,. which he calls
lorioli, the long-spined form of canaliculaia, which the " Challenger " collected off
the mouth of the River Plate (Station 320). The Museum of Comparative Zool-
ogy contains one of the " Challenger " specimens from St 320, and also a large
CLARE: THE CIDA.RIDAS. 213
series of specimens from Patagonia. The latter so completely yet gradually con-
nect the indiyidaals having primaries 3.5 h. d. with those from the Falkland Islands
in which the primaries are only .65 h. d*, that their identity cannot be doubted.
Had Mortensen carefully examined an interambulacrnm, he probably would not
have been misled. Although usually about 25-30 mm. h. d., there are specimens
of canalieulata at hand 36 and 39 mm. ; the primaries range from 16 to 63 mm.
The color varies from very pale yellowish (with pink necks on the primaries) to
very dark brown. Tliis species is apparently confined to the eastern and southern
coasts of Patagonia and the neighboring isUnds. The bathymetric range is from
the shore to 600 fathoms. A specimen in the National Museum, which was
obviously collected many years ago, is labelled '* Navigator Ishinds."
Austrocidaris nutriz.
Cidaris ntUrix Wy ville Thomson, 1876, Joum. Linn. Soc. London, 13, p. 62.
Figf. 4, p. 63, Joam. Idnn. Soc liondon, 13, WyriUe Thomson, 1876.
There can be little question that this species is quite distinct from the preceding.
Like it, however, it shows considerable diversity in color and the length of the pri-
maries ; some specimens are almost black, with light-colored primaries, while
others have the test and secondaries, as well as the primaries, very light colored-
Mortensen ( : 03) asserts, without offering any evidence to support his view, that
the specimens collected by the " Challenger "at stations 147, 153 and 156 are
not this species because the water was too deep at those stations for a shallow
water species like nutria!* In view of the fact that a number of echinoderms are
known with a very great bathymetric range, we can hardly consider the argu-
ment conclusive. The largest specimen of nutrix at hand is only 30 mm. h. d.,
but the primaries are 66 mm., while a specimen 26 mm. h. d. has primaries only
18 mm. This species appears to be confined to Grozet, Heard, and Kerguelen
Islands, and the neighboring seas.
Austrocidaris mortenseni.
Goniocidans mortenseni Koehler, 1902, Belgica Ech. et Oph., p. 6.
Viga. 1, 11, 17, 29, 30, Belgica Ech., Koehler, 1902.
It is quite possible that this species does not belong here, but so far as can be
judged from the description and figures given it is most nearly allied to the fore-
going species. Koehler says nothing about the secondaries, and as the primaries
are very long, it is possible that the secondaries are not especially peculiar. The
largest specimen was 26 mm. h. d., with primaries 60 mm. The color of the test
and secondaries is very dark, while the primaries are reddish. Koehler says there
was no indication that the species is " viviparous," but as he only had a siugle mature
specimen, and that possibly a male, further light is needed on this point. The
specimens were collected by the " Belgica " in the Southern Ocean, near 70° S.
latitude and 87** E. longitude, in depths of 55-330 fths.
214 . BULLEHN: MUSEUM OF COMPABATIVE ZOOLOGT.
OBNTROCIDARIS.
Centroddaris A. Aga88iz, 1904, Pan. Deep Sea Ech., p. 82.
Test Tery flat, yertical diameter generally less than .60 h. d. ; coronal plates 7 or 8 ;
areolae yery little sunken ; median interambulacral areas narrow^ a little sunken,
and bare ; ambulacra yery broad, .65-.00 of interambulacra ; poriferous zones little
or not at all sunken ; median ambulacral area broad, flat, or somewhat depressed,
with a double marginal series of tubercles, outer much larger ; interyening space
bare, or each ambulacral plate may carry an additional miliary tubercle ; pores
yery large, nearly or quite horinmtal ; distance between two about equal to diamr
eter of pore ; snriace of interval slightly elevated. Abactinal system moderate,
.45-.50 h. d., with few (about 100) tubercles ; genital plates much higher than wide,
narrow, and bluntly -pointed externally ; oculars entirely excluded from anal sys-
tem, very wide and low but sharply pointed, with a markedly concave outer
margin. Actinoetome, .40-.46 h. d. Primary spines straight, cylindrical, slender,
and nearly or quite smooth, about equal to h. d. or somewhat longer; actinal prima-
ries not peculiar save for a wide colkir ; secondaries flat, thin, and narrow. All
three kinds of pedicellariae usually present ; large globiferous ones of two quite
distinct sorts, one with broad, fiat valves and neither lip nor end-tooth, the other
with curved valves (like Cidaris), but with a prominent end-tooth and Up.
This monotypic geuus was established for a very interesting and handsome
cidaroid taken by the " Albatross " in 1891 off Cocos Island, 52 fths. In 1904-
05, the " Albatross *' obtained a dozen additional specimens near Hood Island,
Galapagos, 100-300 fths., so that it is now possible to diagnose the genus fully.
It is quite distinct from Groniocidaris, though it resembles it in the broad ambu-
lacra, but it is doubtful if it is nearer to any other known genus.
Centrocidaris doederleini.
Goniocidaris Doederleini A. Agassiz, 1898, Bull. M. C. Z., 32, 6, p. 78.
CetUrocidai-ia Doederleini A. Agassiz, 1904, Pan. Deep Sea Ech., p. 33.
Plate 14, figs. 1» H, Pan. Deep Sea Ech., A. Agassis, 1004.
In young specimens the primary spines are very white and shiningi and have
8-10 slightly elevated, glassy, longitudinal ridges, but these practically disappear
with age and the spines become dull and yellowish. In alcoholic specimens the
secondaries are green, slightly tipped with dark yellow, while the test is greenish
with the lines between the genital and ocular plates and the bare spaces of ambu-
lacra and interambulacra deep purplish or dull red. The lai^est specimen is 28
mm. h. d. and the longest spines measure 33 mm.
APOROOIDARIS.
Aporocidaris A. Agassiz and Clark, 1907, Haw. Pac. Ech. Cid., p. 86.
Test flattened, .60-.60 h. d. (but abactinal system sometimes so much elevated
that vertical diameter from centre of anal system, .60-.80 h. d.), rather thin and
fragile ; coronal plates 6, rarely 7 ; areolae only slightly sunken ; median interam-
CLARK: THE CIDARIDAE. 215
bulacral area rather wide, bare, and slightly sunken along sutural line ; ambulacra
about .30 of interambulacra ; poriferous zones almost flush with test ; ambulacral
plates few, 30-32 in largest specimens; median ambulacral area somewhat wider
than a poriferous zone ; each ambulacral plate is vertically wide and carries only
a single tubercle, except in large specimens, when a second smaller tubercle is
present and then vertical suture is obscured ; pores very close together, somewhat
oblique. Abactinal system very large, .60-.70 h. d., either flat or more or less ele-
vated, with few or many tubercles. Actinostome .40-.50 h. d., consequently only
.60-.80 of abactinal system, and notable for small number of plates borne by mem-
brane, more or less of which near outer margin is quite bare. Primary spinet slen-
der, straight, and cylindrical, very finely prickly, white or nearly so, 1.5-3 h. d. ;
actinal primaries either coarsely or finely serrate or entire ; secondaries and milia-
ries alike, cylindrical or club-shaped, blunt and more or less erect, rather scattered.
Pedicellariae of only one kind, globiferous, but very variable in size.
The affinities of this interesting genus are rather obscare, for although the sec-
ondary spines resemble those of Austrocidarit nutris, it is hard to believe that
there is any close relationship to that species. There are no other living species
of Cidaridae which approach sufficiently near the three rare species placed here
to give us any real clue to their natural position. Although A. milleri has actinal
primaries similar to those of Porocidaris, there is little else to ally it with that
genus, and the other two species are even more different The large abactinal
system, few ambulacral plates, unsunken poriferous zones, somewhat bare actino-
stome, and the primary spines are striking reminders of Salenia. Two of the
species are discoveries made by the "Albatross" and are found only in the deep
waters of the Pacific Ocean ; although milleri was once taken in 465 fths., most of
the specimens are from over 1,600 fths. Ku&fragilis has been taken only at depths
exceeding 1,500 fths. The third species was found by the " Belgica " in much
shallower water, but in the far Antarctic Ocean. The following key is based on
the examination of 116 specimens of the two " Albatross " species.
Key to the Species.
Test moderately high, .55 h. d. and more ; abactinal system elevated, with
numerous tubercles (250-300 on a system 13 mm. across) ; ambulacral
plates about 20, in a specimen 15 mm. b. d tnUleri
Test flat, about .50 h. d. ; abactinal system not elevated, with comparatively
few tubercles (100-200 on a system 13 mm. across) ; ambulacral plates
about 15, in a specimen 15 mm. h. d.
Color reddish- or yellowish-brown ; arctic . fragilis
. Ck>lor bay or reddish ; antarctic incerta
Aporocidaris milleri.
Porocidaris MiUen A. Agassiz, 1898, Bull. M. C. Z. 32, 5, p. 74.
Aporocidaris Milleri A. Agassiz and Clark, 1907, Haw. Pac. Ech. Cid., p. 37.
Plate 6, Pan. Deep Sea Ech., A. Agassiz, 1904.
. The test of this species is grayish, sometimes with a purple tinge, or yellowish,
and the secondaries are of about the same color or paler. The primaries are
216 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
nearly or quite white. The largest specimen is 31 mm. b. d., while the primaries
are sometimes 75 mm. long. The abactinal system is often elevated 3 or 4 mm.
above the test. The " Albatross " collected this species in 1891, in the deep
water between Acapolco and Panama, and the Galapagos, 466-1880 fths., while
in 1904-05 she found it common in the still greater depths south and south-
west of the Qalapagos, 3005-2153 fths.
Aporooidaris fragilis.
AporocidarUJragilis A. Agassiz and Clark, 1907, Haw. Pac. Ech. Cid., p. 87.
Plate S3, flffs. t^S, Haw. Pao. Bch. Cid., A. Asaisis and Clark, 1907.
Of this species, the only known specimens, the largest of which is 23 mm. h. d.,
with primaries over 40 mm. long, were taken by the " Albatross " in the North
Pacific, south of Akska and southwest of Kamchatka in 1557-1973 fths.
Aporooidari8 inoerta.
Porocidaris incerta Koehler, 1902, Belgiea Ech. et Oph., p. 7.
Fiss. 8, 16, Belgiea Boh., Koehler, 190]l.
Roehler's supposition that this species is related to milleri is quite correct,
though in the shape of the test it is more ]Jikt/ragili8. The position of incerta
in this genus is confirmed not only by Mortensen's (: 03) examination of the pedi-
cellariae, but by a careful comparison of Koehler*s description, with a specimen
oifragilis of the same size (15 mm. h. d.) as his largest specimen. It is difficult
to make out from that description just how much difference there is between the
Arctic and Antarctic species. The latter was taken by the '* fielgica '* about
20 degrees south of Kerguelen Island, in 55-165 fths.
Stereocidaris.
Stereocidaris Pomel, 1883, Class. Math. Gen. Ech., p. 110.
Test very similar to Dorocidaris, but usually flatter (.50-.60 h. d.), with fewer
coronal plates (4-7, rarely 8 or 9) and relatively fewer primary spines (3-7, rarely
8, in each vertical series) ; that is to say, uppermost coronal plat« without primary
spine, and second often, third very rarely, similarly bare. Abactinal system large
(.85-.55 h.d., usually about .53), often convex, and noticeably thick and stout, but
this character varies much within a single species ; abactinal miliaries and second-
aries usually very small, but this character also varies much. Primary spines
usually flaring at tip, or if tapering, provided at base with conspicuous buttress-like
"wings"; *' winged " primaries are usually noticeably compressed, but otherwise
primaries are cylindrical. Globiferous pediceliariae, large and small, commonly
lack a conspicuous end-tooth on valves.
This is the most poorly defined and unsatisfactory genus in the family, and
yet the species contained in it have something about their general appearance
which is distinctive and makes it possible to recognize them usually at a glance.
They show considerable diversity in test, spines, and pediceliariae, and some in-
dividuals are strikingly like Dorocidaris. It is only when a considerable amount
CLARK: THE CIDARIDAE. 217
of material is available for comparison that such indiyiduals can be properly
placed. Unfortunately in preparing the following key there have been available
only five species, represented by 69 specimens, and it is probable that errors have
crept in which might have been avoided had a larger series of specimens been
available. However, Anderson's and Doderlein*s descriptions and figures are
sufficiently complete and accurate to make it possible to include their species.
Doderlein's (:06) measurements and figures have been of the greatest help.
The Japanese species need revision based on plenty of material, and it is possible
that the tliree species here recognized will prove to be simply forms of a single
species, as the differences between them are slight. All the recent species occur
in depths of 40 fathoms or over, and all but one {ingol/Utna) are found only in the
Indo-Pacific region. A number of fossil species from the Cretaceous are referred
tb this genus. How Doderlein (:06) can lay great stress on the form of the
pedicellariae in Stereocidaris and write without qualification '' Grosse und kleinere
globifere Pedicellarien ohne unpaaren Endzahn*' (p. 102), is incomprehensible,
for his own figures (Plates XXXVI and XXXVII) contradict the statement flatly.
Had I examined no specimens, the study of Doderlein's figures would have satis-
fied me that the pedicellariae are no more reliable than the spines.
Key to the Species.
Actinostome very small, .20-.35 h. d., usually under .30 except in young
specimens.
Primary spines often more or less trigonal, but seldom with three con-
spicuous "wings" near base; tridentate pedicellariae wanting;
pedicels contain perforated plates besides thorny curved rods.
Longest primary spines, 1.3-2.7 h. d., thickness commonly less than
8% of length; perforated plates in pedicels small, with few large
holes indica
Longest primary spines about 1.35 h. d., thickness about 10% of length;
perforated plates in pedicels broad, with many small holes . . . capensis
Primary spines commonly with three conspicuous wings near base ;
tridentate pedicellariae common ; pedicels with few or no . per-
forated plates tricarinata
Actinostome larger, almost always over .35 h. d.
Primaries pale pink or reddish, with 10-16 longitudinal series of fine
prickles, which often merge into ridges, and 1 (or more) of these
becomes a conspicuous " wing '* or " buttress " on basal half of spine,
which is also often flattened ; primaries tapering towards tip ; coronal
plates 5 or 6 (rarely 7).
Abactinal system coarsely tubercled ; median ambnlacral- area de-
pressed and bare along vertical suture, each plate with only 1 or
2 tubercles ; color of test and secondaries madder purple . . . cUcocki
Abactinal system with numerous small tubercles ; median ambulacral
area not depressed, often elevated along vertical suture, which is
seldom visible, crowded with tubercles, each plate with 4-6 ; color
of test and secondaries brownish, usually very pale ; no tridentate
pedicellariae ingotfiana
218 BULLETIN: MUSEUM OF COMPARATIVE ZOOLOGY.
Primaries cylindrical, at least near base, nerer proTided with " wings/'
bat with more or less erident, longitudinal series of rounded or sharp
granules, tending to become ridges near tip of spine, which is often
flaring.
Ambulacra very narrow, only .1S-.25 of interambulacra, not deeply
sunlcen ; median area closely covered with 6 series of tubercles ;
all miliaries very minute ; neck of primaries white . . mierotuberculata
Ambulacra .25~.d8 of interambulacra in width.
Secondaries not white; actinostome much smaller than abactinal
system ; tridentate pedicellanae present.
Abactinal system elevated 10% or more above test; abactinal
surface appears very bare fh)m small, wide, closely appressed
miliaries; primaries not white with purple collar .... grandis
Abactinal system flat or little elevated; abactinal snrfiice well
covered with ordinary miliaries; primaries when perfectly
clean, white, usually with a distinct purple collar . . . leucacantka
Secondaries white or whitish; actinostome nearly equal to abac-
tinal system ; no tridentate pedicellariae scepHferoidea
Stereocidaris indioa.
Stereocidaris indica Doderlein, 1901, Zool. Anz., 23, p. 19.
Plate 10, figs. 1, «} Plate 11, Deatsehe TleAee Exp. Bch., IKSderleln, 1900.
This species appears to be very variable, and Doderlein ( : 06), recognizes four
varieties (integra, africana, carinata^ sumatrana), based upon slight differences
chiefly in primary spines and pedicellariae. He says, however, that he doubts the
constancy of any of these varieties except sumatrana^ which appears to be well-
marked. Doderlein's admirable descriptions and his tables of measurements are
all that could be desired, but the figures given often suffer from indistinctness ;
they are quite good enough, however, to reveal the notable diversity in the pedi-
cellariae of this species. The color is yellowish, each of the larger secondaries
with a dark spot and the actinal primaries white. The krgest specimen measured
46 mm. h. d. The distribution of indica b from Somali-Land to the Moluccas in
443-715 fths.
Stereocidaris oapensis.
Stereocidaris indica var. capensis Doderlein, 1901, Zool. Anz., 23, p. 19.
Stereocidaris capensis Doderlein, 1906, Deutsche Tiefsee Exp. Ech., p. 110.
Plate 10, flgrs. 3-6, Deutsche TIeftoe Exp. Ech., Doderlein, 1906.
Altliough closely related to the preceding species, Doderlein considers the
South African form entitled to specific rank. As he finds the chief and most
constant character in the calcareous plates of the pedicels, the species seems
to me open to serious doubt, for I do not consider that any importance can •
be attached to the exact form of the microscopic, calcareous particles of the
Echini. The only known specimens of capensis were taken by the " Valdivia "
CLABK: THE CIDABIDAR. 219
off Gape Colony in 278 fths. The largest measured 36 mm. h. d. The color
is gray, with a brownish tinge, the secondaries with darker tips, and the actinal
primaries whitish.
Stereocidaris trioaxinata.
Stereoddaris indica yar. tricarinata Doderlein, 1901, ZooL Anz., 23, p. 20.
Stereocidaris tricarinata Doderlein, 1906, Deutsche Tiefsee Ezp. Ech., p. 112.
Plato 9, Dentsehe TIeftee Exp. Ecli., Duderleln, 1900.
This species seems to be rather better defined than eapensis, bat as its validity
depends largely on the value assigned to certain features of the pedicellariae, there
is still room for some doubt as to its proper standing. The deformed specimen
to which Doderlein has given the varietal name ieretitpina is indeed very different
from the typical form, but as it was a parasitized individual, its peculiarities may
be pathological. The '* Valdivia " collected tricarinata only in the vicinit][, of
Sumatra in 206-417 fths. The largest specimen was 64 mm. h. d. The color
of the test is dark reddish ; the primaries are gray with rosy necks ; the actinal
primaries whitish ; the larger secondaries have a dark spot.
Stereocidaris alcocki.
Dorocidaris alcocki Anderson, 1894, Joum. Asiat Soc. Bengal, 63, pt. 2, 3, p. 191.
Flato ti, flffs. 3, 3a, HL Inyestiflrator Zo81. Eoh., Aloock and Andenon, 1895.
There can be little question of the validity of this species unless indica proves
to be even more variable than is supposed. If the published descriptions are
accurate (and there is no -apparent reason for doubting them), the two species are
quite distinct. The " Investigator " took alcocki in the Laccadrve Sea in 636
fths. It is a small species, only 25-26 mm. h. d.
Stereocidaris ingolfiana.
Stereocidaris ingolfiana Mortensen, 1903, Ingolf-Exp. Ech., 1, p. 88.
Plato 6, flfft. 1-5, 11, Insolf-Sxp. Ech., Mortonsen, 1903.
It is rather curious that this very distinct and interesting species should not
have been described until so recently, for adult specimens are easily distinguished
from any other North Atlantic or West Indian species. Even when the primary
spines are missing or do not have the " wings " developed, the species may be
recognized by the very numerous slender secondaries and miliaries, and the more
or less elevated median ambulacral area, densely covered with minute tubercles.
Mortensen' s description lacks nothing, but in the table of measurements it is
evident that " height" is estimated in some variable way ; for while in a large
series of tests of such a variable species as D. papillata, for example, there is
sometimes a variation of 20% in the vertical diameter, Mori»nsen's measure-
ments would indicate a variation of 30% amoug 8 specimens of ingolfiana; and
while papillata is occasionally .76 h. d. in height, Mortensen gives one speci-
220 bulletin: museum of compabative zoology.
mea of ingolfiana over .90 b. d., or, in other words, almost spherical ! The
specimens in the Mosenm of Comparative 2iOologj are .54-.58 h. d., while Mor-
tensen's table gives .61-.91 h. d. as the range for bis 8 specimens ; it can hardly
be doubted that this difference is dae to the method of measurement used. In
the diameter of the abactinal system and the actinostome, Mortensen's figures,
.41~.54 h. d. for the former and .36 -.40 h. d. for the latter, accord well with the
measurements of the specimens in the Museum of Comparative Zoology. One
error in his table occurs which may be either a slip of the pen or a misprint ; the
specimen 28 mm. h. d. is said to have the abactinal system only 10.5 (the same as
the actinostome), while examination of the figure given on Plate 6 (which is appar-
ently that specimen) shows the abactinal system to be about 14 mm., which is
what would be expected. The largest specimen recorded is 35 mm. h. d. ; the
color is brownish, but not at all distinctive. The geographical range is from
Iceland to Nevis, in 165-665 fths.
Stereooidaris microtuberoulata.
CtdariM (Stereocidcuris) microtuberculata YoBhlwara, 1898, Ann. ZooL Jap., 2, pt. 2, p. 57.
Plates 1 and 9.
Although this species is closely allied to the following, it is easily distinguished
by the characters given in the key. The test and small spines are yellowish-
brown with a greenish tinge, and the larger secondaries have a median, longitudinal
stripe of a darker shade. The fully developed primaries, when clean, are white.
This is the biggest member of the genus, the diameter of the largest known
specimen being 86 mm.
Stereooidaxis grandis.
DoTocidaris grandis Doderlein, 1885, Arch. Naturg., 51 Jhrg., 1, p. 77.
Stereocidaris grandis Doderlein, 1887, Jap. Seeigel, p. 42.
Plate 1, Plate 2, fiffs. 1-11, Jap. Seeigel, Doderlein, 1887. Plates 33, 36, Haw.
Pao. Eeh. Old., A. AkamIc and Clark, 1907.
The series of specimens at hand from Japan and Hawaii shows that this is a
well-characterized but somewhat variable species. The primaries are quite stout
(the thickness 5-7% of the length), usually deep pinkish, especially at base,
but often brown, gray, or green, while the test is gray, yellowish, or greenish,
and the secondaries yellowish or greenish, often with a broad, longitudinal green
stripe; the general effect is grepnish, more or less inclined towards yellowish.
The largest specimen in the sqries is 40 mm. h. d., but Doderlein's largest
specimen was 61 mm. Specimens of grandis are known not only from Japan and
Hawaii, but also from the Dutch East Indies (de Meijere :04). It is possible
that those to which de Meijere refers as having "die Habe" **hell violet" are
really to be referred to the next species.
' GLABE: THE GIDARIDAE. 221
Sbereocidaris leuoaoantha.
Stereocidaris leucacantha A. Agassiz and Clark, 1907, Haw. Fac. Ech. Cid., p. 28.
.Plates 15, 32, Haw. Pac. Ech. Cld., A. AKassls and Clark, 1907.
Although this Hawaiian species, collected at a number of stations by the
" Albatross,'* is very close to grandis in many ways, the two are easily distin-
guished at a glance, and no intermediate specimens have been seen. The largest
specimen is 67 mm. h. d. The color is somewhat variable, that of the test and
secondaries ranging from almost yellowish-white to deep purplish-gray ; there is
usually a decidedly purple cast actinally. The primaries are longer and more
slender than in grandis (the thickness only 4 or 5% of the length), and are
white when clean. The fully grown ones almost always have the collar deep
purple, sharply contrasted with the white neck. In many specimens the seconda-
ries show an evident green tinge.
Stereocidaris sceptriferoides.
Cidaris (Stereocidaris) sceptriferoides Doderlein, 1887, Jap. Seeigel, p. 6.
Stereocidaris sceptriferoides Doderlein, 1887, Jap. Seeigel, p. 42.
Plato ^3, figs. 13-17, Jap. S«eleel, Doderlein, 1887.
This species, although it appears to be very rare, is well characterized. The
globiferous pedicellariae are very slender, the valves often have a conspicuous
end-tooth, and the opening may be very long a id narrow. The only known
specimens of this species were taken in Japanese waters.
ANOMOCIDARIS.
Anomocidaris A. Agassiz and Clark, 1907, Haw. Pac. Ech. Cid., p. 30.
Test rather flat, vertical diameter about .50 h. d., but sometimes, through eleva-
tion of abactinal system, conspicuously rounded-conical; vertical diameter from
centre of anal system in such cases being about .60 h. d. ; coronal plates 7~9 ; areo-
lae abactinally small, very shallow and indistinct, on the uppermost plates practi-
cally wanting, but at ambitus and below deeply sunken and merging together near
actinostome ; median interambulacral area covered with small tubercles, not at all
bare or sunken, but sutural lines distinct ; ambulacra about .80 of interambulacra ;
poriferous zones not deeply sunken ; median ambulacral area with two or tiiree
series'of tubercles on each side, inner much smaller and more or less incomplete;
vertical sutural line usually distinct; pores nearly horizontal; distance between
two not quite equal to diameter of pore. Abactinal system moderate, about .47 h. d. ;
anal system small, less than .40 of abactinal system and composed of only about
"^u plates and grains ; ociiTars rather small and genitals very widely in contact with
each other. Whole abactinal surface more or less densely covered with very small
secondaries, miliaries, and pedicellariae. Actinostome small, .85 h. d., only about
.75 of abactinal system. Primary spines slender, 1-1.50 h. d. ; thickness 8-5% of
length; cylindrical with longitudinal series of minute granules, sometimes nearly
smooth, often flattened and widened at tip ; actinal primaries very variable, some-
222 BULLETIN: MUSEUM OF COMPARATIVE 'ZOOLOGY.
times flattened, cunred, and entire, slightly notched or eren serrate, hot freqnentij
thick, straight, atad more or less smooth ; secondaries flat, those on ambulacra quite
narrow. Large globiferous sometimes, and tridentate pedicellariae always, want-
ing; small ones sometimes with, more often without, end-tooth on ralTCS.
The above diagnosis of this interesting monotypic genos is based on a large
series of specimens, 11-40 mm. h. d., which admits of little question of the iden-
tity of Doderlein's St. japomea and Yoshiwara's C. tenuispinus. Some of the
peculiarities are given by those writers in their original descriptions of the only
species, which they regarded as a Stereocidaris. While its nearest relatives are
probably to be found in that genus, it is quite distinct from them and is' well
entitled to generic rank. For a full discussion of this genus and its type species,
see A. Agassiz and Gkrk, 1907, Bull. M. C. Z., 51, p. lld-114.
Anomocidaris Japonioa.
Dorocidaris japonica Doderlein, 1886, Arch. Naturg., 51 Jhrg., 1, p. 76L
Stereocidaris japonica Doderlein, 1887, Jap. Seeigel, p. 34.
Cidaris {Stereocidaris) tenuiapinue Yoshiwara, 1898, Ann. Zool. Jap. 2, pt. 2, p. 67.
Anomocidaris tenuispina A. Agassiz and Clark, 1907, Haw. Pac. £ch. Cid., p. 30.
Anomocidaris japonica. A. Agassiz and Clark, 1907, Prelim. Rep. Albatross 1906
Ech., BuU. M. C. Z., 51, p. 112-114.
Plate 31, figs. 5-8, Haw. Pac Ech. Cid., A. Agassis and' Clark, 1907. Plate 3,
Jap. Seeigel, Doderlein, 1887.
The only known specimens of this species have been taken in Japanese waters^
in 40-284 fths. The largest specimen is 40 mm. h. d. The color of test and sec-
ondaries is commonly some shade of brown, often reddish, sometimes greenish,
while the primaries are grayish or brownish, often with a decidedly olive-green
tinge, rarely rosy-reddish ; the neck is brown, usually polished and shining.
ACANTHOCIDARIS.
Acanthocidaris Mortensen, 1003, tngolf-Ezp. Ech., 1, p. 21.
Test high, .60:-.70 h. d. ; coronal plates 7 or 8; areolae not at all sunken and very
distinct, even actinally ; medlati interambiilacral area somewhat sunken and bare
along vertical suture ; ambulacra about .26 of interambulacra ; poriferous «one8
little sunken ; each ambulacral plate slightly curved, with a single large tnbercle
near upper margin of median portion, a much smaller one near lower margin half-
way to inner end, and a very minute one (which usually carries a pedicellaria) just
beneath largest ; this arrangement is remarkably constant, regardless of age and
size; it is well shown in a specimen 9 mm. h. d., and is not essentially different in
one 62 mm. h. d. ; in some very large specimens, however, another small secondary
tubercle may be borne on inner end of plate ; median vertical suture usually visible,
bnt there is no noticeable median bare strip ; pores oblique much as in Cidaris.
Abactinal system about .45 h. d., very flat ; peculiar in that all oculars are broadly.
CLABK: the CIDA&IDAE. 223
in contact with anal plates except right anterior one ; this ocular is wholly or very
nearly excluded ; instead of being an individual peculiarity (as sometimes occurs
in Tretocidaris et al.), this curious arrangement is remarkably constant, and is as
evident in a specimen 17 mm. h. d. as in those over 40 mm. Actinostome .35-
.40 h. d., generally about .90 of abactinal system. Primary spines unique, 2.6-
3.3 h. d., straight or somewhat curved, nearly smooth;' base broad and depressed,
somewhat triangular in cross-section, with more or less evident traces of Icttagitudi-
nal series of granules, but in large specimens these are scarcely visible; collar
enormously wide, .20 or more of length of spine, and abruptly contrasted with
remainder in color ; this remainder bears 10-20 sharply distinct longitudinal ribs,
which are seemingly continuations of series of granules on collar; outer limit of
collar not straight, t. e, forming a ring around spine, but more or less deeply
concave on both sides, especially actinally ; tip of primary blunt or more or less
expanded ; actinal primaries conspicuously capped and serrate as in Stephanoci-
daris, but much stouter than in that genus ; secondaries long, slender, and flat. All
three kinds of pedicellariae present; globiferous, both large and small, lack an end-
tooth on valres ; stalks of large ones usually with a " limb."
This notable genus will be recognized at first sight by the peculiar, handsome
spines somewhat resembling those of Coelopleurus. The above diagnosis is based
upon the examination of fifty fine specimens of hastigera, representing all ages.
The type of the genus is the species named by Bell (*93) curvatiapinisj but nothing
is known of its test or abactinal system, for neither Bell nor Mortensen (: 03)
has attempted any description beyond spines and pedicellariae. It is interesting
to find that the ''Siboga" collected in the East Indies a third species of this
genus, which de Meijere (: 04) has named Cidaris maeulicollu. His careful de-
scription of the primary spines leaves no doubt as to the proper relationship of
this new form, although the describer, in spite of the primaries, places it in the
same subgenus with C. metularia, tribuloides, etc., because he considers the large
globiferous pedicellariae like those of Cidaris. As a matter of fact, however, the
valve of a pedicellaria which de Meijere figures is quite as near Acanthocidaris as
it is to typical Cidaris. On account of the broad collar and the serrate actinal
primaries, de Meijere (:03) originally described maeulicollis as a Porocidaris,
but it really has as little in 9ommon i^th tliat genus as with Cidaris.
Key to the Species.
Collar 'of primary spines very light-colored, unspotted ; remainder of spine
reddish or brownish.
Secondaries cream-color or yellowish ; base of primaries with distinct
angles, which may be somewhat serrate * curvatispinis
Secondaries dark reddish-brown ; base of primaries with rounded angles,
not in tlie least serrate hastigera
Collar of primary spines greenish, with red spots ; remainder of spine whit-
ish with 3 or 4 cross-bands of reddish macuUicollis
224 BULLETIN : MUSEUM OF COMPARATIVE ZOOLOGT.
Aoanthocidaris curvatispixiis.
Cidarii eurvatispinis Bell, 1893, Trans. Zool. Soc., London, 13, p. 303.
Aeanthocidaris curvatispinis Mortensen^ 1908, Ingolf-Exp. Ech., 1, p. 29.
Plate 38, Trans. ZooL Boo. lA>ndon, 13, Bell, 1893.
Nothing is known in regard to this species, except that Bell has figured the
entire animal and Mortensen the pedicellariae. The type specimen in the British
Museum, and a second specimen in Paris, are both from Mauritius and are
the only ones known. The type specimen is about 50 mm. h. d., with primaries
150 mm. long; many of the latter are banded near the tip with brownish
and yellowish.
Acanthooidaris hastigera.
Aeanthocidaris hastigera A. Agassiz and Clark, 1907, Haw. Pac. Ech. Cid., p. 39.
Plates 37-4iB> Haw. Pao. Ech. Cld., A. Agassla and Clark. 1907.
In addition to the differences mentioned above, this species may be distinguished
from the preceding by the stouter primaries and their entire lack of any cross-
barring or bands of color. It was found by the " Albatross " to be common
among the Hawaiian Islands. When cleaned, the test is nearly white in young
specimens, with the median ambulacral area red, the actinostome decidedly green,
and the abactinal system dull greenish-red ; in older specimens the white is re-
placed by reddish-cream color, and there is little green evident anywhere. When
uncleaned the test is, like the secondaries, dark brownish-red, much lighter in
very young specimens. The largest specimen is 52 mm. h.d. ; the longest
primaries are 145 mm. All of the '* Albatross " specimens were, fiiken on sandy
bottom in comparatively shallow water, 23-222 fths.
Aoanthocidaris xnaoulioollis.
Porocidaris maculicoUis de Meijere, 1903, Tijdsch. Ned. Dierk. Vereen. (2) 8, p. 1.
Plate 3» flffs. 18, 19, SIbogra-Ezp. Eoh., de Meljere, 1904.
The secondaries of this species are described as having " a dark longitudinal
stripe," but the ground color is not mentioned. The four specimens collected
by the '*Siboga'* were all small (10-18 mm. h.d) and were evidently young
ones. They were taken at depths of only 39-53 fths , and at each of the
three stations mussel-shells formed a characteristic feature of the bottom.
POROCIDARIS.
Porocidaris Desor, 1854, Syn. Ech. Foss., p. 46.
Test rather high, .60-. 76 h. d. ; coronal plates, 7-9; areolae more or less sunken
and merging actinally ; median interambulacral area with vertical sutural region
somewhat sunken and bare ; ambulacra .18-.34 of interambulacra; poriferous zones
very little sunken; median ambulacral area with a single marginal row of tubercles.
CLARE: THE CJDARIDAE. 225
and even this may be incomplete in small specimens ; bet\¥een are more or fewer
scattered tubercles, but there is nerer a complete second series e^en in yery large
specimens; vertical sutaral line, bare; pores oblique, close together, surface of
interval rough or elevated. Abactinal agrstem variable in size, oculars and especially
genitals with noticeably wide bare margins. Actinostome .30-.45 h. d., with few or
no interambulacral plates. Primary spines, when fully developed, long, 1.5-4 h. d.
cylindrical or nearly so, white (sometimes tinged with rose, purple, or yellow) with
a darker collar; actinal primaries flat, somewhat curved, coarsely and sharply
serrate ; secondaries flat and not peculiar. No globiferous pedicellariae whatever ;
tridentate pedicellariae very variable in size (.30-6.0 mm.) and form, with 2-4
(generally 8) unusually stout, wide valves.
This is one of the most distinct and easily recognized of the genera of recent
Echini, but the species it contains are most perplexing and are exceedingly diffi-
cult to distinguish from each other. The genus has a wide geographical range, as
it occurs in the North Atlantic Ocean, the Caribbean Sea^^ among the Gaiapagog
Islaiidsi the Hawaiian Islands, tbe ^ast Indian Islands, and the Nicobar Islands,
.along the coast of Japan, near Australia, and along the east coast of Africa, it
depths ranging from 16d to 799 fths. Several species from tbe Tertiary have
been named, and serrate spines, like the actinal primaries of Porocidaris, occur in
the Jurassic. There is little diversity of color in the genus, for the test, the collar
of the primaries, and the small spines are commonly some shade of brown, often
becoming very dark or deep purple with age, while the primaries are usually very
white. The following key is based on the examination of 54* specimens representing
all the species, except misakiensis.
m Key to the Speoiea
Pedicellariae all with 2 valves purpurata
Pedicellariae mostly with 3 valves.
Abactinal system .40-.56 h. d. ; primaries rather stout (thickness of large
ones 3-6% of length), finely and sharply thorny. (These prickles
are not always easily seen with the unaided eye, but are so distinct
that a spine cannot be drawn upward between thumb and finger
when lightly closed upon it.)
Small spines in interambulacra, outside scrobicular circles, above am-
bitus, very few ; ambulacra almost wholly bare between marginal
rows of tubercles ; primaries stout, 1.5-2.5 h. d. (thickness 5-6 per
cent of length), often becoming larger and fluted near tip, with
numerous (25-30) longitudinal series of prickles sharreri
Small spines more numerous on upper half of test ; ambulacra usually
with scattered tubercles ; primaries somewhat less stout, with
about 12-15 longitudinal series of prickles, more or less tapering
and never enlarged and fluted at tip, but occasionally with large
projecting thorns near base.
Primaries less stout (thickness 3-4 per cent of length) ; no special
depression on inner surface of valves of large pedicellariae above
hypophysis ; test, secondaries, and collar of primaries light red-
dish- or yellowish-brown degans
VOL. LI. — NO. 7 15
226 bulletin: museum o^ comparativb zoology.
Primaries stouter (thickness about 4.6 per cent of length) ; a distinct
triangular impression on inner surface of yalves of large pedi-
cellarise aboTe hypophysis; test, secondaries, and collar of
primaries deep, dark brown muakienns
Abactinal system .d5-.46 h. d. ; primaries rery long and slender, 2.6-4 h. d.
(thickness only 2^ per cent of length) ; very nearly smooth (slip
easily between thumb and finger).
Large pedioellariae always with 8 Talres, which are distinctly pointed ;
anal system about .60 of abactinal ; median ambulacral area about
.37 of ambulacrum ; size small, under 36 mm. h. d. ; color pale, and
primaries yery white and shining coboti
Large pedioellariae very rariable, sometimes with only 2 or with 4
Talves, wliich are usually broad and are rounded at tip; anal sys-
tem about .46 of abactinal ; median ambulacral area about .60 of
ambulacrum ; suee large, up to 86 mm. h. d. ; color usually very
dark and primaries yellowish variabUU
PorooidariB purpurata.
Poroeidaris purpwuta Wyrille Thomson, 1872, Ann. Mag. Nat. Hist, (4) 10, p. 302.
Plato 59, Porcupine Ech.» WyriUe Thompson, 1875.
One needs only to compare a specimen of this cidaroid with any other member
of the genus to reject Mortensen's ( :03) proposed genus *' Histocidaris/' for
aside from the pedicellariae, the only feature in which purpuraia differs noticeably
from the others is the presence of an exceptionally wide collar on some of the
primaries of some specimens, and that can hardly be considered a very useful
character. Moreover Mortensen's proposed variety talismani, which he thinks
may even be a distinct species, cannot be recognized, for the primaries with
swollen, fusiform, violet collars occur in typical purpuraki^ and one is figured by
Thomson ('75), though they are not present in all specimens. The small spines
and some of the abactinal primaries are light reddish- or purplish-brown. The
largest recorded specimen is 60 mm. h. d. This species is known only from
the North Atlantic, save for the specimen from the Nicobar Islands, collected by
the *' Valdivia," and referred to purpurata by Doderlein.
Porooidaris sharreri.
Porecidaris Sharreri A. Agassiz, 1880, Bull. M. C. Z., 8, p. 7L
Plato 3y Blake Ech., A. Asassiz, 1883.
This handsome West Indian species was dredged by the ** filake " off Georgia
in 279 fths. (in company with St^ ingolfiana ) and also near Barbados in 356 fths.
The general color is red-brown and not at all purplish. The largest specimen is
69 mm. h. d., with spines 114 mm. long.
CLARK: THE CIDARIDAE. 227
Porocidaris elegans.
Porocidaris elegans A. Agassiz, 1879, Proc. Amer. Acad., 14, p. 198.
Plate 3, ChaHenger Ecn., A. Agassiz, 1881.
Originally collected by the " Challenger " oflF New South Wales and southeast
from the Philippines, specimens of Porocidaris, referred to this species, have since
been taken by the " Valdivia " near Sumatra, and off the east coast of Africa, and
by the " Siboga " among the Dutch East Indies. One of the specimens collected
by the latter vessel measured 85 mm. h. d. The specimen from the Bay of Biscay
reported by Koehler ('96) is doubtless not this species ; but probably purpura ta,
though it might be sharreri, with which species elegans agrees in coloration and
many other points. The 5 specimens taken by the '* Siboga " which de Meijere
( : 04) calls " Cidaris elegans juv. ? " are rather peculiar, especially the pedicel-
lariae, and their real relationship' is doubtful. The specimens taken by the
" Valdivia " differ from elegans^ not only in their remarkably light coloration, but
in their small abactinal system, actinostome and anal system, the very thorny
primaries, and their large number of coronal plates. It is quite likely that they
are a distinct species.
Porocidaris misakiensis.
Cidaris {Porocidaris) misakiensis Yoshiwara, 1898, Ann. Zool. Jap., 2, pt. 2, p. 68.
Plate 2, fiif. 16, Slboga-£zp. £ch., de Me^ere, 1904.
This is the most dubious species of the genus, especially as no complete de.
scription or figures have appeared. Aside from the original preliminary descrip-
tion, the only available information about misakiensis is contained in de Meijere's
" Siboga " report (:04). He found one specimen which might be referred to this
species, but the difference between it and elegans is difficult to understand, and it
will be surprising if the two prove to be really distinct. Yoshiwara's specimen was
39 mm. h. d., and de Meijere's was 50 mm. The color is said to be dark brown.
Porocidaris cobosi.
Porocidaris cobosi A. Agassiz, 1898, Bull. M. C. Z., 32, 5, p. 74.
Plate 9, Pan. Deep Sea Ech., A. Ag^assiz, 1904.
This is the handsomest species of the genus, and except purpurata^ the easiest
to recognize. It has been taken only once, and then by the " Albatross," near
Chatham Island, Galapagos, on a rocky bottom in 385 fths. The largest specimen
is only 35 mm. h. d.
228 BULLETIN: MUSEUM OF COMPABATIVB ZOOLOGY.
Porooidaris Tariabilis.
Poroddaru variabUia A. Agassiz and Clark, 1907, Haw. Pac. Ecb. Cid., p. 32.
Plataa 16-»S» S8, fiss. 1-4, Haw. Pi^ ISch. Old., A. A^amU and Clark, 1907.
This species was found by the "Albatross " to be common among the Hawaiian
Islands, and some yery fine specimens were secured. The largest is deep purple,
and measures 85 mm. h. d. ; the others are various shades of brown, and one was
yery light-colored, like codosi. It is possible that if misakietuU is really distinct
from elegant^ this species may prove to be identical with Yoshiwara's.
CLABK: THE CIDA.RIDAE.
229
INDEX
abjssicola (Dorocidaiis).
208
Diplocidaris.
191
Acanthocidaris.
222
Discocidaris.
175
affinis (Tretocidaris).
203
doederleini (Centrocidaris).
214
africana (Stereocidaris indica).
218
Dorocidaris.
207
alcocki (Stereocidaris).
219
dubia (Fhyllacanthus).
187
Anaulocidaris.
175
dubia (Tretocidaris).
204
annnlata (Tretocidaris).
203
1 /T% t Y • V
animlifera (Fhyllacanthus).
Anomocidaris.
188
221
islegans (Forocidaris).
Eocidaris.
227
175
Aporocidaris.
214
Euddaris.
175
assimilis (Schizocidaris).
198
fimbriata (Goniocidaris).
199
Astropyga.
192
florigera (Groniocidaris).
198
Aulacocidaris.
175
fragilis (Aporocidaris).
216
austndis (Phjllacantlias) .
187
Austrocidaris.
212
galapagensis (Cidaris).
186
geranioides (Goniocidaris).
198
baculosa (Fhyllacanthus).
189
gibberub (Tylocidaris).
177
bartletti (Tretocidaris).
203
gigantea (Chondrocidaris).
191
biseriaHs (Goniocidaris).
199
glandulosa (Stephanocidaris).
194
bispinosa (Stepbanocidaris).
194
Goniocidariens.
168
blakei (Dorocidaris).
209
Goniocidaris.
195
bracteata (Tretocidaris).
206
gracilis (Porocidaris).
197
calacantha (Tretocidaris).
Calocidaris.
205
211
graiidis (Stereocidaris).
Gymnocidaris.
220
175
canaliculata (Austrocidaris).
212
hastigera (Acanthocidaris).
224
capensis (Stereocidaris).
218
hawaiiensis (Stephanocidaris).
195
carinata (Stereocidaris mdica).
218
hirsutispmus (Goniocidaris).
197
Centrocidaris.
214
Histocidaris.
175
Cho]idrocidari&
190
hystrix (Cidaris).
167
Cidaridae.
177
Cidarid^.
168
imperialis (Fhyllacanthus).
188
Cidariens.
168
incerta (Aporocidaris).
216
Cidaris.
183
indica (Stereocidaris).
218
cidaris (Echinus).
167
inermis (Orthocidaris).
177
Cidarites.
167
ingolfiana (Stereocidaris).
219
clypeata (Goniocidaris).
197
integra (Stereocidaris indica).
218
cobosi (Porocidaris).
cren iilaris ( Sclil^nitzia) .
227
187
japonica (Anomocidaris).
222
cretosa (Stereocidaris).
177
Leiocidaris.
175
curvatispiuis (Acanthocidaris).
224
Leptocidaris.
175
230
bulletin: museum of comparative zoology.
leacacantha (Stereocidaris). 221
lorioli (Stereocidaris). 212
lutkeni (PhjUacantLufi). 189
maculicollis (Acanthocidaris). 224
niagnifica (Temnocidaris). 177
metularia (Cidaris). 184
niicans (Calocidaris). 811
microtuberculata (Stereocid-
aris). 220
mikado (Goxdocidaris). 199
Mikrocidaris. 175
milleri (Aporoddaris). 216
Miocidaris. 175
misakiensis (Porodidaris). 227
njortenseni (Austrocidaris). 213
multiceps (Polycidaris). 177
nonarias (Polycidaris). 199
nada (Dorocidaris). 209
nutrix (Aufltrocidaris). 213
Orthocidaria. 201
panamensis (Tretocidaris). 204
papillata (Dorocidaris). 209
Paracidaris. 175
parvispina (Phyllacanthus). 187
perplexa (Tretocidaris). 205
Petalocidaris. 175
Phalacrocidaris. 175
Phyllacanthus. 186
pistillaris (Phyllacanthus). 167
Plegiocidaris. 175
Pleurocidaris. 175
Poly cidaris. 199
Porocidaris. 224
Prionocidaris. 175
Procidaris. 1 75
purpurata (Porocidaris). 226
quoyi (Goniocidaris). 167
recens (Rhabdocidaris). 187
reini (Tretocidaris). 207
reynesi (Tetracidaris). 177
Rhabdocidariens. 168
Rhabdocidaris. 175
rugosa (Dorocidaris). 210
Salenia. 215
sceptriferoides (Stereocidaris). 221
Scliizocidaris. 175
Schleinitzia. 175
serrata (Discocidaris). 198
sharrcri (Porocidaris). 226
spinosa (Tretocidaris). 200
Stephanocidaris. 192
Stereocidaris. 216
sumatrana (Stereocidaris indica). 218
talismani (Porocidaris purpu-
rata). 226
Temnocidaris. 195
tenuispina (Anomocidaris). 222
tenuispinus (Stereocidaris). 222
teretispina (Stereocidaris tricari-
nata). 219
Tetracidaris. 192
thomasii (Phyllacanthus). 188
thouarsii (Cidaris). 185
tiara (Tretocidaris). 202
Tretocidaris. 200
Triadocidaris. 175
tribuloides (Cidaris). 185
tricarinata (Stereocidaris). 219
tubaria (Goniocidaris). 198
Turbans. 167
Tylocidaris. 183
Typocidaris. 175
uiiibraculum (Goniocidaris). 198
variabilis (Porocidaris). 227
veronensis (Porocidaris). 177
verticillata (Phyllacanthus). 187
CuJOL. —Hie CiduidM.
EXPLANATION OF PLATES.
PLATE 1,
8tereocidari8 microtuberculata Yosliiwara. Nat size.
Abactinal view.
>.
\\
Clabx. — The OidazidM.
PLATE 2.
Stereocidaris microtaberculata Yoshiwara. Nat. size.
Side view of same specimen as Plate 1.
Olaol ' TlM Oidadda^
PLATE 3.
Caloddaris micans (Mortensen). Nat. size.
1. Ambalacral view of partly cleaned specimen ; all primarj spines broken
2. Abactinal view of same.
Clabk. — The Cidaridae.
PLATE 4.
Doroddaris rugosa, sp. nov. Nat. size.
Abactinal yiew.
Claul — The Ci<Und«&
PLATE 5.
Dorocidaris rugosa, sp. nov. Nat size.
Actinal view.
PLATE 6.
1-2. Tretocidarifl perplexa, sp. noT. Nat nse.
1. Abactinal yiew.
2. Actinal view.
3-4. Tretocidaris dnbia, sp. nov. Nat. size.
3. Abactinal view of fMirtlj cleaned specimen.
4. Actinal view of same.
ot^A^m ^Xte OldMidis*
i
1
PLATE 7.
1-4. Tretocidaris perplexa, sp. nor. Nat. sin.
1. Abactinal view of partly cleaned 8peciineQ»
2. Actinal view of same. j
8. Interambulacral view of same. "^
4. Ambolacral Tiew of same. '
6-8. Dorocidaris rogosa, sp. dot. Nat sin.
6. Abactinal view of partly cleaned test
0. Actinal view of same.
7. Interambulacral view of same.
8. Ambolacral Tiew of same.
4ift €P
i
PLATE 8.
Tretocidjuru butletti (A. Agmniz). Nat. sue.
Ahai'tinal riew of specimen with cylindrical apine^
:>^
» *
OliAUL — TlM OidaiidM.
PLATE 9.
TretocidariB bartletti (A. Agansiz). Nat. size.
InterambuUcnl Tiew of same Bpecimen as Plate &
PLATE U.
1-S. Tretocidarii bncteata (A. Agiatiz). Nat aiae.
1. Abacdnal riew of putlj deaoed f pectmen*
2. Side Tiew of fame.
8-4. Geniocidarif nmbracolnm HatUm. Nat tiae.
8. Ambulacral Tiew of bare test
i. Abactinal Tiew of same.
6. GoniocidariS' tubaria (Lamarck). Nat size.
Interambalacral Tiew of partly cleaned, small
specimen, with slender spines.
Claik. — The CidwidM.
PLATE 11.
Goniocidarifl tubaria (Lamarck). Nat. size.
1. Abactinal yiew of partly cleaned, large
specimen with short, stout spines.
2. Side Tievr of same.
^<y ^Bwnid.
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