OF THE
UNIVERSITY
OF
LIBRARY
SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
Bulletin 88
REVISION OF PALEOZOIC STELLEROIDEA
WITH SPECIAL REFERENCE TO NORTH
AMERICAN ASTEROIDEA
BY
CHARLES SCHUCHERT
Professor of Paleontology, Yale University
Sew Haven
WASHINGTON
GOVERNMENT PRINTING OFFICE
1915
SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
Bulletin 88
REVISION OF PALEOZOIC STELLEROIDEA
WITH SPECIAL REFERENCE TO NORTH
AMERICAN ASTEROIDEA
BY
' '*"
CHARLES SCHUCHERT
Professor of Paleontology , Yale University
New Haven
WASHINGTON
GOVERNMENT PRINTING OFFICE
1915
PALEONTOLOGY
BULLETIN OF THE UNITED STATES NATIONAL MUSEUM.
ISSUED MARCH 20, 1915.
ADVERTISEMENT.
The scientific publications of the United States National Museum
consist of two series, the Proceedings and the Bulletins.
The Proceedings, the first volume of which was issued in 1878, are
intended primarily as a medium for the publication of original, and
usually brief, papers based on the collections of the National Museum,
presenting newly-acquired facts in zoology, geology, and anthro-
pology, including descriptions of new forms of animals, and revisions
of limited groups. One or two volumes are issued annually and dis-
tributed to libraries and scientific organizations. A limited number
of copies of each paper, in pamphlet form, is distributed to specialists
and others interested in "the different subjects as soon as printed.
The date of publication is printed on each paper, and these dates are
also recorded in the tables of contents of the volumes.
The Bulletins, the first of which was issued in 1875, consist of a
series of separate publications comprising chiefly monographs of large
zoological groups and other general systematic treatises (occasionally
in several volumes), faunal works, reports of expeditions, and cata-
logues of type-specimens, special collections, etc. The majority of
the volumes are octavos, but a quarto size has been adopted in a few
instances in which large plates were regarded as indispensable.
Since 1902 a series of octavo volumes containing papers relating to
the botanical collections of the "Museum, and known as the Contribu-
tions from the National Herbarium, has been published as bulletins.
The present work forms No. 88 of the Bulletin series.
RICHARD RATHBUN,
Assistant Secretary, Smithsonian Institution,
In charge of the United States National Museum.
WASHINGTON, D. C., January 27, 1915.
3
M650670
TABLE OF CONTENTS.
Page.
Preface 9
Terminology as applied to the skeleton of Paleozoic Asteroidea 13
Finding list of Paleozoic Stelleroidea 18
Remarks on the development of Paleozoic Stelleroidea 27
Geological 27
Distribution 27
Preservation and occurrence 28
Asterid evolution 30
General observations 30
The most primitive fossil starfish 31
Origin of the wriggling type of starfish 32
The asterid radicle 34
Ontogeny 36
Regeneration 37
Number and character of rays 38
Development of the parts 39
Anus 39
Madreporite 39
Spinosity 41
Ambulacralia 41
Adambulacralia 43
Inframarginalia 43
Supramarginalia 45
Radialia 46
Accessory plates 47
Ocular plates -^, 48
Centro-dorsal 48
Primary disk-plates 49
Axillary ossicles 49
Interbrachial areas 50
Classification 51
Catalogue of Paleozoic Stelleroidea 51
Description of genera and species 51
Class Stelleroidea 51
Subclass Asteroidea 51
Phanerozonia 51
Family Hudsonasteridae 53
Hudsonaster 53
Siluraster 65
Family Palseasteridse 66
Palseaster , 67
Australaster 72
5
6 TABLE OF CONTENTS.
Description of genera and species Continued.
Class Stelleroidea Continued.
Subclass Asteroidea Continued.
Phanerozonia Continued. p age .
Family Promopalaeasteridae 73
Subfamily Mesopalaeasterinae 74
Mesopalseaster 74
Spaniaster 95
Miomaster 96
Devonaster 97
Subfamily Promopalasasterinse 102
Promopalasaster 102
Subfamily Anorthasterinae 125
Anorthaster 125
Family Xenasteridae 128
Xenaster 129
Agalmaster 131
Rhenaster 133
Eifelaster 133
Trimeraster 134
Family Neopalaeasteridse 134
Neopalseaster 134
Family Palasterinidae 138
Petraster 138
Lindstromaster 148
Palasterina 150
Uranaster 154
Palseostella 155
Pseudopalasterina 156
Family Lepidasteridae 157
Lepidaster 158
Helianthaster 159
Lepidasterella 160
Family unknown 161
Astropecten 161
Ataxaster 161
CryjDtozonia .- 162
r~Family Stenasteridae 163
/ Stenaster 163
Tetraster 167
Family Monasteridaa 170
Monaster 170
Family Urasterellidee 172
Urasterella 173
Family Calliasterellidaa 190
Calliasterella 190
Family Compsasteridse 191
JaeJcelaster 192
Compsaster 192
Family Schuchertiidae 194
Schuchertia 195
Family Palasteriscidae 199
Palasteriscus 200
Echinasterella... 200
TABLE OF CONTENTS. 7
Description of genera and species Continued.
Class Stelleroidea Continued.
Subclass Asteroidae Continued.
Cry ptozonia Continued .
Family Palasteriscidae Continued. Page.
Loriolaster 201
Cheiropteraster 202
Family Schoanasteridae . 202
Schcenaster 202
Family Palseosolasteridae 207
Palxosolaster 209
Echinasterias 211
Echinodiscaster 211
Echinostella 212
Medusaster 212
Subclass Auluroidea 213
Order Lysophiurae 215
Family Protasteridae 215
^fseniaster 216
[_ Eophiura. . 222
Bohemura 223
L-Palseum 223
Protester 224
Akpidaster 228
Gregoriura 233
Bunderibachia 234
Palxophiomyxa 234
Family Palaeophiuridae 235
Palseophiura 235
Sturtzura 236
f-JSygasterella 237
; Ptilonaster 239
Family Encrinasteridse 241
Encrinaster 241
Order Streptophiurae . 246
Family Ophiurinidae 246
Ophiurina 246
Tremataster 247
Family Lapworthuridae 248
Squamaster 249
^Lapworthura 250
r Sturtzaster 252
_ Bdellacoma 254
THhopalocoma 254
Hallaster 254
Sympterura 256
Furcaster 261
Palastropecten 261
Family Eoluidiidae 262
Eoluidia 262
Eospondylus 263
Miospondylus 263
Family Aganasteridae 263
Aganaster 263
Family Cholasteridae 265
Cholaster. . . 265
8 TABLE OF CONTENTS.
Description of genera and species Continued.
Class Stelleroidea Continued. p age .
Subclass Ophiuroidea 267
Family Onychasteridee 268
Onychaster 268
Forms whose relationships are unknown 274
Cribellites carbonarius 274
Order Ophiocistia , 275
Family Eucladiidse 276
Eucladia 276
Euthemon 279
Bibliography 281
Explanation of plates 289
Index.. 303
REVISION OF PALEOZOIC STELLEROIDEA WITH
SPECIAL REFERENCE TO NORTH AMERICAN
ASTEROIDEA
By CHARLES SCHUCHEET
Professor of Paleontology, Yale University, New Haven.
PREFACE
This memoir had its inception in 1896, and came about through
the force of circumstances. The writer, at that tune assistant curator
in the United States National Museum, was made aware of the desire
of Mr. I. H. Harris, of Waynesville, Ohio, to present to that institu-
tion his extensive collection of Ordovicic fossils. He had long known
Mr. Harris, having met him as a young man, about the year 1876,
at which time the Harris collection was already widely known, not
only for its quantity, but more especially for its many choice speci-
mens of the rarer species of starfishes, crinids, and trilobites. Twenty
years later came Mr. Harris's desire to present to the National
Museum the collection upon which he had been at work for fifty
years. This gift brought to the National Museum not less than 40
asterids, 35 (31 on one slab) ophiurids from the Cincinnatic strata
(the great majority from the Waynesville formation), besides an
asterid and 12 specimens of Ony chaster flexilis from the Keokuk
formation at Crawfordsville, Indiana. Some years later the National
Museum acquired the very valuable E. O. Ulrich collection, in which
there were also many good starfishes. Accordingly, it is safe to state
that in no other museum are there so many Ordovicic asterids, in so
great a variety and in such excellent preservation.
The material of the Harris collection seemed to show that its com-
plete study would not only reveal much new knowledge, but give an
insight as well into the probable evolution of the Paleozoic starfishes.
When these facts were made known to the then director of the National
Museum, Dr. G. Brown Goode, he asked the writer to invite Mr. Harris
to join him in a work on the Ordovicic starfishes. This was done and
Mr. Harris was greatly pleased to undertake the joint authorship.
He did all he could to further the work, but it went very slowly,
because the present writer was so much occupied with curatorial
work as well as the arranging of museum exhibits for the expositions
in which the United States Government participated and which came
9
10 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
nearly every year. Unfortunately, Mr. Harris died on October 17,
1897, when the work was still in the first stages.
At odd times during subsequent years the writer kept up his studies
and in the spring of 1904 the work on the Asteroidea was practically
completed. Then came his appointment to the chair of paleontology
at Yale University. Here new duties and new lines of thought kept
him away from the study of the Paleozoic ophiurids which had been
planned, but the hope was strong that somehow the time would be
found to complete the work. However, this hope has not materialised,
others have gone to work on the starfishes, and one genus and general-
ization after another has slipped away. Therefore it was thought
best to revise what had been done, bringing the asterids up to date.
In regard to the ophiurids, it was found impossible to restudy the
specimens, but as the writer had kept abreast of the literature, this
was put into order and is here presented, in the hope that subsequent
workers will find it useful in enabling them to find the references on
any species and the taxonomic status of any form.
A study of the specimens in the Harris collection made it plain
that no satisfactory taxonomic and systematic results would follow
unless the undertaking also embraced many other Paleozoic forms,
of Europe as well as of America. Species and genera had been pro-
posed in a very loose manner; in fact, most of the Paleozoic litera-
ture on asterids and ophiurids up to the tune this work began had
no other value than to force the reviser under the rules of nomencla-
ture to recognize the names proposed. The International Rules of
Zoological Nomenclature have here been strictly followed. Under
these circumstances, all of the type-specimens had to be seen, and most
of them have been accessible. In this way the work finally came to
be extended to all Paleozoic Stelleroidea. To make it plain how little
the American Paleozoic starfishes were known in 1889, it will suffice
to give one example: Miller in his North American Geology and
Palaeontology lists 23 species under Palseaster'j in the present work
there is but a single form of this genus, the genotype, P. niagarensis,
the remainder having been distributed among seven genera, all new
and proposed in the past few years or at this tune. As a result, every
paleontologist will be obliged to disregard what he has learned re-
garding American Paleozoic starfishes and start anew. To facilitate
this, a list has been prepared and is here presented of all the names
so far given to Paleozoic asterids and ophiurids, with cross references
to the names adopted in this work.
Now that this study is finished, at least for the present, it is plain
to the author that his species and genera are also in some cases too
comprehensive. The future student will restudy the specimens along
with the new material and go more deeply into the detailed structure
of the parts. He who attempts this, along with painstaking recon-
EEVISION OF PALEOZOIC STELLEROIDEA. 11
struction of the skeleton as it was in life, will learn where to separate
and what to reject.
During the past 10 years, Schondorf has revised most of the German
asterids and has studied deeply into the structure of the ophiurids.
His work is very detailed and the best extant on the Paleozoic forms.
Every student of Stelleroidea intending to do thorough work will
have to examine his results. In classification he has laid the founda-
tion, and has shown that some asterids are not Asteroidea at all, but
that they and the bulk of the so-called ophiurids of the Paleozoic must,
because of their peculiar structure, ambulacral and otherwise, be
referred to another class, the Auluroidea.
In regard to the evolution of the Asteroidea, the writer hoped
to find the time to present his views concerning it in detail in this
memoir, but that also is not possible. However, the main lines
and often the generic directions of organic change are stated and
will be found either in the introductory pages or scattered through-
out the work, generally in the remarks under the generic discussions.
These results in connection with the work of Schondorf, it is thought,
ought soon to place the Paleozoic Stelleroidea in such order that
a final classification, based on ontogeny, chronogenesis, and phyl-
ogeny, and embracing not only the Paleozoic forms but those of
Mesozoic, Cenozoic, and Recent time as well, can be made.
Many museums and individual paleontologists have loaned the
writer the specimens in their collections or in their keeping, and he
wishes here to thank them, one and all, for these loans. The greatest
bulk of the material studied is of course in the United States National
Museum in the Harris and Ulrich collections; a number of Trenton
specimens and some from the English Siluric have also been given
to the National Museum by Mr. Walter R. Billings, of Ottawa.
Since the writer has been in New Haven, Dr. R. S. Bassler, of the
National Museum, has often kindly helped him to further his studies,
and during the past year has made a large number of photographs
of the specimens under his ' charge. The many Mississippic speci-
mens in the Frank Springer collection have not been studied.
The author is greatly indebted to Dr. F. A. Bather, of the British
Museum (Natural History), for a large number of gutta-percha and
wax squeezes made by him of certain species in that great museum.
These casts, which are all in the United States National Museum,
have enabled the writer to understand several genera that otherwise
he could not have worked out. He is also indebted to him for many
bibliographic corrections and suggestions.
The second largest American collection is at Harvard University,
in the Museum of Comparative Zoology. Most of the material is
in the Charles D. Walcott, Charles B. Dyer, and Charles Wachsmuth
collections. In the first-named collection there is excellent material
from the Middle Ordovicic; the second has many fine specimens
12 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
from the Cincinnatic formations; while the Wachsmuth material has
species from the Mississippic. To all of this material the author
has had access through the kindness of Director Samuel Henshaw
and Profs. R. T. Jackson and P. E. Raymond.
At the University of Chicago there is a great deal of excellent
material in the W. F. E. Gurley, Charles Faber, and U. P. James
collections, to which Weller has added other specimens. All of this
material has been studied through the courtesy of Prof. Stuart
Weller. Most of it is from the Cincinnatic formations.
All of the specimens described by E. Billings and now in the Vic-
toria Memorial Museum in Ottawa, Canada, have been seen at one
tune or another. In this matter the writer was much aided by the
late Dr. J. F. Whiteaves and more recently by Drs. P. E, Raymond
and E. M. Kindle, to the latter of whom thanks are also due for
several photographs reproduced in this memoir. Most of the speci-
mens are from the Ordovicic.
In the University of Toronto there is much material from the
Ordovicic and Siluric in the Sir Edmund Walker collection, and there
may be studied a great many specimens of Stenaster salteri. All of
this has been seen through the great liberality of Prof. William A.
Parks.
The genotype of Pakeaster, from the museum of Cornell University,
was loaned to the author at different times by Prof. H. S. Williams.
It is one of the best preserved of Paleozoic specimens and has become
very valuable because of the continual discussion that centers
around the species, P. niagarensis.
To all the specimens in the Peabody Museum of Yale University
access was had through the kindness of the late Prof. Charles E.
Beecher. Among them is the unique branching form, Eucladia
beecfieri.
Photographs have been made for this memoir by Dr. John M.
Clarke of specimens in the New York State Museum, and by Prof.
George H. Hudson, of Plattsburg, New York, of other material.
Prof. Arthur M. Miller loaned two specimens that he collected and
deposited in the State University, Lexington, Kentucky. Prof. George
W. Harper, of Cincinnati, loaned a specimen of Palseaster darkana
( = Hudsonaster incomptus) .
Nearly all the original drawings were traced from the specimens
by aid of the camera lucida. These outlines were then inked and
shaded by the late Dr. J. C. McConnell, a draftsman and artist of
the first rank.
To Miss Clara Mae LeVene the writer is especially thankful for
the great interest she has taken hi getting the manuscript into order
for the printer, keeping the bibliography up to date and correctly
citing it, and for the careful reading of the proof.
REVISION OF PALEOZOIC STELLEBOIDEA. 13
TERMINOLOGY AS APPLIED TO THE SKELETON OF PALEOZOIC
ASTEROIDEA. 1
Abactinal.
See Body-wall. The integumentary skeleton opposite to that having the ambu-
lacral grooves, or that of the upper or dorsal side of the animal. Also known
as the aboral surface. See Actinal.
Aboral.
See Abactinal.
Accessory plates.
This term refers to the abactinal longitudinal columns, or transverse rows, of small
plates between the radials and supramarginals (=radial accessory plates or dorso-
laterals), or between the inframarginals and supramarginals (=ambital plates or
intermarginals), or where there are no radials, to all the ossicles between the two
supramarginal columns. There may also be accessory actinal plates which are
termed accessory interbrachials, accessory actinals, or inter actinals. The accessory
plates form the secondary skeleton and are inserted between the columns of
the primary skeleton. It is in the secondary skeleton that much of the
specific and generic evolution takes place. See Interbrachial.
Actinal.
See Body-wall. The integumentary skeleton with the ambulacral grooves, or
that of the under or ventral side of the animal. Also known as the oral surface.
Adambulacrals.
The single columns of prominent plates adj oining the ambulacrals . There are never
more than two of these columns in a ray, one on each outer side of the depressed
ambulacrals. They are generally large and thick and bear the larger, or adam-
bulacral, spines . These spines may be inwardly directed and serve as a protection
for the tube-feet. The oral armature in Paleozoic species usually consists of five
pairs of modified adambulacrals, each pair being the oral projection of two united
columns of adjoining rays. They are also called Adambulacralia. Also see Inter-
brachial.
Ambital.
The abactinal space between the infra- and supramarginal columns, occupied by
accessory plates. These are best developed in the axillary disk areas (disk ambi-
tals), while those of the rays are referred to as ray ambitals. They are also
known as intermarginals.
Ambulacral.
Ambulacral grooves (five or more), plates, or ossicles are situated actinally and per-
radially along the center of the rays and disk. They are also called ambula-
cralia. The ambulacral columns are in pairs, the interlocking movable plates
of adjoining columns may be alternate or opposite (always so in recent forms),
and the plates of a column may be superposed like the tiles on a roof. The
podial openings through which the fleshy tube-feet protrude are situated be-
tween the sutures of adjoining plates. In early forms there are but two columns
of podial openings in each ray, one on each side of the axial line, but in later
forms there may be as many as eight columns. The ambulacral grooves may
be tapering or petaloid.
Anal opening.
The only Paleozoic form in which an anal opening may exist visually is Hudson-
aster. Here it is on the disk between the central plate and the madreporite.
The anal pore is nearly always obscured in fossil starfishes, and is absent in some
recent species.
1 Also in the main applicable to the Auluroidea and somewhat to the Ophiuroidea.
14 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Arcs.
See Axil.
Axil.
The angle formed by the junction of the rays. Also known as arcs. In primitive
forms each axil is occupied by a single plate, the axillary inframarginal. In
others this plate is crowded orally and its place occupied by two prominent
proximal inframarginals. In forms with large disks having well-developed
marginals these are termed disk marginals. Also see Ambital and Interbrachial.
Bivium.
The two rays of a starfish that inclose between them the madreporite. When
oriented toward the student, the ray on the left is numbered 1 and the other on
the right is numbered 5. A line drawn through the madreporite and the center
of the upward directed ray (ray 3) divides the animal into symmetrical halves
(=left and right sides). The upper left-hand ray is numbered 2 and the one
opposite is 4. Rays 2, 3, and 4 form the trimum.
Body-wall.
The outer surface of both disk and rays, made up of movable calcareous plates
which are either closely adjoining or form a more or less loose network of ossicles.
These are held together in the living animal by connective tissue and muscular
fibers, which after death soon decompose, freeing the plates. It is for this reason
that fossil starfishes are so rarely found entire. Imbedded in the body-wall
everywhere are blunt, short, calcareous spines, arranged in a more or less
definite order, and often movable upon the underlying plates.
Covering the whole surface of the body, including the spines and pedicellarise,
is a delicate membrane or skin, clothed externally with closely placed vibratile
cilia, which keep the integument clean. The calcareous skeleton of starfishes
is therefore integumentary.
Csecal pores.
The openings between reticular plates for the protrusion of the respiratory csecal
processes. These pores are either restricted to the abactinal area or may be
distributed over the entire body. The processes are also known as papulx.
Central disk.
See Disk.
Central disk plate.
A prominent plate, most conspicuous in primitive forms, which often appears in
the center of the disk. It holds the same position as the centro-dorsal plate in
embryonic crinids of the genus Antedon. Also called the abactinal-central plate
or simply central plate.
Centro-dorsal plate.
See Central disk plate.
Disk.
The central portion of a starfish. In this work, the term generally applies only to
the abactinal central area formed either by the union of the rays, or by the rays
plus the axillary areas. The mouth is in the center of the disk on the under or
actinal side . Disk plates are the small plates over this area on the abactinal side .
Central disk plate refers to a larger primary plate in the center of the disk. There
may be other large primary plates, but these are more properly referred to the
rays. See Radial, Interradial, Marginal, and Ambital.
Inframarginal.
See Marginal plates.
Interbrachial.
The actinal and marginal areas between the rays . These spaces have plates derived
from various regions. In primitive forms, but a single plate is present, the
axillary interbrachial, derived by crowding orally the axillary inframarginal.
In other genera, inframarginals are crowded in pairs orally and form interbrachial
REVISION OF PALEOZOIC STELLEBOIDEA. 15
areas, the interbrachial marginals. Or the adambulacrals may enter into the
construction of these areas, forming interbrachial adambulacrals. When the adam-
bulacrals and inframarginals are separated by small plates, the latter are called
accessory interbrachials; they may extend almost to the distal ends of the rays.
Interradial.
A point half-way between the perradii. The interambulacral areas are inter-
radial in position. See Perradial.
Interradial plates.
These abactinal plates are rarely retained in living mature starfishes. In many
Paleozoic genera they are large, interradial in position, five in number, and
lie inside the basal or proximal supramarginals and between the primary radials.
They may be homologous with the basals of crinids. They are also known as
Madreporite ormadreporic plate.
A sieve-like or spongy plate, with many irregular openings for the circulation
of sea water into the so-called "stone canal, " or better, into the water- vascular
system. It is commonly granular or striated, and in Asteroidea is abactinal
on the disk, basally situated between two rays, but in the Auluroidea is prob-
ably always actinal in position. In recent multiradiate forms there are species
with more than one madreporite.
Marginal plates.
In general the plates which make up the marginal columns bounding the rays,
or disk, or the entire animal. They are parts of the primary skeleton. In the
early Paleozoic genera the inframarginal and supramarginal columns are usually
not directly superposed, making one column of superposed halves as in Meso-
zoic and Recent starfishes, but the former column lies outside or laterally of
the latter. The inframarginals are then the true marginals, and form a part
of both the abactinal and actinal sides. In the primitive arrangement the
inframarginals adjoin the adambulacrals, but in derived forms with large pen-
tagonal disks they are separated from the latter by accessory interbrachial plates.
The situation of the marginals is, however, not always at the margin of the
disk and rays, but they may retain their primitive position beside the adam-
bulacrals; this is more especially true in forms having a well-developed pen-
tagonal disk. These plates are also referred to as marginalia, supramarginalia,
and inframarginalia.
In some forms the supramarginals are not recognizable as such, while the infra-
marginals are well-developed; and in others none of the marginals are distinctly
discernible. Sometimes the plates of the inframarginal columns adjoin one
another, while those of the supramarginal series are separated by small acces-
sory supramarginal pieces. Then the inframarginals may be separated from
the supramarginals by ambital ossicles (see Ambital) . A proximal supramarginal
is the proximal plate of a column and may not be homologous with the primary
supramarginal in primitive forms. Proximal inframarginal is used in a similar
way and refers to the proximal axillary plate, two of which of adjoining col-
umns occupy an axil or lie on either side of the axillary marginal. The latter
in primitive forms alone occupies the angle between the rays. The marginals
in large disks without angles may be termed disk infra- and supramarginals.
The large disk of some species is formed by the oral crowding of pairs of
proximal inframarginals, the interbrachial marginals.
Measurements.
The size of starfishes is usually given in millimeters along the greater and lesser
radii, that is, from the center of the mouth or disk to the tips of the rays. The
symbol for this line is R. The smaller radius is from the center of the animal
to the center of the interradii; the symbol is r.
50601 Bull. 8815 2
16 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Mouth angle plate.
See Oral armature.
Ocular plate.
A single large, grooved eye-plate occupying the distal ends of rays and support-
ing a sensory tentacle at the base of which occurs the eye-spot. They appear in
the larval stage as the primary radials and with growth pass outward, remain-
ing at the tip of the rays as the oculars. They are also known as terminals.
These large plates are not present .as such in early Paleozoic genera, and are
unknown before the Carboniferous.
Oral.
See Actinal.
Oral angles.
The interradial actinal areas around the mouth.
Oral armature.
The pairs of plates, usually five in number, composing the apices around the
central actinal opening or mouth. In most Paleozoic forms the armature
consists of the proximal modified adambulacral plates. These pieces are also
known as Mouth angle plates.
In some Paleozoic forms (Hudsonaster) there lies in front of each pair of mouth
angle plates a single plate; this is known as the Torus.
Gregory (1900, p. 241) writes that "the Oral Skeleton (or actinostomial ring)
consists of a solid calcareous ring around the mouth. It is composed of thirty
plates in a quinqueradiate starfish, there being always six times as many plates
as there are rays. Each segment of the oral skeleton consists of two pairs of
ambulacral, and of one pair of adambulacral ossicles. In Asterias [a cryp-
tozonian] the ambulacral plates are more prominent than the adambulacrals,
and project into the oral cavity."
When the ambulacral elements are the more prominent, the oral skeleton is said
to be of the ambulacral type. This is only present in cryptozonians. When
the adambulacral ossicles are the most prominent, the oral skeleton is of the
adambulacral type. This latter construction is the more primitive and occurs
in all the Phanerozonia, but is also present in some of the cryptozonians.
Ossicles.
See Plates.
Papulae.
In living Asterias, a cryptozonian, from between the spicular plates there rise
from all parts of the external surface short and small integumentary protuber-
ances that are used for respiration. In the more heavily plated Phanerozo-
nia they are limited to the abactinal surface enclosed between the supramar-
ginal plates. It is probable that papulae were present in all Paleozoic forms
having rounded or spicular ossicles, issuing in the open spaces between the
plates. They probably were absent in the earliest closely plated starfishes,
such as Hudsonaster. See Csecal pores.
Paxillse.
Minute calcareous processes arranged around large spines. None are known
in Paleozoic genera.
"Another type of spines occurs as part of the structures known as 'paxillse.'
Each paxilla consists of a thick plate supporting a number of short, calcareous
pillars, the summit of each of which is covered by a group of small spines.
In some Phanerozonia * * * the paxillse occupy almost the whole abactinal
surface of the Asteroid" (Gregory 1900, p. 247).
Pedicellarids.
Pincer or scissors-like calcareous appendages, attached to the spines, the plates,
or the skin, which keep the body-wall clean of parasites. None have as yet
KEVISION OF PALEOZOIC STELLEROIDEA. 17
been detected with certainty in Paleozoic genera. In the primitive Recent
forms they are also absent, the integument being protected by an abundance
of closely set spines.
"The Pedicellarise of Asteroids are of four main types. The simplest form con-
sists of a row of pairs of small, sessile, opposable spines; these are the 'pseudo-
pedicellariae.* The members of the second set are ' sessile.' The next advance
is the development of a stalk; of these pedunculate pedicellariae there are two
varieties: (1) the 'forficiform,' in which the two hooks are attached to the near-
est end of the basal plate nearest to them; (2) the 'forcipiform,' in which the
two hooks cross one another and are attached to the end of the basal plate far-
thest from them " (Gregory 1900, p. 247; also see Verrill 1914, pp. 25-34).
Perradial.
The area along the center of the rays. The ambulacra are perradial in position.
See Interradial.
Plates.
Some writers restrict the term plates to the five primary radials, five interradials,
and the central disk piece, and refer to all the other calcareous parts as ossicles.
The word plates is here used in the widest sense, and as interchangeable with
ossicles. For the sake of brevity the plates will be often referred to as margi-
nalia, radialia, etc.
Podial openings.
The rounded openings between the ambulacral plates for the protrusion of the
locomotor organs, the podia or water- tube-feet. See also Ambulacral.
Primary radial.
See Radial.
Primary skeleton.
The skeleton of a starfish is made up of primary and secondary ossicles. The
primary skeleton consists of the longest inherited and the first appearing plates
in the young, as the ambulacral, adambulacral, marginal and radial columns,
plus the primary pieces of the disk. All the other ossicles inserted between
these are of the secondary skeleton.
Radial
Refers only to the prominent abactinal primary plates holding a radial position.
Where present, they form columns of continuous reticular or separated radialia,
situated along the radial center of the rays. Proximal radials are those on or
near the disk and should not be confounded with the primary radials, the
probable homologues of the radials in crinids. These plates are also known as
median dorsals, dorsals or carinals; also as radialia.
Rays.
The radial "arms " of starfishes. These are hollow and contain the digestive,
reproductive, and water-vascular systems and other organs. The rays merge
gradually and without sharp demarcation into the central disk.
Respiratory pores.
See Csecal pores.
Secondary skeleton.
See Primary skeleton and Accessory plates.
Spines.
See PaxillsR and Pedicellarids.
Supramarginal.
See Marginal plates.
Terminals.
See Ocular plate.
18 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Torus.
See Oral armature.
Trimum.
See Bivium.
Vertebral ossicles.
In the Ophiuroidea each, pair of adjoining ambulacralia is united into a single
plate, the vertebra of the arms. As the arms are very flexible, these ossicles
articulate upon one another as do the bones in the vertebral column of verte-
brates.
FINDING LIST OF PALEOZOIC STELLEROIDEA.
Page.
Acroura (Ophiura) schlotheimii Minister. Known to the writer only from the list
in Woodward, Geol. Mag., dec. 2, vol. 1, 1874, p. 7. Siluric.
Agalmaster Schondorf 131
A. grandis Schondorf 132
A. intermedius Schondorf 133
A. miellensis Schondorf 132
Aganaster Miller and Gurley 263
A. gregarius (Meek and Worthen) 264
A. (?) ep. Miller and Gurley =Lapworthura (f ) ep . 251
Alepidaster Meek 228
A. flexuosus (Miller and Dyer) 231
A. granuliferus (Meek) 230
A. gregarius Meek= Aganaster gregarius 264
A. miamiensis (Miller) 233
A. f, new species -.- 230
Anorthaster, new genus 125
A. miamiensis (Miller) 127
Archaeasterias 'M.\JlleY=Xenaster 129
? Argaster "0.2^1 Mesopalxaster 74
A. antiqua Gregory = Mesopalxaster (?) antiquus 86
Aspidosoma Goldi\i88=Encrinaster 241
A. arnoldi Goldf\i8s=Encrinaster arnoldi 243
A. eifelense SchondoTi=Encrinaster eifelensis 244
A. goldfussi Sch6ndori=Encrinaster goldfussi 243
A. grayss. Spencer=Encrinaster (?) grayse 245
A. petaloides SimonovitBch=Encrinaster petaloides 243
A. petaloides goslariensis Halfar=J!?icrmaser petaloides goslariensis 243
A.f pontis Cla,Tke=Encrinaster pontis 244
A. roemeri Schdndori=Encrinaster roemeri 244
A. schmidti Sch6ndoTi=Encrinaster schmidti 244
A. tischbeinianum Roemer=Jncrwaster tischbeinianus 244
Asterias Graham, Anthony, and 3ameB=Petraster (?) americanus 146
A. acuminatus Simonovitsch= Mesopalseaster (?) acuminatus 93
A. anthonii Dana=Pefr aster (?) americanus 146
A. antiqua ILi8mger=Lindstrdmaster antiquus 149
A. antiqua Troost= Mesopalseaster (?) antiquus 86
A. antiquata Ijocke=Promopalseaster spedosus 109
A. asperula Roemer= Urasterella asperula 188
A. asperula Roemer (pait)=Eoluidia decheni 262
A. constellata Thorent= Urasterella (?) constellata .' 187
A. matutina 'BsX\.=Hudsonaster matutinus '. 57
A. montanus Stschurowsky= Urasterella montana 189
A. primseva Salter and Soweiby=Stenaster (?) obtusus 167
REVISION OF PALEOZOIC STELLEROIDEA. 19
Page.
Asterias primordialis Anon.=Promopalseaster, sp. undet 108
A. (Archseasterias) rhenana M.ulleT=Xenaster (?) rhenanus 131
A. spinosissima Roemer. An unsolved starfish from the Lower Devonic of
Bundenbach, Germany. See Roemer, Palaeontographica, vol. 9, 1863, p. 147,
pi. 29, fig. 4; and Schondorf, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 62, 1909, p. 33.
A. sp. undet. Hall 1847Ccelaster tenuiradiatus .
Asteriscus antiquus Pictet=Lindstromaster antiquus 149
Astropecten Link . . 161
A. (?) schluteri Stiirtz 161
Ataxaster Jaekel 161
A. pygmxus Jaekel 161
Australaster, new genus 72
A. giganteus (Etheridge) - - 73
A. (?) stutchburii (Etheridge) 73
Bdellacoma Salter 254
B. vermiformis Salter 254
Bohemura Jaekel 223
B. jahni Jaekel 223
Bundenbachia Stiirtz 234
B. beneckei Stiirtz 234
B. grandis St\\rtz=Palaophiomyxa grandis 235
Calliaster Trautschold=CaZtoerea 190
C. mirus TT&\itschold=Calliasterella mira 190
Calliasterella, new name 190
C. mira (Trautschold) 190
Cheiropteraster Stiirtz 202
C. giganteus Stiirtz 202
Cholaster Worthen and Miller 265
C. peculiaris Worthen and Miller 266
Ccelaster Sandberger=$/>ara'aster 95
C. americanus D'Orbigny=Peraser (?) americanus 146
C. latiscutatus Sandberger=parmister latiscutatus 96
C. tenuiradiatus D'Orbigny, Prodr. de Paleont., vol. 1, 1849, p. 22. Based on
an isolated madreporite described as Asterias, sp. undet., Hall, Pal. New
York, vol. 1, 1847, p. 18, pi. 4, figs, lla, 116.
Compsaster Worthen and Miller 192
C. formosus Worthen and Miller 193
C., new species 194
Cribellites carbonarius Tate 274
Devonaster, new genus 97
D. chemungensis, new species 101
D. eucharis (Hall) 98
E chinas terella Stiirtz 200
E. ? darwini Clarke 200
E. sladeni Stiirtz 200
Echinasterias Stiirtz 211
E. spinosus Stiirtz 211
EchJLnodiscaster Delage and Herouard 211
E. multidactylus (Stiirtz) 211
Echinodiscites SchucheTt=Echinodiscaster 211
Echinodiscus Stwrtz=Echinodiscaster 211
E. multidactylus St\ulz=Echinodiscaster multidactylus 211
20 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Page.
Echinostella Stiirtz 212
E. traquairi Stiirtz 212
Eifelaster Schondorf 133
E. follmanni Schondorf 134
Encrinaster Haeckel 241
Encrinaster Haeckel (p&rt)=Protaster. . . : 224
E. arnoldi (Goldfuss) 243
E. eifelensis (Schondorf) 244
E. goldfussi (SchSndorf). . r 243
E. grayse, (Spencer) 245
E. petaloides (Simonovitsch) 243
E. petaloides goslariensis (Halfar) 243
E. pontis (Clarke) 244
E. roemeri (Schondorf) 244
E. schmidti (Schondorf) 244
E. tischbeinianus (Roemer) 244
Eoactis Spencer. See Urasterella 173
E. simplex Spencer. See Urasterella girvanensis 186
Eoluidia Stiirtz 262
E. decheni Stwtz 262
Eophiura Jaekel 222
E. bohemica, new species 222
Eophiurites Stiirtz =Eoluidia 262
E. decheni (Stiirtz) =Eoluidia decheni 262
Eospondylus Gregory 263
E. primigenius (Stiirtz) 263
Etheridgaster Gregory= Monaster 170
E. clarJcei Gregory = Monaster clarJcei 172
Eucladia Woodward 276
E. (?) beecheri, new species 278
E. johnsoni Woodward 276
E. woodwardi Sollas 277
Eugaster ILal\=EiLgasterella 237
E. continnus Ringueberg=Eugasterella ( ?) concinna 239
E. logani H&\\=Eugasterella logani 238
Eugasterella, new name 237
E. ( ?) concinna (Ringueberg) 239
E. logani (Hall) 238
Euryak annulatum DeKay. Apparently a lysophiurid. See Hall, Pal. N. Y.,
vol. 3, 1861, p. 134 (nomen nudum).
Euthemon Sollas , 279
E. igerna Sollas 279
Furcaster Stiirtz 261
F. (?) daoulasensis (Davy) 261
F. palseozoicus Stiirtz 261
Gregoriura Chapman 233
G. spryi Chapman. . .- 234
HaUaster Stiirtz 254
#./or&m(Hall) 255
Helianthaster Clarke (part)==PaZa20$oZoster 209
Helianthaster Roemer 159
H. filiciformis Woodward 160
H. gyalum Clarke =PaZasoso Zoster ( ?) gyalum 210
REVISION OF PALEOZOIC STELLEKOIDEA. 21
Page.
Helianthaster gyalum Clarke (paTtyLepidasterella babcocki 160
H. , new species Clarke Lepidasterella babcocki 160
H. rhenanus Roemer 159
H. roemeri Clarke PaZasoso Zoster roemeri 211
Hisingeraster Stiirtz =Lindstromaster 148
H. antiqua Sturtz=Lindstrdmaster antiquus 149
Hudsonaster Stiirtz 53
H. batheri, new species 65
H. incomptus (Meek) 61
H. matutinus (Hall) 57
E. milleri, new species 60
H. narrawayi (Hudson) 59
H. rugosus (Billings) 64
Jaekelaster Stiirtz 192
/. petaliformis Stiirtz 192
Lapworthura Gregory 250
L. cylindrica Parks= Tseniaster cylindricus 220
L. miltoni (Salter) 251
L. sollasi Spencer 251
L. (?)sp 251
Lepidaster Forbes 158
L. grayi Forbes 158
Lepidasterella, new genus 160
L. babcocki, new species 160
Lindstromaster Gregory 148
L. antiquus (Hisinger) 149
Loriolaster Stiirtz 201
L. mirabilis Stiirtz 201
Medusaster Stiirtz 212
M. rhenanus Stiirtz 212
Mesopaleeaster, new genus 74
M. ( ?} acuminatus (Simonovitsch) 93
M. (?) antiquus (Troost) 86
M. bellulus (Billings) 91
M. caractaci (Gregory) 92
M. ( ?) cataractensis, new species 89
M. (?) clarki (Clarke and Swartz) 94
M . ( ?) dubius (Miller and Dyer) 85
M.finei (Ulrich) 81
M. granti (Spencer) 89
M. intermedius, new species 79
M. ( ?) lanceolatus, new species 82
M. (?) parviusculus (Billings) 87
M. proavitus, new species 83
M. sha/eri (Hall) ; 77
M. ( ?) wilberanus (Meek and Worthen) 84
Miomaster Schondorf 96
M. drevermanni Schondorf 96
Miospondylus Gregory 263
M. rhenanus (Stiirtz) 263
Monaster Etheridge 170
Monaster Gregory= Australaster 72
M. darkei (De Koninck) 172
22 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Page.
Monaster giganteus 'Ethendge=Australaster giganteus 73
M. stutchburii JZtheridge=Australaster stutchburii 73
Neopalseaster, new genus 134
N. crawJordsmlUnsis (Miller) 136
Onychaster Meek and Worthen 268
0. asper Miller 272
0. barrisi (Hall) 272
0. confragosus Miller 273
0. demissus Miller 274
0. flexilis Meek and Worthen 270
Ophiopege ~Bohm=Aganaster 263
Ophiura obtusa Eichwald= Pro taster (Ophiura) obtusus.
0. (?) ramosa Fahrenkohle. Unknown to writer. See Bull. Soc. Imp. Nat.
Moscow, n. ser., vol. 17, 1844, p. 208, pi. 3, figs. 1-3; and Woodward, Geol.
Mag., dec. 2, vol. 1, 1874, p. 7 (listed only). Coal Measures.
0. rhenana Sturtz= Miospondylus rJienanus 263
0. salteri Salter and Sowerby =Protaster ( ? ) salteri 227
0. schlotheimii Munster= Acroura (Ophiura) schloiheimii.
Ophinrella Stwrtz=Eospondylus 263
0. primigenia St\irtz=Eospondylus primigenius 263
Ophiurina Stiirtz 246
0. lymani Stiirtz 247
Palseaster Hall 67
Paleeaster of authors =Hudsonaster } Mesopalseaster, Devonaster, Promopal-
seaster, Anorihaster, Neopalseaster, Petraster, Palasterina, Monaster, Uras-
terella.
Palseaster (Monaster) Etheridge= Monaster 170
Paleeaster (Monaster) Etheridge (part) =Australaster 72
P. (Argaster) antiqua Hall= Mesopalseaster ( ?) antiquus 86
P. antiquatus H&ll=Promopaldeaster speciosus 109
P. antiquus Miller= Mesopalseaster ( f) antiquus 86
P. asperrima Salter= Urasterella (?) asperrima 187
P. caractaci Salter= Mesopalseaster caractaci 92
P. clarkana Miller =Hudsonaster incomptus 61
P. clarkei De Koninck= Monaster clarkei 172
P. clarkei Mi\ler=Hudsonaster i/icomptus 61
P. clarki Clarke and Swartz= Mesopalseaster ( f) clarki 94
P. coronella Salter=Stenaster (?) coronella 167
P. crawfordsvillensis Miller = Neopalseaster crawfordsmllensis 136
P. dubius Miller and Dyer= Mesopalseaster (?) dubius 85
P. dyeri 1&eek=Promopal3easter dyeri 120
P. eucharis Il f d,ll=Devonaster eucharis 98
P. exculptus M.i\\eT=Promopalseaster exculptus 117
P. find Ulrich= Mesopalseaster finei 81
P. ( Monaster) giganteus Etheridge =^Lttsratoer giganteus 73
P. granti Spencer= Mesopalseaster granti 89
P. granulosus ILall=Promopalseaster granulosus 112
P. granulosus MeekPromopalseaster speciosus 109
P. harrisi Miller= Urasterella grandis 180
P. hirudo Salter= Urasterella hirudo 188
P. imbricatus Salter= Tetraster (?) imbricatus 169
P. incomptus Meek.=Hudsonaster incomptus 61
P. jamesi Hall Petraster ( ?) americanus 146
REVISION OF PALEOZOIC STELLEROIDEA. 23
Page.
Paldeaster longibrachiatus Miller =Promopalxaster spinulosus 115
P. magnificus Miller =Promopal9easter magnificus 122
P. matutinus Hall=Rudsonaster matutinus 57
P. miamiensis Miller =Anorthaster miamiensis 127
P. montanus Trautschold= Urasterella montana 189
P. niagarensis Hall 69
P. obtusus Salter=Stenaster (?) obtusus 167
P. parviusculus Billings= Mesopalseaster ( f ) parviusculus 87
P. pulchellus Billings= Urasterella pulchella 178
P. pygmsea Eichwald. Not known to writer. Middle Ordovicic, Pulkowa,
Russia.
P. ruthveni Salter= Urasterella ruthveni 187
P. shajferi Hall= Mesopalseaster shajferi 77
P. simplex Miller and Dyer=Hudsonaster incomptus 61
P. spedosus Meek=Promopalseaster spedosus 109
P. spinulosus Miller and ~Dyer=Promopalxaster spinulosus 115
P. squamatus Salter. Not known to writer. See Cat. Woodwardian Mus., 1873,
p. 47; and Woodward, Geol. Mag., dec. 2, vol. 1, 1874, p. 7 (listed only).
Bala of England.
P. ( Monaster] stutchburii Etheridge=Australaster ( ?) stutchburii 73
P. wilberanus Hall= Mesopalseaster ( ?) wilberanus 84
P. ? wilsoni Raymond Promopalseaster wilsoni 106
P. wyJcoffi Miller and Gurley=Promopalseaster wykoffi 119
Palaeasterina Gregory =Palasterina 150
Paleeasterina of authors =Petraster 138
P. antiqua S<er=Lindstr6master antiquus 149
P. approximata Miller and Dyer=Petr aster spedosus 142
P. fimbriata Meek and WorthQn=Schcenaster fimbriatus 204
P. rugosa Billings Hudsonaster rugosus 64
P. spedosa Miller and Dyer=Petr aster spedosus 142
Palaenectria Stmtz=Palseostella 155
P. devonica StiiTtz=Palssostella solida 156
Palaeocoma Miller (p^Tf)=Ptilonaster 239
Palseocoma S<er=Sturtzaster 252
Palaeocoma (Bdellacoma) Salter =Bdellacoma 254
Palaeocoma (Bhopalocoma) SsilteT=Rhopalocoma 254
P. colvini SalteTStiirtzaster colvini 253
P. cygnipes Salter Sturtzaster cygnipes 253
P. cylindrica Billings= Txniaster cylindricus 220
P. marstoni Salter= Sturtzaster marstoni 253
P. princeps MilleT=Ptilonaster princeps 240
P. pyrotechnica S<er=Rhopalocoma pyrote'chnica 254
P. spinosa Billings= Tssniaster spinosus 219
P. vermiformis Salter = Bdellacoma vermi/ormis 254
Palseodiscus ferox Salter=an echinid. See Jackson, Mem. Boston Soc. Nat.
Hist., vol. 7, 1912, p. 250.
Palseophioinyxa Sturtz 234
P.grandis (Sturtz) 235
Palseophiura Sturtz 235
P. simplex Sturtz 235
Palaeosolaster Sturtz 209
P. gregoryi Sturtz 209
P. (?) gyalum (Clarke) 210
24 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Palxosolaster roemeri (Clarke) 211
Paleeospondylus Stuitz=Palastropecten 261
P. zitteli Sturtz=Palastropecten zitteli ". 262
Paleeostella Stiirtz 155
P. solida Stiirtz 156
Paleespondylus Stiirtz =Palastropecten 261
Palaeura Jaekel 223
P. neglecta, new species 223
Palasteracanthion Stiirtz= Urasterella 173
Palasterina Billings (p3tft)=Schuchertia 195
Palasterina McCoy 150
P. antiqua SsdteT=Lindstr omaster antiquus 149
P. bonneyi Gregory 153
P. follmanni Sturtz=Pseudopalasterina follmanni 157
P. (?) jamesi D&na l =Petraster (?) americanus 146
P. kinahani Baily= Uranaster kindhani 155
P. primseva (Forbes) 153
P. (?) ramseyensis Hicks 154
P. rigidus Billings =Petraster rigidus 141
P. rugosa Billings =Hudsonaster rugosus 64
P. stellata BillingB=Schuchertia stellata 196
Palasteriscus Stiirtz 200
P. devonicus Stiirtz 200
Palastropecten Stiirtz 261
P. zitteli Stiirtz 262
Palmipes antiqua Forbes= Lindstromaster antiquus 149
Petraster Billings 138
P. (?) americanus (D'Orbigny) 146
P. (?) antiqua Shumard= Mesopalxaster (?) antiquus 86
P. bellulus Billings = Mesopalseaster bellulus 91
P. rigidus (Billings) . % 141
P. rigidus (Billings) (p&Tt)=Hudsonaster matutinus 57
P. smyihi McCoy 147
P. spedosus (Miller and Dyer) 142
P. wilberianus Meek and Worthen= Mesopalseaster ( ?) wilberanus 84
Promopalaeaster, new genus 102
P. bellulus, new species 113
P. ^m'(Meek) 120
P. exculptus (Miller) 117
P. granulosus (Hall) 112
P. magnificus (Miller) 122
P. prenuntius, new species 107
P. spedosus (Meek) 109
P., sp. undet 108
P. spinulosus (Miller and Dyer) 115
P. wilsoni (Raymond) [ 106
P. wykoffi (Miller and Gurley) 119
Protaster Forbes 224
Protaster Kall=Hallaster 254
P. f barrisi TL2il\=Onychaster barrisi 272
P. biforis Gregory 226
P. brisingoides Gregory=Sturtzura brisingoides 236
P. daoulasensis Davy = Fur caster (?) daoulasensis 261
REVISION OF PALEOZOIC STELLEROIDEA. 25
Protaster decheni Dewalque= Tremataster (?) decheni 248
P. elegans Parks= Txniaster elegans 221
P. flexuosus Miller and DyeT=Alepidaster flexuosus 231
P. forbesi Hall= Hallaster forbesi 255
P. (?) granuliferus lAeek=Alepidaster granuliferus 230
P. (?) gregarius Meek and Worth en =Aganaster gregarius 264
P. groomi Sollas and Sollas 227
P. leptosoma Salter =Sturtzur a leptosoma 237
P. miamiensis Miller =Alepidaster miamiensis 233
P. miltoni SalterLapworthura miltoni 251
P. (Ophiura)obtususEichwald. Unknown to the writer. See Schichtensystem
Esthland, p. 193; Lethsea Rossica, p. 661; and Woodward, Geol. Mag., dec.
2, vol. 1, 1874, p. 7 (listed only). Siluric of Russia.
P. petri Salter. Unknown to the writer. See Salter, Cat. Mus. Pract. Geol., p. 30;
and Woodward, Geol. Mag., dec. 2, vol. 1, 1874, p. 7 (listed only). Probably
Bala, Church Stretton, England.
P. ( f ) salteri (Salter) 227
P. sedgwickii Forbes 225
P. (?) stellifer Ringueberg 228
P. (?) whiteavesianus Parks 227
Protasteracanthion Stiirtz= Urasterella 173
P. primus Stiirtz= Urasterella asperula 188
ProtasterinaUlrich==.47epw/aser 228
P . fimbriata Ulrich=Alepidaster fiexuosus 231
P. flexuosa 3aio.eB=Alepidaster fiexuosus 231
Protoeuryale C. F. Roemer=a crinid.
Protopalseaster Hudson= Hudsonaster 53
P. caractaci Spencer= Mesopalseaster caractaci 92
P. nanawayi Hudson = Hudsonaster narrawayi 59
Pseudopalasterina Sturtz 156
P.follmanni (Sturtz) 157
Ptilonaster Hall 239
P. princeps Hall 240
Bhenaster Schondorf 133
R. schwerdi Schondorf .,*. 133
Rhodostoma Sollas and Sollas =Sturtzura .7. 236
R. leptosoma Sollas and Sol\&s=Sturtzura leptosoma 237
Rhopalocoma Salter 254
R. pyrotechnica Salter 254
Roemeraster Stiirtz= Urasterella 173
R. ( ?) acuminatus Sturtz = Mesopalseaster ( ?) acuminatus 93
R. asperula Stiirtz= Urasterella asperula 188
Salteraster Stiirtz= Urasterella * 173
S. asperrimus Stiirtz= Urasterella (?) asperrima 187
Schoenaster Meek and Worthen 202
S. fimbriatus Meek and Worthen 204
S. (?) legrandensis Miller and Gurley 206
S. (?) montanus Raymond 207
S., new species 206
S. (?) wachsmuthi Meek and Worthen 205
Schuchertia Gregory 195
S. laxata, new species 198
S. ordinaria, new species 199
26 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Page.
Schuchertia stellata (Billings) 196
Siluraster Jaekel 65
S. perfectus Jaekel 66
Spaniaster Schondorf 95
S. latiscutatus (Sandberger) 96
Squamaster Ringueberg 249
S. echinatus Ringueberg 249
Stenaster Billings 163
Stenaster Billings (part)= Urasterella 173
S. (?) confiuens Trautschold 167
S. (?) coronella (Salter) 167
S. grandis Meek= Urasterella grandis 180
S. huxleyi Billings= Urasterella huxleyi 182
S. (?) obtusus (Forbes) 167
8. pulchellus Billings= Urasterella pulchella 178
S. salteri Billings 165
Stiirtzaster Etheridge 252
S. colvini (Salter) 253
8. cygnipes (Salter) 253
8. marstoni (Salter) 253
S. (?) mitchelli Etheridge 254
Stiirtzura Gregory 236
S. brisingoides (Gregory) 236
S. leptosoma (Salter) 237
S. leptosomoides Chapman 237
Sympterura Bather 256
S. minveri Bather 256
Tseniaster Billings 216
T. australis McCoy =Sturtzura brisingoides 236
T. cylindricus (Billings) 220
T. elegans Miller 221
T. meafordensis Foerste 221
T. schoharise. Ruedemann 220
T. spinosus (Billings). 219
Taeniura Gregory= T&niaster 216
T. cylindrica Gregory = Tseniaster cylindricus 220
Tetraster Nicholson and Etheridge 167
T. asperrimus Nicholson and Etheridge = Urasterella (?) asperrima 187
T. (?) imbricatus (Salter) 169
T. sp. indet. Nicholson and Etheridge= Urasterella girvanensis 186
T. wyville-thomsoni Nicholson and Etheridge 168
T. wymlle-thomsoni Nicholson and Etheridge (part) = Hudsonaster batheri 65
Tremataster Worthen and Miller 247
T. (?) decheni (Dewalque) 248
T. diffidlis Worthen and Miller 247
Trentonaster Sturiz=Schuchertia 195
T. stellata Sturtz== Schuchertia stellata 196
Trichotaster plumiformis Wright. A 10-rayed asterid from the Wenlock lime-
stone at Dudley, England, in the collection of Dr. Grindrod. Insufficiently
described to be regarded as defined. See Quart. Journ. Geol. Soc., vol. 29,
1873, p. 421; also Woodward, Geol. Mag., dec. 2, vol. 1, 1874, p. 7 (listed as
Trochitaster plumiformis).
Trimerast er Schondorf ... 134
BEVISION OF PALEOZOIC STELLEKOIDEA.
27
Page.
Trimeraster parvulus Schondorf 134
Trochitaster plumiformis WTight=Trichotaster plumiformis.
Uranaster Gregory 154
U. elizse, Spencer 155
U. kinahani (Baily) 155
Uraster hirudo Forbes= Urasterella hirudo 188
U. obtusus ~FoTbes=Stenaster (?) obtusus 167
U. primsevus Forbes=PaZasmna primseva 153
U. ruthveni Forbes= Urasterella ruthveni 187
Urasterella McCoy 173
Urasterella Stiirtz I8$3=Stenaster 163
U. (?) asperrima (Salter) 187
U. asperula (Roemer) 188
U. (?) constellata (Thorent) 187
U. girvanensis, new species 186
U. grandis (Meek) 180
U. hirudo (Forbes) 188
U. huxleyi (Billings) 182
U. montana (Stschurowsky) 189
U., new species 189
U. pulchella (Billings) 178
U. ruthveni (Forbes) 187
U. selwyni McCoy 188
U. ulrichi, new species 183
Xenaster Simonovitsch 129
X. dispar Schondorf 131
X. elegans Schondorf 131
X. eucharis Schondori=Devonaster eucharis 98
X. margaritatus Follmann=^". elegans 131
X. margaritatus Simonovitsch 131
X. margaritatus Simonovitsch (part)=J5T. dispar and Agalmaster grandis 131, 132
X. (?) rhenanus (Miiller) 131
X. simplex Simonovitsch =Spaniaster latiscutatus 96
REMARKS ON THE DEVELOPMENT OF PALEOZOIC STELLEROIDEA.
GEOLOGICAL.
Distribution. This memoir treats of the following Paleozoic
Stelleroidea :
Species.
North
Ameri-
can
species.
Euro-
pean
species.
Southern
Hemis-
phere
species.
Asteroidea...
Auluroidea. .
Ophiuroidea.
Total. . .
110
59
5
174
11
28 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Arranged geologically the distribution is as follows :
Ordo-
vicic.
Siluric.
Devonic.
Lower
Carbon-
iferous.
Upper
Carbon-
iferous.
Permic.
Asteroidea
46
16
36
6
6
Auluroidea
15
17
24
3
OphiuToidea
5
o
o
Total
61
33
60
14
6
Of all of the Paleozoic Stelleroidea (174 species), not a single one
is common to North America and Europe. Of genera having specific
representation in more than one continent there are the folio whig:
Hudsonaster, 5 in America, 1 in Scotland; Mesopalseaster, 13 (7 doubt-
ful) in America, 1 in England, ? 1 in Germany; Petraster, 3 in America
(1 doubtful), 1 in Australia; Stenaster, 1 in America, ? 1 in Ireland and
Wales, ? 1 in England, ? 1 in Kussia; Urasterella, 5 in America, 1 in
Scotland, ? 1 in Wales, ? 1 in France, 2 in England, 1 in Australia,
1 in Germany, 1 in Russia; Paldeosolaster, 1 in Germany, ? 1 in
America; Protaster, ? 2 in America, 1 in England, 2 (1 doubtful) in
Wales; Tremataster, 1 in America, ? 1 in Belgium. Of Ophiuroidea,
none have wide distribution.
Nearly all of the species have very short geologic ranges, but very
few occurring in more than a single formation. Mesopalxaster
intermedius ranges from the ? Utica to the Maysvillian, M. sliafferi
from the Maysvillian to the Richmondian, Alepidaster flexuosus from
the basal Edenian to the upper Maysvillian, Hudsonaster incomp-
tus from the Edenian into the Richmondian, Promopalxaster mag-
nificus and Petraster spedosus from the Maysvillian to the Rich-
mondian.
These tables also show that the Ophiuroidea do not begin earlier
than the Lower Carboniferous (Burlington) and that the Auluroidea
cease to exist after the same period, for none are as yet known in the
Upper Carboniferous or Permic. The latter attained their maximum
of development in the Siluric and Devonic, with very rapid decline
in the Lower Carboniferous.
The great development of Asteroidea in the Ordovicic is due to the
marked specific evolution of the genera Mesopalseaster and Promo-
palseaster in North America. This knowledge is the result of care-
ful work by local collectors in areas abounding in good fossils. On
the other hand the marked rise hi specific representation in the De-
vonic is due to the protracted collecting in the earlier beds of this
period, the roofing slates of Bundenbach, Germany.
Preservation and occurrence. Starfishes are most often preserved
in sandstones and muds tones and least often in limestones. The
REVISION OF PALEOZOIC STELLEROIDEA. 29
reason why they occur less frequently in limestones is partially
accounted for by the fact that during the weathering process they
are almost always ruined. Those found in such deposits nearly
always occur in the thin shale partings between the bedded limestones.
In coarse sandstones they are less often seen, probably because these
sediments are so much moved by the storm waves action that
destroys the skeleton by separating the plates. Unless a starfish
is quickly covered by sediment it is sure to be broken up and jumbled
into a mass of unrelated ossicles, because the skeletal parts are rarely
coossified. Hence the best preserved specimens are found in mud-
stones and especially in the fine-grained, somewhat muddy sand-
stones. Here they usually occur as fine molds, since all of the calcium
carbonate has been dissolved out by the atmospheric waters. Such
material is apt to be fairly abundant in individuals, and although a
little difficult to study is often well preserved. Its interpretation is
dependent on artificial casts or squeezes in wax or gutta-percha.
When found in black slates, as at Bundenbach, the skeleton is pre-
served in iron pyrite, and even though these sediments have been
subjected to mountain making, the specimens can be cleaned me-
chanically of the adhering slate. The process is, however, a laborious
one and has been successfully used only by Stiirtz and his two prep-
arators. In the calcareous shales asterids are often well preserved,
with the original skeleton more or less permineralized and the ossicles
cemented together so that parts of the individuals weather out as
free fossils. This is particularly the case in the Bichmondian deposits
of Ohio and Indiana.
As a rule, starfishes are obtained in single specimens and as acci-
dental finds, and for this reason they are among the rarest of known
Paleozoic animals; they are the " fancy fossils" of the local collectors
and the " choice specimens" of the museums. All of this is, however,
due to the accident of preservation plus their great destruction
through weathering. That starfishes and ophiurids were common,
though probably not so abundant as aulurids, is proven when they are
located in their entombing sediment and then quarried for. This
is best seen in the well-known Lower Devonic slates of Bundenbach,
where the quarrying for roofing material has produced as many
starfishes as all other localities put together. A great variety
has also been secured here, so that it is the only locality and time
of which we can say that we know the starfish fauna. The most
remarkable starfish find, however, is that made near Saugerties,
New York, where over 400 examples of the Middle Devonic
Devonaster eucharis were found in a fine-grained, somewhat muddy
sandstone, extending over an area of 200 square feet. They occur
as natural molds, and as the animals are found closely associated
with Grammy sia, it is thought that while feeding on these bivalves
30 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
they were suddenly entrapped by the engulfing sand. In the
crinid bed of the Keokuk formation at Crawfordsville, Indiana, the
ophiurid Ony chaster flexilis is also often met with.
Aulurids, although as a rule not well preserved, are probably more
often seen than starfishes, and certainly are far more abundant than
ophiurids. The latter condition is, however, probably explained by
the fact that no ophiurid is known older than the Carboniferous.
Of the aulurid Tseniaster elegans 31 individuals occur on one small
slab. It is probable that if aulurids and asterids were systematically
dug for, a far greater harvest would result than that now at hand.
Whenever such specimens are found in place, such prospects should
be followed up by digging or by prolonged search on the part of the
local workers.
It is very seldom that a Paleozoic asterid is so well preserved that
all of its parts can be made out, and this is especially true as regards
the dorsal side. During the decay of the animals it is probably
true that in most cases the individual lies with its ventral side down,
and as the soft parts vanish the dorsal skeleton sinks in irregularly
over the actinal skeleton. It is the central region of the disk that
suffers dismemberment most. Then, as the two sides of the skeleton
finally come to be fully compressed, the rays take on an unnaturally
great width, so that in many of the fossils preserved in shales the
ambulacral furrow is considerably wider than in nature. For
these reasons it is often impossible to say whether the ambulacralia
are of the opposite or the alternate arrangement, and what was the
original structure of the disk and the interambulacral areas.
ASTERID EVOLUTION.
GENERAL OBSERVATIONS.
The majority of the starfishes studied by the writer are from
the Ordovicic strata of North America. Collectively these show
considerable evolution, for at this early time in the history of the
earth Phanerozonia, Cryptozonia, and primitive ophiurids ( = Aulu-
roidea) are present. Even near the base of the Middle Ordovicic
the two former groups appear to be equally common, while the
aulurids are as a rule rare and small, though there are occasional
large ones. Here the primitive phanerozonian asterids are also small,
hardly ever exceeding 15 mm. in diameter, while the derived and
far more complex Cryptozonia are large, some of them having rays
exceeding 50 mm. In the Lower Ordovicic of America no starfishes
are known, while the few that have been reported from Great Britain
appear to the writer to be from the Middle Ordovicic.
These facts show that in spite of their absence in Lower Ordovicio
rooks tjfcere must have lived at that time various kinds of starfishes^
REVISION OF PALEOZOIC STELLEEOIDEA. 31
Not only this, but we may confidently add that the varied organi-
zation of those of the Middle Ordovicic proves that there is a long
previous history of which paleontology knows nothing. The origin
of the Asteroidea goes back in all probability to the Proterozoic, as
may be inferred from the complex metamorphosis of the starfish
larva. In all of the Cambric, however, not a single starfish has been
found, and this is the more surprising when one considers the abun-
dance of life unearthed from these sediments in so many places
throughout the world. This may mean that a preservable starfish
skeleton was not evolved until Lower Ordovicic time, where they
will surely be found.
TJie most primitive fossil starfish. Of the Ordovicic starfishes, the
genus Hudsonaster is the most primitive, but with the full quota of
parts in the primary skeleton that an unmodified or unspecialized
starfish must have. Among living starfishes this primitive skeleton
is rarely seen, but occurs in Heterasterias volsellata, "and in a few
other species, especially when young" (Verrill 1914:27). There are
other genera with a simpler skeleton (Stenaster and Tetraster), but these
are clearly cryptozonian forms that have originated in a phanerozonian
stock not unlike Hudsonaster. The oldest Hudsonasters (see plates 1 to
6) are small animals about 10 mm. in diameter, with thick, highly con-
vex, closely adjoining plates, all of which are devoid of spines ex-
cepting the adambulacrals, which have simple, minute, articulating
spines. On the ventral side there are in each radius double columns
of rectangular ambulacralia, with the pieces of each ambulacrum
arranged opposite to one another, or nearly so. Outside of these are
single columns of adambulacralia of about the same number as the
ambulacrals; they are, however, somewhat larger and more trans-
verse. These are laterally bounded by much larger, highly convex,
granulated, very prominent marginals, which, as they border the
animals and are simple ossicles (there are not here two superposed
marginalia as is so common in living forms), are the inframarginal
columns. In the axils of the rays lies a single large marginal plate,
the axillary, and these occupy the entire interbrachial areas. Around
the inner sides of these axillaries the adambulacralia continue, the
two basal pieces being the largest and the essential elements of the
oral armature. All of the ventral ossicles are, therefore, seen to be
of the primary skeleton.
On the dorsal side of Hudsonaster, the rays have medially promi-
nent columns of radials, while on each side of these are other columns
of thick ossicles, alternating with the radialia, and these are the
supramarginals. The former ossicles continue closely adjoining to
near the center of the disk, where lies a single large plate usually called
in this memoir the central disk plate, or, more rarely, the centro-dorsal.
In the same way, the supramarginals abut in the axils upon a single
50601 Bull. 8815 3
32 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
large ossicle, the basal supramarginal plate. It is thought that the
ancestors of Hudsonaster had a dorsal disk skeleton made up of a
centro-dorsal, a first ring of 5 primordial radials, and a second ring
of 10 plates, 5 of which are" the second radials, and 5 interradial
pieces, the primordial supramarginals. In Hudsonaster, however,
some progression has taken place in the disk in the way of increase
of size. Therefore additional ossicles of the secondary skeleton have
developed to take up this space, these accessory disk pieces being
inserted between the centro-dorsal and the first ring of primordial
radials. Such* a development of a small number of accessory pieces
here is a hint of future complexity and the rise of the secondary skel-
eton, and we shall see how in later and derived genera their number
becomes multitudinous and their places of insertion nearly everywhere
in the older portions of the skeleton between the columns of ossicles.
The supramarginals do not margin the animals, but lie inside the
inframarginals and axillaries which do margin Hudsonaster and most
of the Paleozoic asterids.
From Hudsonaster to all other progressive asterids of the Phane-
rozonia type, the change lies mainly in the increasing number of the
ossicles, relative decrease in the size of the plates, introduction of
many new series of Accessory pieces, absorption and removal of
others, with a marked general tendency to break up the stiff and
ponderous inherited skeleton into one of small pieces, thus affording
greater flexibility and greater podial strength through the endless
duplication of ambulacral parts. These developmental tendencies
take place more especially on the dorsal area and are further accent-
uated through increase of body cavity, which demands an enlarged
skeletal covering. The disk widens, and along with it the proximal
parts of the rays, so that interbrachial areas are also affected, until
finally in more than one phylum pentagonal asterids result. The
interbrachial areas are dorsaUy increased by the insertion of acces-
sory pieces between the infra- and supramarginalia, and ventrally
by the crowding into these areas of, first, the single axillaries, and
then more and more of the oldest inframarginals in pairs, assisted
also here by the development of accessory ossicles.
Origin of the wriggling type of starfish. From Hudsonaster to the
other progressive asterids of the Cryptozonia type the evolutionary
tendencies are in the same directions, but here even greater flexibility
appears to be the main stimulus. Accordingly, the entire dorsal skel-
eton tends to break up into small loosely adjoining pieces and finally
even into a spicular spinose mesh. In these forms the ossicles of the
primitive columns are no longer discernible as such, and this tendency
is very apt to be likewise true of the primordial disk plates. On the
ventral side the inframarginals are no longer wholly present as such,
but may be here mixed up with the dorsal skeletal complex. How-
EEVISION OF PALEOZOIC STELLEKOIDEA. 33
ever, some of these plates are at times retained in the interbrachial
areas as large pieces, and in many forms they are still to be made
out clearly on the sides of the growing distal ends of the rays. Here
the inframarginals occur as columns of tiny closely adjoining ossicles
situated directly beside the adambulacrals, and in a few cases the
whole five primary dorsal columns (one radial, two supramarginal, and
two inframarginal) can be made out without any accessory pieces be-
tween them. In other words, a Paleozoic cryptozonian may retain
the marginals throughout life, but because of their small size and
isolation one from another by accessory pieces or because of intense
spiculization, they are no longer recognizable as such. The classifies -
tory value of the presence or absence of marginalia is discussed else-
where (under Cryptozonia), and as the inframarginals are seemingly
or actually lost independently in a number of phyla the term is
here used as expressive of this condition, and not necessarily of
relationship.
Spencer (1914:9) takes up the origin of the wriggling habit from
another point of view, that is, from a study of living Stelleroidea as
described by MacBride. The former states that the living forms can
be grouped into two divisions as follows :
The graspers. Asteroid forms in which the tube-feet are used for walking, and for
grasping and pulling open the bivalve shells of the mollusks upon which they usually
feed. The ambulacralia form an arch to take the pull.
The wrigglers. Ophiuroid forms in which the tube-feet have lost locomotory powers
and become much reduced. The animals progress by wriggling movements of the
arm, and the ossicles of the ambulacral groove are extensively modified for this pur-
pose. The food is pushed into the mouth by the first two pairs of tube-feet, which
become considerably enlarged and are known as buccal tentacles.
If we trace the history of the forms backward we find that the difference between
them tends to disappear. Both the ' ' graspers } ' and the ' ' wrigglers ' ' descended from
a third group, which I call provisionally "the primitive Asterozoa."
The postulated " primitive Asterozoa" above referred to are based
upon the earliest stages of growth of starfishes, and had the following
characters. The animals were attached to the ground by a well-
developed, flexible stalk (seen in Asterina and Asterias); the disk
was small compared with the five arms. In connection with the
water-vascular system there were two series of ossicles, (1) a double
column of flooring pieces forming the sides of the ambulacrum, and (2)
a paired covering series as a protection to the soft structures under-
neath. The podia emerged between the flooring pieces and these at
first "were arranged not exactly opposite to each other, but slightly
alternating, and in consequence we find that both the flooring and
covering plates, which are in direct association with the podia, are
not exactly opposite, but arranged alternately. "
34
BULLETIN 88. UNITED STATES NATIONAL MUSEUM.
The rays terminated in a single ossicle with an unpaired tube-foot
that later developed at its outer end a pigment spot and finally an
"eye."
At last the animal lost the stalk and moved about freely as the
primitive asterozoon. The flooring plates changed into ambulacralia
and the roofing plates into adambulacralia, while the tube-feet were
used not for grasping and crawling but for passing small particles of
food to the mouth.
Inf
Ax
FIGS. 1 AND 2. VENTRAL AND DOKSAL VIEWS OF THEORETIC PHYLEMBRYO OF STELLEROIDEA, INDICATING
HOW THE RAYS ARE INTRODUCED. BASED ON HUDSONASTER. A d, ADAMBULACRALS; Am, AMBULA-
CKALS; AX, MARGINAL AXILLARIES; CZ>,CENTRO-DORSAL; /72/,INFRAMARGINALS; Rl , PRIMORDIAL RADIALS:
R2 TO Rll, SUBSEQUENT RADIALSJ Sml , DORSAL INTERRADIALS OR PRIMORDIAL SUPRAMARGINALSJ Sm2
TO SmlO, SUBSEQUENT SUPRAMARGINALS.
The asterid radicle. Hudsonaster is held to be very near the radicle
that gave rise through modification and inheritance to all subsequent
Stelleroidea. A study of the various species and specimens of Hud-
sonaster, reinforced by the chronogenesis of this genus and the de-
velopmental stages in the individuals, which are discernible in the
youthful distal ends of the rays when contrasted with the mature
proximal region of the same, has led the writer to speculate on the
probable skeletal characters of the radicle of the Stelleroidea. An
analysis of mature H. matutinus and of half-grown H. incomptus
shows that their ancestors must have been devoid of all disk accessory
pieces, or for that matter of all secondary ossicles, since it appears
that it is on the disk that these plates first arise. Further, the
same material indicates that the younger the individual specimen,
or the older the species geologically, not only the smaller is the
specimen, but the fewer plates has it in any column. As the
REVISION OF PALEOZOIC STELLEROIDEA.
35
ossicles, other than those of the disk, the interbrachial areas, and the
accessory pieces, are introduced in all the columns at the distal ends
of the rays, we are permitted, in theorizing as to the characters of the
radicle, and from what has been learned from Hudsonaster, to elimi-
nate all of the rays with their ventral and dorsal ossicles down to
the disk. If this is done, there finally results a tiny depressed and
mammillated spheroid, made up dorsally of 11 primary plates, and 5
marginal axillaries, and ventrally of a limited number of adambu-
lacrals and ambulacrals around the mouth. To make this matter
clearer, two diagrams are presented of the probable characters of the
direct ancestor of Hudsonaster (see
figs. 1 and 2) .
By taking away the rays down to
near their bases, i. e., to R2, and
correspondingly decreasing the size
of the primordial supramarginals, we
get the dorsal structure of a late stage
of the probable typembryo of the
Stelleroidea, as illustrated in fig. 3.
If all of the ray plates are com-
pletely removed, we apparently get
the typembryo of the class Stelle-
roidea, as iUustrated in figs. 4 and
5. It Should be Stated here, how- FlG - S.-DORSAL VIEW OF THEORETIC TYPEM-
,, t , , . ,, BRYO OF STELLEROIDEA AT THE BEGINNING
ever, that the number of the adam-
bulacrals and ambulacrals in figure
5 is entirely conjectural, though
these plates and those of the mouth
frame are among the first to develop
in living forms.
If the developmental views just stated are correct, it follows that
the five branches of the water- vascular and nerve rings of the typ-
embryo of Stelleroidea grow outward along the radii, developing as
they grow the rays or arms with their dorsal and ventral skeletons.
In other words, the growing tips of the radii spread outward beyond
the disk, and each one gives rise to nine columns of ossicles (one
radial, two supramarginal, two inframarginal, two adambulacral and
two ambulacral) . The radial plates continue as simple columns, while
upon the primordial supramarginals and the axillaries rest pairs
of columns, the supramarginals and inframarginals, respectively.
In Echini, however, the growing tips of the radii do not spread
beyond the disk (apical disk of oculars and genitals) but remain
internal and give rise to new ossicles, the ambulacrals (same ossicles
in Stelleroidea) and interambulacrals ( = adambulacrals of Steller-
OF RAY DEVELOPMENT. THE PRIMORDIAL SUP-
RAMARGINALS HERE HAVE ON EACH SIDE THE
BASAL PIECES OF TEN SUPRAMARGINAL COL-
UMNS. CD, CENTRO-DORSAL; Rl, PRIMORDIAL
RADIAL; R2, SECOND RADIAL; Sml, DORSAL IN-
^ERRADIAL OR PRIMORDIAL SUPRAMARGINAL;
82, BEGINNING OF SUPRAMARGINAL COLUMNS.
36
BULLETIN
UNITED STATES NATIONAL MUSEUM.
oidea), all of which appear from underneath the ventral edges of
the oculars, each plate in turn being pushed orally 1 .
In any event, whatever the pre-Hudsonaster evolution was, we
are on safe ground in indicating what the developmental changes
were, starting with this genus. We will not take up all of the skele-
tal parts and discuss them in detail, but will attempt only the more
essential portions.
Ontogeny. Ontogenetic stages of development in the ossicles
of asterids can readily be made out to some extent in any well pre-
served specimen. The variations can best be seen in those forms
with the least number of plates and therefore in the more primitive
genera. Here all of the primary ossicles are introduced at the tips
of the rays in the seven primary columns. The youngest pieces are
also the smallest and the simplest. At first they are globular,
FIGS. 4 AND 5. DORSAL AND VENTRAL VIEWS OF THEORETIC TYPEMBRYO OF STELLEROIDEA. Ax, VEN-
TRAL OR MARGINAL AXILLARIES OR BASAL INFRAMARGINALS; CD, CENTRO-DORSALJ R, PRIMORDIAL
RADIALSJ Sml, DORSAL INTERRADIALS OR PRIMORDIAL SUPRAMARGINALS.
smooth, and loosely adjoining. Tracing them down the columns,
the specific characters are seen to develop and finally the basal pieces
are found to be the most modified of all. This is particularly true
of the dorsal columns where the most ornate plates are the oldest.
In the different species of Hudsonaster one can also trace the changes
chronogenetically, as for instance in the basal radialia and supra-
marginalia. In the oldest species these plates are not at all, or but
very slightly stellate, more so in H. incomptus, and most so in the
youngest and largest species, H. rugosus.
In the stocks with more complex skeletons, the ontogenetic varia-
tions in the plates can be traced in the same way. However, as these
forms almost always have more or less of accessory plates that are
introduced nearly throughout the entire skeleton during the juvenile
growth and even at maturity, the study is complicated by the inter-
mixture of ossicles of varying age.
i See Jackson, Phytogeny of the Echini, Mem. Boston Soc. Nat. Hist., vol. 7, 1912.
REVISION OF PALEOZOIC STELLEROIDEA. 37
Spinosity can also be studied ontogenetically because the youngest
ossicles do not have spines, and when these appear they are smaller
and less characterized than the older ones situated in the proximal
portions of the columns.
Many of the ontogenetic changes observed by the writer are re-
corded throughout this work, and their significance in related species
and genera has been made use of in discerning the phylogenetic
lines of descent.
Regeneration. Among living starfishes it is common to regrow
arms that have been lost through accidental causes. From the
base of the severed ray a new growing tip is established, forming
a juvenile arm that gradually grows to full size and assumes
mature characteristics. Schondorf (1909a: 96-97) states that this
habit has been pronounced since the Jurassic but that he has failed
to find marked regeneration in Paleozoic asterids. He did, how-
ever, note partial replacement of minor losses among the Devonic
species. Stiirtz, who has handled more Paleozoic asterids than any
other paleontologist, also has not noted a single case of marked
regeneration. The same is true for the 400 Devonaster eucJiaris
found in a limited area of the Middle Devonic of New York.
Clarke, in describing this find (1912: 44-45), however, does note a
few specimens " which show the existence of only four instead of
the normal five arms." These are the only examples of four-rayed
Paleozoic starfishes so far recorded.
The writer has also been unable to find a single case of regenera-
tion, but in the Middle Ordovicic cryptozonian Urasterella ulrichi
he describes a specimen with two normally developed rays and three
short stumps. All of the arms are normal for the species, except
for the length of three rays and their terminations, which are blunt.
(See pi. 29, fig. 1.) It seems to him that this occurrence is not due
to the accident of fossilization or weathering, but is an actual case
of loss in life with subsequent healing of the wounds, but without
regeneration of the lost parts.
During most of the Paleozoic, the starfishes could have had no
carnivorous enemies other than the cephalopods; as for marine
fishes, the armored Arthrodires did not appear until the Middle
Devonic, while the ancient sharks were not common until Lower
Carboniferous (Mississippi) time. It is possible that regeneration
among the starfishes is connected with the rise of carnivorous ene-
mies, but as the habit is so common among living forms it is more
probable that this power has always been inherent in the class.
Regeneration among the crinids has been noted in several cases
where lost distal ends of arms were being replaced by immature
growths. Such have been seen in the Lower Carboniferous (Bur-
lington and Keokuk formations) of America.
38 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Number and character of rays. In the geologically oldest and the
most primitive asterids as well there are always 5 rays, no multirayed
form being known back of the middle of Siluric time. In the most
primitive genus of Phanerozonia, Hudsonaster, the rays are short and
stout, there is a large disk without interbrachial arcs, and the skele-
ton is simple but ponderous. From this type originate those with 5
slender rays and also forms with more or less large interbrachial arcs,
the pentagonal starfishes.
Long, slender, and flexible-rayed genera without marginalia are
present as early as the most primitive Phanerozonia, i.e., as early as
the older Middle Ordovicic, indicating that the two main types of
asterid development arose still earlier. The same is true for the
ophiurid ancestors (Auluroidea). In fact, slender-rayed Crypto-
zonia are more commonly seen in the Middle Ordovicic than are the
Phanerozonia.
Multi-rayed Phanerozonia with primitive arm construction appear
in the Middle Siluric (Wenlock) in Lepidaster. This genus has rays
which are short and stout, reminding one in this respect strongly
of the primitive 5-rayed forms. An insufficiently described 10-
rayed form (Trichotaster plumiformis) is recorded from the Middle
Siluric (Wenlock) of England. In Helianihaster of the Lower Devonic
there are from 14 to 16 rays and here they are long and slender,
while in the Upper Devonic there is a form with only 1 1 rays. Lepid-
astereUa of the Upper Devonic has 24 arms.
All of the multi-rayed Cryptozonia have slender rays and rather
large disks. None are older than the Lower Devonic, where Medusaster
has from 12 to 15 rays, Echinasterias 25, Palxosolaster 27 to 29
(an Upper Devonic species referred doubtfully to this genus has 24
or 25) , Echinostella 28 or 29, and Echinodiscaster 29.
Curiously, no multi-rayed starfish is known hi the Paleozoic later
than the Devonic. Further, the arms may be either odd or even in
number in the same genus or even species. How the living multi-
rayed starfishes introduce their rays is explained under Palseosolas-
teridse (p. 207).
Verrill (1914: 12-17) holds that the multi-rayed forms arose as
" ' sports/ which have persisted by heredity and natural selection
because they were advantageous." The advantage lay in "the
increased number of ambulacral sucker-feet. " Most of these forms
are now living in shallow water among the rocks exposed to the surf.
He states: " This is true of the seven species of Heliaster, with very
numerous rays; and of Pycnopodia, with 20 to 24 rays; and of the
various shallow-water and littoral species of Solaster and Crossaster,
which usually have 9 to 15 rays (rarely 8 or less). It is also true of
the numerous 6-rayed species of Asterias, Pisaster, and allied genera."
REVISION OF PALEOZOIC STELLEKOIDEA. 39
On the other hand, the increase of rays is more probably due " to
the advantage gained in holding their food securely, and in opening
bivalves, than for holding to the rocks, though both go together."
"The ability to cling tenaciously to rocks may be perfected in
other ways, involving equally an increased number of sucker-feet.
This is often attained by lengthening the rays, * * * by crowding
the suckers into more than four rows, * * * and by increasing
the size and strength of the suckers."
DEVELOPMENT or THE PARTS.
Anus. In all living starfishes the anus apparently lies in the next
interradius to the right of the madreporite, or in the direction as the
clock goes, but in no Paleozoic asterid is there positive evidence of
an anal opening. However, it is extremely rare in these forms to find
the disk so well preserved as to make it possible to locate so small
an orifice. In certain specimens of Hudsonaster, immediately adj acent
to the centro-dorsal plate, there is an open space that either is the
anal opening or where a plate has dropped out in fossilization.
Schondorf has also failed to find this structure in the German Paleo-
zoic asterids, but has indicated its probable position in Spaniaster
(see fig. 8) and Calliasterella.
Madreporite. In more than half of the Paleozoic asterids the
madreporic plate has not been seen. In many cases this is due to the
infallen condition of the disk skeleton and the jumbled state of the
ossicles; in others it is due to the fact that it is so small as not to be
readily distinguished among the mass of other small plates. Finally,
a great number of asterids are known only from the actinal side.
The madreporite in size varies from minute to very large and
ponderous. It may be round, oval, or many-sided, concave, and
depressed beneath the disk pieces, or protruding cone-shaped (Petras-
ter) . As a rule, it is marked by striations that are coarse or very fine,
straight or wavy lined, or it may be crenulostriate or with granular
surface. In Petraster the surface canals enter into two spirals that
carry the water into the stone canal. In living forms with more
than six rays there may be two or more madreporites, but in no
Paleozoic species has more than one been seen.
In all of the Paleozoic phanerozonians the madreporite appears
to be always dorsal in position. In the most primitive genus, Hud-
sonaster, the plate may be large and conspicuous or at times can not
be made out even when the disk is fairly well preserved. In no
Mesopalseaster has this sieve plate been seen, although good speci-
mens are at hand, a condition probably due to its small size among
the small skeletal pieces of this genus. In Spaniaster it is very small.
In Promopalseaster the madreporite is usually conspicuous and large,
40 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and the same is true for Xenaster and Agalmaster. The plate is also
known in Palseaster, Devonaster, Neopalseaster, and Palasterina.
In multi-rayed phanerozonians the madreporite is unknown in
Lepidaster and Lepidasterella. In Helianthaster it is large, conspic-
uous, and situated on the edge of the disk, more actinal than abac-
tinal. On the other hand, in the cryptozonian multi-rayed genera
the madreporite is apparently always actinal and more or less large,
as in Palseosolaster and Echinodiscaster. This plate is probably also
always actinal on large-disked Cryptozonia such as Palasteriscus
(plate enormously large), and OJieiropteraster (here small and situ-
ated near the mouth) .
Among the five-rayed Paleozoic cryptozonians the madreporite
is known only in Monaster (large) , Urasterella (medium) , and Jaekel-
aster (small) . In all it is always on the abactinal area.
From this evidence we see that in the five-rayed Phanerozonia,
forms that are more or less rigid, the madreporite is always on the
dorsal or upper side of the animals, this being true for the deeply
stellate as well as for the pentagonal species. In the multi-rayed
genera, however, there is a marked tendency for this plate to move
outward to the edge of the disk. In the five-rayed cryptozonians
the madreporite so far as known is always dorsal, but in the large-
disked or pentagonal forms and the multi-rayed species this plate is
apparently always ventral.
Spencer (1914:35-38) says that "many Paleozoic Asterozoa do not
appear to have a madreporite" and he is "inclined to regard this,
in these old forms, as usually a primitive feature." The present
writer, it is true, states above that in more than half of the Paleo-
zoic asterids the madreporic plate has not been seen. This is due in
most cases, however, to poor preservation, and in others to the fact
that the plate is so much like the other disk plates as not to be dis-
tinguishable from chem. Spencer concludes further that it appears
that the madreporite originated on the ventral side, or at least was
marginal, in the primitive forms, and that it passed over in later
Asteroidea to the dorsal side. In Eudsonaster we have the most
primitive known starfish, and here in H. incomptus it is a large and
conspicuous plate on the abactinal side. Further, the evidence of
the Paleozoic starfishes (Phanerozonia), so far as the present writer
knows them, is that this plate is always dorsal in position, and he is
inclined to the view that all of them had the madreporite, that it
originated on this side, and that in later cryptozonian multiradiate
forms it moved outward to the margin of the disk or to the actinal
surface. In regard to the Auluroidea, the writer has seen the
madreporite so rarely that no deductions as to its original position
in these forms can be made.
REVISION OF PALEOZOIC STELLEROIDEA. 41
Spinosity. In general, it can be said that spines are least developed
in the heavily plated Phanerozonia, more so in those forms where
the dorsal skeleton is made up of small ossicles in abundance, and
fully in most of the Cryptozonia stocks.
In primitive Hudsonaster, all of the plates other than the am-
bulacrals are more or less well tuberculated, and these granules do
not now carry spines, if they ever did. In later forms, however, as
Promopalseaster, they may all have borne spines, some of them of
large size. Small club-shaped spines occur here in considerable
abundance along the sides of the inframarginals and the adambula-
crals. In Palseaster there are brushes of delicate spines along the inner
edge of the adambulacrals. In Mesopalseaster the ventral spines are
better developed, fully so in Promopalseaster, and probably equally so
in most post-Ordovicic forms.
Among the primitive cryptozonian genera Stenaster and Tetraster
no spines are now seen, but undoubtedly such were present on the
actinal side. In Ur aster ella the entire dorsal skeleton is more or less
spinose, consisting of long, nonarticulating, slender rods, the dorsal
extensions of the ossicles. In the multi-rayed Cryptozonia of the
Devonic, spinosity is at its greatest development in the Paleozoic.
Ambulacralia. Undoubtedly the most important skeletal parts of
the Stelleroidea are the ambulacrals. In general they undergo the
least alteration during geologic time of the entire asterid skeleton,
and therefore any marked variation must be of broad classificatory
value. Schondorf * is well aware of this fundamental value and has
made full use of it in denning his three " classes, 57 Asteroidea, Aulu-
roidea, and Ophiuroidea. In the Asteroidea the ambulacralia of ad-
joining columns are nearly always placed directly opposite one
another, with the podia issuing through openings that are not in the
plates themselves, but laterally between two adjoining ambulacrals
and the corresponding adambulacrals. Among the Paleozoic asterids
one is not always certain whether the ambulacralia are arranged
lt opposite" or "alternate," because the specimens in nearly all cases
have suffered more or less from distortion. In many good specimens
they are very slightly alternate, but in all such cases the arrange-
ment is said to be alternate. In other fine fossils they are now
alternate, but a close study seems to show that originally they were
practically opposite in arrangement. On the other hand, certain of
the species with wide ambulacral furrows and large ambulacralia, as
in Promopalseaster, Anorthaster, and Urasterella, have an alternate
arrangement, while in other species of the first and last named genera
they are just as certainly opposite. The writer therefore does not
i Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, pp. 206-256; Palaeontographica, vol. 57, 1910,
pp. 1-56.
42 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
lay much stress upon this feature among the Ordovicic asterids, nor
does he regard it even necessarily as of specific value, but in the later
Asteroidea the arrangement is always opposite or nearly so. Ver-
rill (1914:20) states that he has noticed the alternate arrangement
in living Pycnopodia "as an abnormal variation in some of the
rays, and also that it may occur from lateral bending." Evi-
dently the alternate disposition is a fixed character among some
Ordovicic forms, as in AnortJiaster, but more will certainly be dis-
covered in these old strata and probably rather among the Crypto-
zonia than the Phanerozonia. The writer is led to this view because
of the alternate arrangement in many Auluroidea, a subclass of
Stelleroidea that seems to have had its origin in some cryptozoniaii
asterid. Out of the Auluroidea with opposite ambulacralia came the
Ophiuroidea with their highly modified ambulacrals which are
coossified into " vertebral ossicles." In the Auluroidea the arrange-
ment of the ambulacrals is either alternate or opposite, but they are
never coossified but are so modified as to suggest the Ophiuroidea
rather than the Asteroidea. The structure of these plates is de-
scribed elsewhere and need not be repeated here.
The ambulacrals are very variable in number throughout the
genera. In Hudsonaster, Stenaster, and Tetr aster there are as few as
10 in a column, while in Promopals&aster there are certainly 50 and
possibly as many as 60. The cryptozonian genus Urasterella has
certainly over 100 in a column. In general, these ossicles abut
against each other, but in Urasterella and other genera they overlap
proximally to a certain extent. As yet the writer has seen no
Paleozoic asterid with more than 2 columns of ambulacrals, though
in Promopalseaster magnificus the proximal areas of the rays for a
short distance have 4 columns of podial openings. Here, however,
the ossicles have not yet wedged themselves into 4 columns of
ambulacralia.
In shape and surface contour there is also considerable variation
among the ambulacralia. In Hudsonaster, Tetraster, and several
Devonic genera they are more or less rectangular, either squarish or
drawn out laterally. In other forms of Hudsonaster, along with
Stenaster and Urasterella, the ossicles are h- -shaped, with the lateral
shaft thin, thus allowing for large, elongate podial openings. The
more primitive genera have the outer surfaces plain or slightly ridged,
while in derived forms like Promopalseaster the ridges are high and
straight or have their medial terminations orally directed. In
P. magnificus, the ridges in the proximal region are arranged in pairs
having the shape of a tuning fork; in P. wyleqffi one sees that these
ridges approach one another in pairs transitional to their transforma-
tion into the tuning-fork form.
REVISION OF PALEOZOIC STELLEROIDEA. 43
Adambulacralia. Adambulacralia are present in all Stelleroidea.
They are best and typically developed in the Asteroidea, more or
less modified in the Auluroidea, and completely altered into the thin
scalelike side plates of the Ophiuroidea. In number and relative
size they vary as do the ambulacralia, being least numerous and
largest in the most primitive genera. Originally they probably
bore no spines at all, the plates being practically smooth. In Hud-
sonaster and most Ordovicic and Siluric, and some Devonic genera,
their surfaces are granular to tubercular. These protuberances in the
younger genera may have borne small spines, but long before these
appeared each adambulacral bore two or more stout or slender, com-
paratively long spines along its ambulacral edge.
In most of the Paleozoic genera there are in each column as many
adambulacrals as there are ambulacrals, and both series are as a rule
arranged opposite one another. In Anorfhaster, however, there are
fewer and therefore larger adambulacrals than ambulacrals, and
these latter ossicles clearly alternate with one another.
In the Phanerozonia the adambulacrals never margin the rays
but always lie inside of the bordering inframarginals. In the Crypto-
zonia, however, the adambulacrals margin the animals and here the
ossicles are usually small, though at times they are relatively large
and make a stout outer skeletal frame as in Stenaster, Tetraster,
and Schcenaster (?} montanus. The same is true in Encrinaster
of the Auluroidea.
The adambulacrals in probably all the Paleozoic Phanerozonia
continue into the oral region and each two adjoining columns meet
here in a pair of modified, elongate, pointed pieces, the most promi-
nent ossicles of the oral armature. No other skeletal parts lie in
front of these oral ossicles except in the phanerozonian Hudsonaster
narrawayi (pi. 1, fig. 1) and in the young of the cryptozonian
Urasterella ulrichi (pi. 30, fig. 7). Both are primitive forms of
their respective phyla and whether these five pieces or tori are to
be interpreted as five spines or whether they represent five primi-
tive orals is not yet determinable.
Inframarginalia. In the great majority of Paleozoic Phanero-
zonia the inframarginals alone margin the animals, and only in a
few forms (Spaniaster, Miomaster, RJienaster, and Neopalseaster)
have the supramarginals moved outward and completely covered
the inframarginals, so that the two columns together equally bound
the rays and disk. That condition is a peculiarity common to most
Paleozoic phanerozonians, while the wholly superposed arrange-
ment distinguishes nearly all the Mesozoic, Cenozoic, and Recent
genera of the same kind of Asteroidea.
The inframarginals are usually the most conspicuous ossicles of
the Paleozoic Phanerozonia, and this is especially true where the
44 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
skeleton is strong and devoid, or nearly so, of accessory ossicles.
They are ponderous in Hudsonaster, Palseaster, Australaster, Devon-
aster, Xenaster, Trimeraster, Petraster, etc. When the smaller
accessory plates are introduced, the inframarginals also reduce in
size; this development is best seen when passing from the heavily
plated Hudsonaster to the more abundantly plated Mesopalseaster
and finally Promopalseaster. The next step in this phyletic line
would be pronounced a cryptozonian because the infra- and supra-
marginals could not be distinguished from the abundance of acces-
sory ossicles. In fact, it is not always easy to make out the various
categories of plates even in Promopalseaster, and here one must
resort at times to a study of the tips of the rays where the seven pri-
mordial columns are not yet disturbed by the introduction of accessory
pieces.
In primitive Hudsonaster, where the inframarginals are compara-
tively very large, there are only half as many of these ossicles as
there are of adambulacrals, but in the great majority of Paleozoic
genera of Phanerozonia this marked difference tends to be less
decided and toward equalization. In certain phyla, as, for instance,
from Hudsonaster to Mesopalseaster and Promopalseaster, the equal-
ization has gone on rapidly, hi fact, in but a short part of Ordovicic
time. In more conservative stocks, as from Hudsonaster to Devonic
Xenaster and Trimeraster, the equalization is far slower, as is seen
in the following figures: Hudsonaster, 12 inframarginals and 24
adambulacrals; Trimeraster, 14 and 23; and Xenaster, 20 and 32,
respectively. In nearly all the genera these columns of plates alter-
nate with one another.' This of course must be so where the number
of ossicles is different in the two columns, and complete alternation
is retained even in certain species where the numbers of ossicles are
alike, as for instance in Promopalseaster spinulosus. On the other
hand, in Mesopalseaster sJiafferi, where there are 16 inframarginals
and 18 adambulacrals, the plates alternate in the proximal half of
the ambulacra and are opposite one another distally. A study of
the growing ray tips shows that the various ossicles all appear
practically of one size and where there are more adambulacrals than
inframarginals, the former are crowded orally. The latter develop-
ment appears to be the primitive type of growth, but this mode is
rapidly changed to one in which the seven columns of primordial
ossicles appear in cycles or rings. This type of growth is easily made
out in primitive forms such as Hudsonaster and Mesopalseaster, and
in some species of Promopalseaster. However, when accessory plates
are developed in abundance, and especially in forms where the seven
primordial columns of plates remain of one size, their introduction
in cycles can only be made out in very young individuals or at the
KE VISION OF PALEOZOIC STELLEKOIDEA. 45
extreme tips of the rays, conditions not often attained among these
fossils.
In the great majority of Paleozoic Phanerozonia the inframar-
ginals and adambulacrals lie closely adjacent to one another. In
typical asterids they are not separated by the introduction of acces-
sory interbrachial pieces until in the Devonic. Here in Xenaster
and Trimeraster isolated pieces, and finally a column of them, are
inserted in the proximal half of the rays. In the large-disked asterids,
this separation occurs much earlier, in the Middle Ordovicic in
Petraster.
In Paleozoic Phanerozonia where the skeleton is made up of an
abundance of more or less equal-sized ossicles, one notices also on
the distal portions of the rays a tendency for the inframarginals to
appear on the dorsal side and not to remain on the ventral, as in
primitive stocks. This alteration brings the adambulacrals to the
margin of the rays, and is a hint as to how Phanerozonia gradually
change into Cryptozonia. It is best seen in Promopalseaster, Petras-
ter, and Mesopalxaster. In Australaster it is a marked feature, in
fact, one can say that the distal parts of the rays in that genus are
cryptozonian while the bulk of the arms are still phanerozonian.
This ontogenetic appearance is in harmony with phylogenetic devel-
opment and chronogenesis. The living Echinaster sepositus is in
youth also a phanerozonian, but at maturity is a typical cryptozo-
nian. In so typical a cryptozonian as Ur aster ella, one still finds in
half -grown U. ulrichi a number of inframarginals in the interradial or
axillary areas. The same retention of the first formed inframargin-
als is also met with among the multi-rayed Cryptozonia in Helianih-
aster, and even among the Auluroidea in Encrinaster.
It has just been pointed out how the inframarginal columns are
eliminated as marginal plates. They are not, however, removed
from the skeleton in these early forms by resorption or by failure
of development, but for want of special growth. In other words,
the inframarginals are probably present in all Paleozoic Asteroidea,
but because of lack of specialization through the developmental
tendency to greater skeletal flexibility, remain small and are lost
as such in the mass of the dorsal plates. This apparent eh'mination
of the inframarginals has gone on independently in various stocks
as pointed out elsewhere, and therefore the absence of large marginals,
either infra- or supramarginals, or both series, is not of ordinal value.
Supramarginalia. In Hudsonaster the prominent supramarginal
plates of the dorsal side are placed decidedly inside of the inframar-
ginals, though the former clearly overlap the latter. This primitive
position is retained in many Paleozoic genera, and apparently not
before the Devonic do these two columns of ossicles come to lie
wholly upon one another, and then they together margin the animals.
46 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
It is apparently always a rare condition in the Paleozoic, but as the
post-Devonic asterids are as yet little known, we can more accurately
say that the modern type of Phanerozonia is unknown before Devonic
tune. It should be added here that complete superposition of the
supra- upon the inframarginalia undoubtedly took place in more
than one stock and at different times. Accordingly, to group all
asterids with large marginalia into the " order" Phanerozonia is
further proof that these terms cannot be used in the phyletic sense.
Phanerozonia and Cryptozonia are, however, of use in a descriptive
way.
The supramarginals, and the inframarginals as well, in primitive
forms gradually attain larger size down the columns from the tips
of the rays toward the disk and mouth. In other words, the older
the ossicle in primitive genera, the larger is its size, and in all such
forms the pairs of supra- and inframarginal columns terminate in
single plates, the primary or basal supramarginals and the primary
or basal inframarginals. The latter ossicles in the most primitive
genera are situated in the axils of the rays and are therefore also
known as axillary marginals, and in the more specialized genera the
axillaries and even pairs of inframarginals are crowded orally into
the interbrachial areas. The basal supramarginals in primitive
forms always occur prominently on the disk, in derived genera they
are, however, less conspicuous, and where the disk skeleton is made
up of small ossicles, they are generally indistinguishable in the mass
of plates.
We have seen that accessory interbrachial plates in some genera
are developed between the inframarginals and adambulacrals, and
to a greater extent the same is true of the development of ambital
accessory pieces between the former columns and the supramarginals.
In probably all cases where the rays and disk grow wider the addi-
tional space is largely covered in by the development of accessory
plates. As it is in the disk area, and therefore necessarily in the
basal portion of the rays as well, that the main increase of body
extent takes place, here are developed the greatest number of these
pieces. Rarely are the ambital accessory pieces crowded in between
these columns beyond the proximal third of the rays. This is best
seen in Devonaster and Xenaster, but in the large-disked forms, as
Petraster, they extend farther out and are present to within the
outer third of the rays. Ambital accessory plates may also be
present in Mesopal&aster, but are more definitely so in Promopalse-
aster (especially in P. magnificus) , and in both genera they occur
throughout the greater part of the rays.
Radialia. In many Paleozoic genera of Phanerozonia and even
among some of the Cryptozonia, a more or less prominent column of
ossicles is seen down the center of the dorsal side of the arms. These
REVISION OF PALEOZOIC STELLEEOIDEA. 47
are the radial columns of ossicles, which are especially well seen in
Hudsonaster, Spaniaster, Devonaster, Xenaster, CaUiasterella, and
less prominently in Mesopalseaster, Promopalseaster, and Petraster.
The prominence of the radials is lost in the same way as has been
described for similar reduction in the supramarginals and inframar-
ginals.
In primitive genera the supramarginals and radials lie closely
adjacent as prominent columns, but in the great majority of Paleo-
zoic Asteroidea these columns are more or less extensively separated
from one another by the introduction of radial accessory plates. The
latter are always small pieces and in general one can say that their
number increases with geologic time and with the evolutionary
complexity of the various stocks. At first the accessory pieces appear
singly between the plates in the proximal region of the rays, where,
increasing in numbers, they separate the primary columns more and
more and finally also crowd in between the individual ossicles, es-
pecially the proximal radials, pushing these apart more or less com-
pletely. This is true not only in chronogenesis but also in onto-
genesis. It is best seen in a study of the dorsal areas beginning with
Hudsonaster and Spaniaster, and proceeding to Mesopalseaster,
Devonaster, Promopalseaster, and Urasterella. In the forms with
great disks the development of accessory plates is extreme, as may be
seen in Petraster speciosus.
The columns of radial plates are less persistent in development
than any other of the primary columns. Like all of the primary
plates, the radials are apt to lose their ponderosity and in the course
of geologic time become smaller. In this way they may lose their
individuality in the mass of dorsal plaies, but in some cases radialia
appear to fail to develop throughout the greater part of the rays,
although one or more primordial ossicles of each radius are present
on the disk. In certain species of Mesopalseaster, and more especially
in Promopalseaster, the radialia are either not present or have been
squeezed out of the mid line of each radius so as to be unrecognizable.
In other forms of the latter genus they are well developed and arranged
in linear columns. Promopalseaster as a rule has no radialia on the
rays and the space between the supramarginals is occupied by many
small accessory radial pieces. Among the Cryptozonia, radialia are
usually not discernible as such on the rays, though present on the disk.
Accessory plates. These small pieces of irregular form are space
fillers between all of the primary columns of plates, with the exception
of the ambulacrals and adambulacrals. As the body cavity grows
larger, skeletal increase takes place either through enlargement of
the seven primordial columns of ossicles or by the intercalation and
formation of new pieces, the accessory plates. The former method
50601 Bull. 8815 4
48 BULLETIN
is the one least often resorted to. The accessory pieces appear first
chronogenetically on the disk, separating the primordial ossicles
one from another in various ways, and as the body cavity grows
larger in the course of geologic time, they wedge all of the primary
columns apart proximally except the ambulacrals, which compensate
for the widening of the rays by growing in size transversely. Some
accessory plates also develop in the interbrachial areas separating
the interbrachial marginals, as in Trimeraster and Xenaster, while
in the large-disked forms, as Petraster, their number becomes excessive.
Ocular plates. The large ocular plates seen in so many living
Phanerozonia are very rarely developed in Paleozoic forms. In
fact, the writer knows of but a single occurrence, in the Lower Car-
boniferous genus Neopalseaster.
It seems to the author that these ossicles should have considerable
significance in phylogeny because their large growth indicates that
they are much older plates than any of the small adjacent pieces.
In other words, in all genera where the ocular or sensory tentacle
does not lie in an enlarged and grooved plate, the newly developed
ossicles of either the ventral or dorsal side appear distally to the pre-
viously borne pieces, i. e., at the tips of the rays, whereas in the stocks
having ocular plates the new skeletal arrivals appear not at the tips
of the rays but on the proximal side or inside of the eye ossicles
Therefore, where ocular plates are present, these pieces are not only
enlarged in size with age, but progressively pushed farther and farther
outward as well by the growing ray tips that lie on their inner sides.
This mode of growth may have originated once or several times,
there being in the latter case parallel developments of ocular plates.
This is a study that can not be pursued far among Paleozoic genera,
but in the later forms may be productive in the discerning of
phylogenetic lines.
After the above was written the writer read in Verrill (1914: 20-22)
that Fewkes has shown the ocular plates to be the first to appear in
the young, and that with growth they are pushed distaUy because
of "the budding in of new plates between the apical plate and the
one next to it. 7 ' These observations on living starfishes are therefore
in harmony with the study of Neopalseaster, and go to show that the
bulk of Paleozoic starfishes insert the plates of the primary columns
in a different way, i. e., always distally. On this ground alone most
of the Paleozoic starfishes should be grouped by themselves and
apart from most of the subsequent Phanerozonia.
Centro-dorsal. In the center of the disk of many Paleozoic genera
of Asteroidea and of some Auluroidea is seen a more or less prominent
plate, which is thought to be the equivalent of the centro-dorsal of
the embryo of the living crinid Antedon (Comatula). It is a promi-
nent though small plate in most of the forms of Hudsonaster and in
REVISION OF PALEOZOIC STELLEEOIDEA. 49
Palxaster; it is also well developed though less conspicuous in Meso-
palseaster, Spaniaster, Xenaster, Agalmaster, Devonaster, Neopalseaster,
Palasterina, Monaster, CalliastereTla, and Urasterella.
As the disk plates are rarely well preserved, and as the centro-
dorsal has been seen in many Paleozoic species, the writer thinks-
it will be found in most of the forms of this era.
Primary disk plates. In the most primitive and oldest Paleozoic
asterids the centro-dorsal is surrounded by a variable but small
number of diminutive accessory disk pieces. In mature Hudsonaster
they form a single ring, but in the young of the cryptozonian Uras-
terella ( U. ulrichi) and in mature CalliastereTla there are none of these
accessory pieces present. For these reasons it is thought that in
the Ordovicic there will be found a small asterid, even more primitive
than Hudsonaster, that will be devoid of accessory disk pieces.
Around the centro-dorsal of young Urasterella and in mature
Calliasterella, there follows directly a ring of five larger plates. The
same five plates are also seen outside of the single ring of accessory
pieces in Hudsonaster and Spaniaster, and directly beyond the two
rings of accessories in Mesopalseaster. These more or less large
plates are radial in position and are the basal radialia above which
continue the columns of successively formed radials.
Keturning to Hudsonaster, we see that the first ring of prominent
plates has 10 pieces, 5 of which are radial in position, the remainder
interradial. The same is true for Spaniaster and Xenaster, while
Calliasterella has an equal number of plates in the second ring. An
analysis of these disks shows that 5 of the plates are either primary
or secondary radialia, while the remainder are situated interradially,
and upon them rest the 10 columns of supramarginals. The first
prominent ring in Mesopalseaster sJi'Offeri and Devonaster and the
second ring in Urasterella ulriclii each have 15 plates, 5 of which
are radialia, while the other 10 are supramarginals. Simplifying
these statements and cutting out the accessory pieces, we learn that
the disks of these genera and others could be added are composed
of a centro-dorsal, a first ring of 5 radialia, and a second one with 5
radialia and 10 basal supramarginaiia. This, then, is the same struc-
ture postulated for the hypothetic phylembryo of Asteroidea and
deduced through reduction of ossicles, as gleaned from a partial
ontogenesis of Hudsonaster.
Axillary ossicles. In the most primitive asterid, Hudsonaster, one
sees in each axilla a single large plate, on either side of which adjoin
the 10 columns of inframarginals. The same arrangement obtains
in Siluraster, Palseaster, and Australaster (here the genus occurs in the
Permo-Carboniferous, and the axillaries are very large contrasting
with the smaller ones of the Ordovicic Hudsonaster). In no other
Paleozoic genera does this same development hold, though the single
50 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
axillaries occur in many other forms where they are, however, pushed
orally into the skeleton of the interbrachial areas. Single, not double,
axillaries, therefore, make the simplest and smallest type of inter-
brachial areas. Further complexity and enlargement of these areas
are produced by crowding into them orally, first the axillaries and
then one or more pairs of proximal inframarginals.
In Hudsonaster narrawayi one sees tiny single plates proximal to
each axillary. These are clearly of no marked morphologic value and
are to be interpreted as accessory plates. Similar single ossicles are
also seen in genera with large interbrachial areas, as in Promopalseaster.
They are regarded as space-fillers and without ancestral significance.
Interbrachial areas. The simplest type of interbrachial area is that
just described. Increase in size and complexity of interbrachial struc-
ture is a natural result from an enlarged body cavity. The first change
in this direction is seen in Mesopaldeaster, Spaniaster, Miomaster, and
Devonaster, where the axillaries are crowded orally and surrounded
distally by a pair of adjoining basal inframarginals, and proximally by
a variable number of adambulacrals. The same development is seen
but hi somewhat modified form in the cryptozonian genera Urasterella
(U. ulrichi and U. girvanensis) and CaUiasterella. Here the axillaries
are present only in the young, but may be retained to maturity; in
either case they may or may not be margined distally by basal infra-
marginals. As the inframarginals in these forms are vanishing (pri-
mitive Cryptozonia), we should expect to find related genera devoid
of even the axillaries, and such is the case in Stenaster and Tetraster.
The next step in the enlargement of the interbrachial areas consists
of the inward crowding of pairs of basal inframarginals, one or more
at a time. In Trimeraster these areas have the single axillaries and a
pair of basal inframarginals, with the second pair of proximal infra-
marginals making the margin of the axils. In Xenaster the first and
second pairs of proximal inframarginals are enclosed, while in Pro-
mopalseaster there are from one to three pairs in each interbrachial
area. As these areas enlarge, it follows that more and more of the
adambulacrals must also take part in the expansion. These ossicles
are therefore called interbrachial adambulacrals. Further enlarge-
ment, but not to any marked extent, takes place through the insertion
of a variable number of accessory disk interbrachials. This is best
seen in Trimeraster and Xenaster.
Abnormal enlargement, but to a small extent, of the interbrachial
areas takes place through oral crowding entirely of adambulacrals in
pairs and through the increase in size of some of these ossicles. This
type of development is known only in Anorthaster.
It has been shown how enlargement of the interbrachial areas comes
about through oral crowding of the axillaries, proximal inframargi-
nals and adambulacrals pieces of the primary skeleton but the
REVISION OP PALEOZOIC STELLEROIDEA. 51
greatest expansion is brought about by the introduction of more and
more accessory disk pieces between the adambulacrals and inframar-
ginals. The beginning of this development of the secondary skeleton
is seen in Petr aster and is variously progressive in the different species.
They finally crowd more and more distally into the rays, as may be
seen in Uranaster and Palseostella. Among the large-disked Crypto-
zonia, as Schuchertia and the Palasteriscidse, it is the ambital and
abactinal accessory pieces that go to enlarge the arcs and body cavity.
CLASSIFICATION.
A good history of the various attempts at classifying the fossil
starfishes is given by Spencer (1914: 39-52). How recent starfishes
are classified may be learned from Verrill (1914:24), and it will be
seen that his principles are not always applicable to fossil forms be-
cause of the loss of parts through fossilization. In subsequent pages
the author will give his groupings up to superfamilies, but the time
is not yet at hand for an ordinal classification applicable alike to
fossil and recent starfishes.
Catalogue of Paleozoic Stelleroidea. A catalogue of all Paleozoic
starfishes was published by the writer in April, 1914, under the title
"Fossilium Catalogus, Anirnalia, part 3, Stelleroidea palseozoica," pp.
53 (Junk, Berlin). It gives the entire bibliography of the genera and
species up to that date, and an outline of the classification adopted.
The actual date of several new genera is of the work above cited,
although they are also printed in the present book as new. This
Bulletin is there cited as of 1914 and is now to be corrected to 1915.
The above-mentioned catalogue will always be useful for ready refer-
ence to the literature, although the larger volume repeats all of the
essential references, corrected to November, 1914.
DESCRIPTION OF GENERA AND SPECIES.
Class STELLEROIDEA.
Subclass ASTEROIDEA.
PHANEROZONIA Sladen.
Asteroidea nearly always with large marginal plates. These consist
in the Mesozoic and later asterids Qf completely superposed inframar-
ginals and supramarginals; in the Paleozoic the rule is that the in-
framarginals alone border the animal, while the supramarginals are
always dorsal, generally alternate with the inframarginals, and are
placed farther inward though resting more or less upon the latter.
Ambulacrals always present and opposite in arrangement in post-
Paleozoic forms. In the oldest and most primitive Paleozoic forms
they are apt to be alternate throughout the rays, or only so distally
in the young parts. By far the greater number of Paleozoic forms,
however, have the opposite arrangement.
52 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Oral armature always with the adambulacral pieces most con-
spicuous.
This order is not known to appear before the Middle Ordovicic,
and in the Paleozoic contains the following families:
Hudsonasteridae. Xenasteridae.
Palaeasteridae. Neopalaeasteridae.
Promopalaeasteridae. Palasterinidae.
Mesopalaeasterinae. Lepidasteridae.
Promopalaeasterinae.
Anorthasterinae.
These seven families of Paleozoic asterids are all related to one
another and seemingly had their origin more or less directly in
Hudsonaster or a form very similar to it. Therefore the family
Hudsonasteridae is the primordial stock out of which all asterids
progressively developed. Hudsonaster is massively and simply
built, with the spines rudimentary and restricted to the adambula-
crals and inf ramarginals ; it is a small rigid asterid with few and
comparatively large ossicles, nearly all of which are of the pri-
mary skeleton. From it the evolutionary tendency was to get rid
of the rigidity and ponderosity through the comparative reduction
in size of the plates and the introduction of many smaller sec-
ondary pieces, along with an abundance of spines. With this ten-
dency once established, there arose several others: (1) elongation
of rays, with greater flexibility and greater power of locomotion
through the indefinite duplication of pairs of podia; (2) increase of
body cavity in three different ways. This took place through the
development of interbrachial areas (a) by the inward crowding of
the single interbrachial axillaries, followed by more and more pairs
of proximal inf ramarginals; (&) through the insertion of an indefinite
number of accessory ossicles between the adambulacrals and the
inf ramarginals, forcing the latter more and more outward; and
(c) through the introduction of supernumerary rays. The latter
development is probably abnormal, in that the tendency arose in
the later larval life as is explained elsewhere. (See p. 207.) With
these changes of the actinal side naturally must arise compensating
growths of the abactinal surface and accordingly here is seen the
introduction of accessory ossicles in various places.
Starting with Hudsonaster, there is one phyletic line developed
through the elongation of rays, flexibility, and the increase in size
of the oral region by the (a) method from the primitive Hudson-
asteridae into the derived Palaeasteridae, Promopalaeasteridae, Xen-
asteridae, and Neopalaeasteridae. This is the most satisfactory line
of evolution, resulting in the greatest variety of genera. Again
with Hudsonaster as the radicle, another phyletic line is established
through the increase of the body cavity by the (6) method, giving
REVISION OF PALEOZOIC STELLEROIDEA. 53
rise to the Palasterinidse. This line of change is not at all so pro-
ductive of genera as the former. Finally, a third phyletic line is
established through larval adaptation, causing a sixth ray to develop
and thereafter pairs of rays, resulting in the multi-rayed asterids of
the family Lepidasteridae.
These phyletic lines may be expressed taxonomically as follows:
Superfamily Promopalseasteracea, new.
Includes families Hudsonasteridae, Palseasteridse, Promopalseasteridae, and
Xenasteridae. The Neopalseasteridae developed out of the Promopalaaasteridse.
Superfamily Palasterinacea, new.
Includes family Palasterinidse.
Superfamily Lepidasteracea, new.
Includes family Lepidasteridse.
HUDSONASTERID.E, new family.
Most primitive known Phanerozonia.
Small, five-rayed, heavily plated asterids, with narrow ambulacral
furrows and slightly alternate ambulacralia. The incipient inter-
brachial arcs are occupied by single axillary marginal plates. Abac-
tin ally the rays have five columns of ossicles, the radials in the center
being bounded on either side by the supramarginals and inframarginals;
the latter margin the rays and the ossicles of adjoining columns alter-
nate with one another. There are no accessory ray plates of any
kind. The disk has a prominent central disk piece separated by a
small number of accessory disk plates from a ring of five large basal
radials and five large basal interradial ossicles. Spines rudimentary
and apparently restricted to the adambulacrals and inframarginals.
This family contains the genera Hudsonaster Stiirtz and Siluraster
Jaekel.
Genus HUDSONASTER Sturtz.
Plates 1-6.
Palseaster (part) of authors.
Hudsonaster STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
pp. 224, 225.
Protopalseaster HUDSON, Ottawa Naturalist, vol. 26, 1912, p. 25; vol. 27, 1913,
pp. 77-84. RAYMOND, Ottawa Naturalist, vol. 26, 1912, p. 105 (genoholo-
type, P. narrawayi). SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palaeontgr.
Soc. for 1913), 1914, p. 21.
The name is derived from the " Hudson River formation/ 7 an
Ordovicic series of strata of long endurance. If the stratigraphic
term is used correctly, then these rocks have as yet furnished no
starfishes of this genus; for this reason Hudsonaster is a misnomer
and has no significance, but under the rules of nomenclature it must
be accepted.
Generic characters. Disk small and without actual interbrachial
arcs. Rays five, short, tapering rapidly.
54 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Abactinal area of rays occupied by three columns of large, thick,
more or less tumid or carinate, closely adjoining, tuberculate plates;
more or less of the inframarginal plates are also to be seen, making
five visible columns of plates on each ray abactinally. There is,
therefore, a single radial column bounded on each side by two columns
of supramarginal and two of inframarginal plates. The latter
columns are the marginal plates both actinally and abactinally, but
on the dorsal side are overlapped by the supramarginals. In the
center of the disk there is a small but very tumid plate (central
disk plate) around which is a nearly complete circle of smaller acces-
sory plates and an open space which may represent an anal opening.
Outside of this first circle of accessory plates there is a ring of 10
very large plates that in the geologically later species are stellate
on the surface, 5 of which are radial and 5 interradial in position.
The radial plates are the basal radial plates of the 5 medial columns,
while the 5 interradial plates give rise to the 10 larger and conspicu-
ous supramarginal ossicles. The basal supramarginals adjoin one
another in the axillary areas, and in the geologically later forms
these also have stellate surfaces. Between each of the 5 inter-
radial plates there is inserted a single plate, making a second but
disconnected ring of accessory plates.
Madreporite abactinal, more or less large, adjoining one of the
large interradial plates and between the basal plates of two supra-
marginal columns of ossicles of adjoining rays.
No accessory ambital or abactinal plates.
Spines appear to be absent everywhere except on the inframarginal
and adambulacral ossicles. Otherwise the surface is granular or
tuberculate.
Inframarginal plates well-developed, tuberculate, and common
to both the actinal and abactinal areas. These proximally adjoin the
five isolated and large axillary marginals, each one of which occupies
.an axil of the rays. Each plate, except the axillary ones, bears on
its outer edges one to three apparently smooth, small spines.
Axillary plates quite conspicuous, marginal, and one in each
actinal axil. In at least one of the species there is a single tiny acces-
sory piece orad to each axillary (H. narrawayi), and it is only seen
on the inside of the skeleton.
Adambulacral plates numerous, one or two to each inframarginal
plate and two on each side of each axillary marginal plate. All of
these pieces bear at least two spines, one on the inner and another
on the outer edges.
Oral armature consisting of at least five pairs of pointed and elon-
gate basal adambulacral plates; each bears spines like the other
ambulacrals. In rare cases issuing from beneath these ossicles may
be seen five single stout spines or plates (tori) that are connected
REVISION OF PALEOZOIC STELLEKOIDEA. 55
with the pairs of ambulacral armature pieces. In H. narrawayi above
or dorsal to the pair of large oral pieces there is another but smaller
and modified pair of ambulacral ossicles clearly also a part of the oral
armature.
Ambulacral grooves originally narrow and regularly tapering. Am-
bulacral plates rectangular, practically opposite or very slightly
alternating, one to each adambulacral plate. The ambulacralia
of each column may be so closely adjoining as to leave almost no
space for the podial openings, or there may be laterally between
adjoining ossicles very large openings.
Genoholotype. The genotype selected by Sttirtz is Pdlasterina
rugosa Billings. The above generic definition is, however, largely
based upon Palxaster incomptus Meek and P. matutinus Hall.
Distribution. Restricted to the middle and upper portion of the
Ordovicic. The following are the known species :
H. matutinus (Hall). Trenton.
H. narrawayi (Hudson). Black River.
H. milleri, new species. Trenton.
H. incomptus (Meek). Maysvillian and Richmondian series.
H. rugosus (Billings). Lower Richmondian, Charleton forma-
tion.
H. ~batheri, new species. Upper Ordovicic, Girvan, Scotland.
Remarks. Sttirtz is unfortunate in selecting as the genotype of
Hudsonaster, Palasterina rugosa Billings, a very poorly known starfish
from the Richmondian of Anticosti Island. Therefore his original
definition of Hudsonaster is not only very imperfect, but the specimen
is wrongly interpreted. His characterization of the genus is as
follows :
Of Palasterina rugosa only the abactinal side is known. On the disk occur orna-
mented, deeply crenulate, star-shaped plates. Four [the specimen is distorted and
has forced into view one of the inframarginal columns, while the one on the other side
of the ray is shoved out of sight] regular and decided columns mark the rays, and of
these the outer rows are stronger than the inner columns. The outer columns consist
of unmistakable marginal plates [one is inframarginal, the other is a supramarginal
column]. According to Billings there are even traces of the actinal marginals to be
seen from the abactinal side. Spines absent [the usual spines of this genus are also
present here]. The arms terminate sharply in points.
Sttirtz is correct in saying that this species can not be included in
Palasterina and also that the genus has come to embrace a variety
of unrelated genera. However, he does not say what are the
actual generic characters of Hudsonaster, but as P. rugosa is not a
Palasterina he concludes that as "new generic names are required
I propose Hudsonaster for Palasterina rugosa Billings. " This
species is undoubtedly congeneric with Palseaster incomptus, a form
fully described in this work both from the actinal and abactinal sides,
and is therefore taken as the genotype for the above generic defini-
tion.
56 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
The adopted genotype of Protopalseaster (P. narrawayi) is described
at length and in detail by Hudson, but, thinking the specimen
showed the actinal side, he described the ambulacrals as epineural
plates roofing over the ambulacral furrow, a character that he of
course recognized as wholly anomalous for starfishes. Raymond
later on showed that the actinal side of this specimen lies buried in
the rock and that the entire abactinal plates are now gone, so that
the view is of the actinal plates from their inner side, i. e., an inter-
nal view of the ventral skeleton (later Hudson combats this view).
Under these circumstances nearly all of Hudson's generalizations
are fallacious and especially his statement that "we are dealing
with an unrecognized and very archaic morphological type which
links the Edrioasteroidea with the Stelleroidea " (1912 : 24) . A proper
interpretation of the specimen shows it to be a primitive but other-
wise a normal early Paleozoic starfish. Even though Hudson's
genus is far better determined generically than that of Stiirtz, it
must give way to the older name Hudsonaster. No one regrets this
more than the writer, because the generic name Protopalseaster ex-
presses the phyletic relationship of these primitive starfishes and is a
name that he has had in manuscript for the past 15 years.
Hudsonaster has its nearest known relationship in Palseaster, but
differs at once from the latter in that the rays have abactinally five
columns of plates, while Palseaster has but four. In other words,
Palseaster has no radial columns of ray plates, but their place is oc-
cupied by numerous small accessory pieces. The abactinal disk
of Palseaster is also quite different hi having numerous small acces-
sory ossicles, while in Hudsonaster this area is devoid of these pieces
and instead there are disk plates that are larger, far less numerous,
and with a definite arrangement.
The plate arrangement of Hudsonaster is simple and primitive
in that it distinctly retains to maturity much of the larval plate
structure of recent species and does not develop accessory ambital
or accessory abactinal plates as do its descendants Mesopalseaster
and particularly Promopalseaster. In Hudsonaster the axils are
occupied by single, large, axillary marginals, against which rest
the columns of inframarginal plates of adjoining rays. In the other
two mentioned genera, interbrachial arcs begin to develop by crowd-
ing orally the axillary marginals, when, as in Mesopalseaster , the
proximal plates of the inframarghial series of adjoining rays abut
against each other. In Promopalseaster this crowding of the infra-
marginal plates into the interbrachial areas is continued, always a
pair of plates at a time passing orally. For other remarks on the
development and distinction between these genera, see Mesopalse-
aster and Promopalseaster.
REVISION OF PALEOZOIC STELLEKOIDEA. 57
In the Lower Carboniferous genus Neopalseaster the primitive
disk structure of Hudsonaster is also retained in its essential fea-
tures. Otherwise these genera are easily distinguished, as is pointed
out in the remarks on Neopalseaster.
HUDSONASTER MATUTINUS (Hall).
Plate 2, fig. 2; plate 3, fig. 2; plate 5, figs. 1, 2.
Asterias matutina HALL, Nat. Hist. N. Y., Pal., vol. 1, 1847, p. 91, pi. 29, figs.
5o, 56.
Palxaster matutinus HALL, Twentieth Eep. N. Y. State Cab. Nat. Hist., 1868,
p. 283, pi. 9, fig. 2; rev. ed., 1868=1870, p. 325, pi. 9, fig. 2.
Petraster rigidus (part) BILLINGS, Geol. Surv. Canada, Can. Org. Bern., dec. 3,
1858, pi. 10, fig. 36 (not fig. 3a). HALL, Twentieth Rep. N. Y. State Cab.
Nat. Hist.,1868, p. 294; rev. ed., 1868=1870, p. 337.
Original description. "Body small, with five radiating arms,
arms elongated (length twice and a half the width of the body),
terete, acute, composed of three rows of plates [abactinally], which
join above by their lateral margins. Beneath each upper lateral
row of plates there is an inferior lateral range [inframarginal] vis-
ible, leaving the sulcus beneath the middle row, which is often
depressed. The dorsal places are somewhat hexagonal; those of the
arms quadrangular; surface punctate or granulate, perhaps from the
removal of the spines covering the surface.
"The specimen is considerably crushed, and two of the arms
broken off at the base; the madreporiform tubercle upon the back
is not visible in our specimen. From its condition, the structure
can not be entirely made out, but it is sufficiently clear to enable
any one to recognize the species."
Emended description. Specimen of the usual size (No. 3, Mus.
Comp. Zool.) measures: R, = 14 mm., r = 3.5 mm., R = 4r. The larg-
est specimen (Ottawa, Billings collection): 11 = 16 mm., r = 5 mm.,
R = 3.1r. The smallest New York specimen (No. 26, Mus. Comp.
Zool.): R = 7 mm., r = 2.5 mm., R = 2.8r. Width of rays at base
from 2.5 mm. to 5.5 mm.
Rays short, tapering rapidly, convex abactinally and flat actinally.
Disk comparatively large, probably convex abactinally during life.
Abactinal area of rays consisting of 5 ranges of large, thick,
and tumid granular plates. The 3 medial ranges, when not
distorted by pressure, conspicuous, with the 11 to 12 plates of
each column closely adjoining and considerably elevated above
the inframarginal ranges. The proximal plates of each supra-
marginal range lie against the large interradial plates and are
more or less superposed on the large axillary marginals. In the
center of the disk there is a distinct highly convex plate, around
which is arranged a nearly complete ring of seven smaller pieces
58 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and a vacant space which may be an anal opening. Five other
small quadrangular plates are intercalated, one proximally to each
of the large radial basal plates. Outside of these 12 intercalary
plates there is a ring of 10 quite large and conspicuous plates, 5 of
which are radial and 5 interradial in position. The radial plates are
largest and are the basal pieces of 5 medial or radial columns, while
against the 5 interradial plates rest the 10 inframarginal columns.
Madreporite abactinal, quite small, situated immediately above a
large interradial plate and between two basal supramarginal plates.
Its position in relation to the anal opening is apparently as in P.
incomptus.
Inframarginal plates in mature examples about 14 to a col-
umn, and when not displaced completely, inclosing the adarnbu-
lacral plates and margining the rays. The axils are occupied by a
very large, oval, axillary marginal plate. Each inframarginal bears
upon its outer surface one to three short, slender, pointed spines,
which are finely striated longitudinally.
Adambulacral plates small, subquadangular in outline and about
19 in a column in specimens having 13 inframarginals. Each plate
bears on both the outer and inner edges a single spine and is like
those of the inframarginal ranges.
The oral armature consists of 5 pairs of elongate adambulacrals,
each pair being situated directly orad of the large axillary marginal.
Ambulacral plates I -shaped, practically opposite or but very
slightly alternating, one to each adambulacral plate. In the ex-
panded medial portion the ambulacralia are slightly superposed one
above the other proximally, while the lateral portion of each plate
is very slender, leaving between adjoining pieces comparatively large
podial openings.
Locality and formation. From the Trenton limestone, as follows:
The type-specimen was found in the " shelly layers" at Trenton
Falls, New York (Amer. Mus. Nat. Hist., No. 199), and another
specimen was secured there by Mr. Rust (New York State collection).
Dr. Walcott collected a slab with three individuals in the Lower
Trenton at Rathbone Brook, near Newport, Herkimer County, New
York; and Mr. Taylor got two specimens at Deerfield, Oneida County,
in the same State (Mus. Comp. Zool., Nos. 26 and 3). Mr. W. R.
Billings found a specimen in the Trenton at Lachine, Quebec, and
another from Government House Bay, Ottawa.
Remarks. H. matutinus is intermediate in structure between the
older E. narrawayi and the younger H. incomptus. It is larger than
the former, has more plates in each of the columns, while the ambu-
lacrals are deeply cut out for the podia, which is not the case in H.
narrawayi. As H. matutinus is most closely related to H. incomptus
more detailed comparisons must be made between them. Actinally
KEVISIOISr OF PALEOZOIC STELLEEOIDEA. 59
the former differs in the wider ambulacral grooves. The difference
of greatest value, however, is found on the abactinal area of the
disk. In both species there is a central disk plate surrounded by a
ring of accessory plates. In H. matutinus there are eight plates in
this ring and in H. incomptus there are seven. Between all the basal
radials in the latter species there is an accessory plate, or these are
five in number, while in the former there are only two such plates.
Again, the proximal plates of the supramarginal columns in H.
incomptus are larger than any others of the abactinal area, while in
H. matutinus they are smaller than the basal radials and their form
and ornamentation are quite different. The madreporites are also
quite different. In general, H. incomptus is larger, stouter, the rays
broader and the disk has a greater number of accessory pieces. These
features are still more pronounced in H. rugosus (Billings). Viewing
these differences developmentally, the distinctions are slight, which
proves the three forms to be genetically related, but specifically the
differences are sufficiently marked to be regarded both morpholog-
ically and geologically as of specific value.
Apotype.Csii. No. 60603, U.S.JST.M.
HUDSONASTER NARRAWAYI (Hudson).
Plate 1, fig. 1; plate 2, fig. 1; plate 4, fig. 1.
Protopalseaster narrawayi HUDSON, Ottawa Naturalist, vol. 26, 1912, p. 25, pis.
1-3; vol. 27, 1913, pp. 77-84, pis. 8, 9. RAYMOND, Ottawa Naturalist, vol. 26,
1912, pp. 105-107. HUDSON, Bull. N. Y. State Mus., No. 164, 1913, pi. 5
SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914,
p. 21, fig. 19.
Description. Specimens small, an average individual from St.
Paul, Minnesota; measuring: R = 5 mm., r = 3 mm., R = 1.6r. Width
of ray at base 2.7 mm. The three good specimens show only the
actinal side while the fourth free siliceous one is too poorly preserved
to make out more than the form.
Inframarginal columns have from 6 to 9 plates, while the adambu-
lacrals and ambulacrals each have from 11 to 12. Ambulacral plates
large and solid, rectangular, with slightly rounded ends. As these
plates are not t- -shaped as in other species of this genus it may be
shown that this character is of generic value, in which case Proto-
pal&aster will have to be revived.
Inside of each of the large axillary plates lies a pair of oral armature
pieces that are in form very much like the adjoining adambulacrals.
From beneath these ossicles projects a stout and long, rounded, arma-
ture rod, the torus (a tooth or chisel), the 5 teeth meeting together
in the center of the mouth. These armature rods and more especially
the 10 pairs of ambulacral armature pieces are seldom seen in fossil
starfishes, and none shows it more clearly than the Yale specimen
60 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
of H. narrawayi. In the holotype of this species, here illustrated,
are seen additional pairs of oral armature ossicles. These are situated
directly above or dorsal to the pairs of large armature plates, and are
probably overlapping ambulacralia modified into the actinostomial
ring.
Locality and formation. In the Black Kiver formation, as follows:
The holotype was found by Mr. J. E. Narraway at City View, near
the Central Experimental Farm at Ottawa, Canada; it is now in his
private collection. An excellent specimen was found by Mr.
Townshend near Kirkfield, Ontario, and is now in the Peabody
Museum of Yale University. Many years ago Dr. Ulrich found a
fine specimen at St. Paul, Minnesota, and another good one in the
Lower (Glade) Lebanon limestone of the Stones River series at
Shelbyville, Tennessee. These two specimens are in the United
States National Museum (Nos. 60602, 60619). Mr. Moritz Fischer
secured a coarsely silicified individual of this genus at Curdsville,
Kentucky, and this is now in the Beecher collection in the Yale
Museum; it may, however, prove to be of another species, distin-
guished by its stouter appearance, wider disk, and smaller axillary
plates.
Remarks. Originally the writer included specimens of this form
under P. matutinus, regarding them as but young individuals of
Hall's species. However, as the specimens are all geologically older,
are always smaller and stouter, and with fewer plates in all of the
columns, Hudson's species is retained as valid. It is the ancestral
form and the most primitive species of Hudsonaster and hence the
most primitive Paleozoic starfish. It is primitive because of its
extreme simplicity of structure, in that the columns and plates are
reduced to the smallest number and all of them are closely adjoining
and without intercalary plates of any kind. Then the spines are
developed only on the actinal side and as protection for the podia.
For further information see generic remarks under Hudsonaster.
Cat. Nos. 60602, 60619, U. S. N. M.
HUDSONASTER MELLERI, new species.
Plate 4, fig. 2.
Description. Of this form there is at hand but a single poorly
preserved individual showing the actinal side. It is a more slender
species than H. matutinus, has a larger and more rectangular axillary
plate, and the proximal inframarginals have each about 13 to 15
plates and the adambulacrals from 18 to 20. No ambulacrals are
preserved. The specimen measures: R = about 12 mm., r = 3 mm.
Width of ray at base 4 mm., at mid-length 3 mm.
Locality and formation. Found by Prof. Arthur M. Miller, after
whom the species is named, in the Lower Lexington (Wilmore)
REVISION OF PALEOZOIC STELLEKOIDEA. 61
limestone of the Trenton, in Fayette County, Kentucky. The
holotype is in the collection of the State University of Kentucky at
Lexington.
HUDSONASTER INCOMPTUS (Meek).
Plate 6, figs. 1, 2.
"Goniaster" of AGASSIZ, GRAHAM, ANTHONY, and JAMES, Amer. Journ. Sci.,
ser. 2, vol. 1, 1846, p. 441 (nomen nudum).
Palseaster incomptus MEEK, Amer. Journ. Sci., ser. 3, vol. 4, 1872, p. 275; Geol.
Surv. Ohio, Pal., vol. 1, 1873, p. 64, pi. 4, figs. 5a, 5&.
Palseaster simplex MILLER and DYER, Journ. Cincinnati Soc. Nat. Hist., vol. 1,
1878, p. 29, pi. 1, fig. 6. MILLER, N. Amer. Geol. Pal., 1889, p. 266, fig. 380.
Palseaster darlcei MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 1, 1878, p. 102,
pi. 3, fig. 5.
Palseaster clarkana MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 3, 1880, p. 236.
Original description (P. incomptus). " Small; rays rather short, or
only about once and a half as long as their breadth at their inner ends,
and rapidly tapering to their outer extremities, which are somewhat
obtusely angular. Disk equaling in breadth the length of the rays.
Dorsal side of the rays composed each of three rows of pieces that are
wider than long, about nine in each row, and increase rather rapidly in
size inward to the margin of the disk, which is composed of smaller
pieces; a few very minute pieces apparently some times occur between the
rows on the dorsal side of the rays [none are apparent in our material].
Surface of the dorsal pieces a little granular, but apparently without
spines. Madreporiform piece rather small, a little oval, or almost
circular [or trapezoidal], nearly flat, and marked by fine, irregularly
interrupted, radiating striae. Ventral side unknown."
Emended description. B, = 11 mm., r = 6 mm., R = 1 .8r. Width of
ray at base 6 mm. Another larger specimen measures: R = 16 mm.,
r = 7 mm., R = 2.3r. Width of ray at base 8 mm. Smallest speci-
men: 11 = 4.5 mm., r = 2 mm.
Rays short, stout, tapering rapidly, convex abactinally and longi-
tudinally ridged, flat actinally. Disk large, probably very convex
abactinally during life.
Abactinal area of rays consisting of five columns of large, thick,
tumid, alternating, granulose plates. In the center of the disk there
is a distinct, highly convex plate around which is arranged a nearly
complete ring of seven smaller pieces and a comparatively large open
space which possibly represents an anal opening. This ring is fol-
lowed by another composed of five large interradial plates between
each of which laterally and somewhat centrally is intercalated a
smaller plate radial in position. Laterally and distally to each of
the five large interradial plates are five large radial plates which are
the basal pieces of the five radial columns. Against the basal plates
of the medial ranges there adjoin laterally two large stellate ossicles,
62 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
each of which is the proximal plate of a supramarginal column.
The rays are margined by the inframarginal plates. There are 10
plates in each radial and 9 in each supramarginal column.
Madreporite abactinal, situated interradially and dis tally against
the large interradial plate of the second ring, which is in front of the
anal opening, and between two of the proximal supramarginal plates.
It is trapezoidal in form, flat, and marked by very fine, dichotomous,
radial striae.
Inframarginal plates completely inclosing all other plates and com-
mon to both the abactinal and actinal surfaces. Usually there are
9 or 10 in a column, not counting the axillary plates. Spines are
present at least along the inner edges of the inframarginals.
Abactinally the axillary marginals are not prominent but actinally
they are very conspicuous. No spines appear to be connected with
these ossicles.
Adambulacral plates small, subquadrate, usually 2 to each
inframarginal or from 18 to 22 plates in each column. From 4 to
6 are situated around the inner edge of each axillary, of which the
2 central ones are largest, being one of the 5 pairs of plates
making the oral armature. The adambulacral plates originally bore
on their lateral and inner edges numerous, very short, thick spines.
Ambulacral plates nearly entirely concealed by the adambulacral
plates, apparently as numerous as the latter, and, so far as can be
determined, like those in P. matutinus Hall.
Locality and formation. In the lower, middle, and upper beds of
the Cincinnatic (Ordovicic). Ten specimens have been studied:
Four in the Harris collection in the United States National Museum
(No. 40882) from the Richmond formation (Waynesville division)
about Waynesville, Ohio; another from the same place and formation
in the University of Chicago Museum (Gurley collection, No. 10977);
one from the hill quarries in the Maysvillian series (Vaupel collection,
U. S. Nat. Mus., No. 60616) back of Cincinnati; another, said to be
from Cincinnati, in the University of Toronto (Walker collection,
No. 691 H. R.); the type-specimen, now in the Museum of Com-
parative Zoology, also from Cincinnati; one in the Ulrich collection
of the United States National Museum (No. 60617), found back of
Covington, Kentucky, in the Maysville formation, about 315 feet
above low-water mark in the Ohio River; and the type of Palxaster
simplex Miller, found near Raysville, Ohio (Faber collection, Uni-
versity of Chicago Museum, No. 8830). The holotype of P. darkana
Miller is in Professor Harper's collection at Cincinnati, while another
specimen is in the Ulrich collection of the National Museum (No.
60618).
Remarks. The writer has seen the type-specimen of Palseaster
simplex Miller, which agrees with the description and figure given
KEVISION OF PALEOZOIC STELLEEOIDEA. 63
by its author, and it is like other specimens in the Harris collection
known to be H. incomptus (Meek) . Of the latter, Meek only described
the abactinal area, while Miller's specimen shows the actinal side.
The four specimens in the Harris collection unmistakably affirm
that both forms are identical, and since Meek's name has priority
it is here accepted. *
For remarks as to the relationship of this species see H. matutinus.
Through the kindness of Prof. George W. Harper, principal of the
"Woodward High School, Cincinnati, Ohio, the writer was enabled to
study the type of Palseaster darkana Miller." The specimen is a
little abraded, but shows all the characters of H. incomptus except that
it is much smaller in size. The abactinal area on a cursory examina-
tion shows but three columns, but in several places can be seen pro-
jecting small parts of the inframarginal columns. The disk also shows
the same general arrangement and number of plates as in H. incomptus.
There is therefore nothing except the smaller number of plates in the
columns to distinguish P. darkana from H. incomptus. Bearing
in mind that the former is but one-third the size of the latter, it is
not remarkable that P. darkana should have but 6 or 7 plates
while H. incomptus has 9 to 10 plates in the same columns. The
entire difference is accounted for when one regards P. darkana as
the young of H. incomptus.
Another small but very poorly preserved specimen showing the
actinal area is in the Ulrich collection of the National Museum and
agrees with the type of P. darkana. Its measurements are:
11 = 4.5 mm., r = 2 mm., K = 2.2r.
The only ontogenetic character observable in these small speci-
mens of H. incomptus is that the columns have fewer ossicles, and the
same is true of the abactinal area of the disk. It has a central disk
plate, the first ring of small plates and the second ring of basal radial
and interradial plates. But no intercalated small plates radial in
position are present. The second ring of 10 plates and the central
plate are the most prominent. If one were theoretically to reduce
the size of the animal still more, the plates of the first ring would be
the first to go and the rays would be simultaneously very much
shortened, probably obsolete, resulting in a hemispheric nonstellate
starfish. The abactinal area seemingly will then have but a central
disk plate and 10 others, 5 radial and 5 interradial in position. On
the actinal side there will be but the 5 axillary marginal plates
separated from one another by the ambulacra! furrow and the
ambulacralia. This hypothetic animal therefore closely resembles
a very young Comatula without the stalk or column, and also the stage
of living asterids following the brachiolarial.
Cat. Nos. 40882, 60616, 60617, 60618 U.S.N.M.
50601 Bull. 8815 5
64 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
HUDSONASTER RUGOSUS (Billings).
Plate 3, fig. 1.
Palasterina rugosa BILLINGS, Geol. Surv. Canada, Rep. of Progress for 1853-1856,
1857, p. 291; Geol. Surv. Canada, Can. Org. Remains, dec. 3, 1858, p. 77, pi.
9, figs. 2a-c.
Palasterina rugosa WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1
(Paleeontogr. Soc. for 1861), 1862, p. 27.
Hudsonaster rugosa STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
pp. 218, 224, 225.
Original description. "Two inches [not over 47 mm.] in diameter,
rays five, acute at their apices and rapidly enlarging to a breadth
of four lines [about 11 mm.] at the disk, which is eight lines [about
17 mm.] in width. The specimen shows the upper [or abactinal]
side of the fossil only; some of the plates are absent from the center
of the disk, but those which remain are very prominent in their
centers, and roughly ornamented by four or five deep crenulations
or furrows from near the center to the edges, producing a star-like
appearance resembling a half-worn plate of Glyptocrinus decadac-
tylus; their diameter is from one to two lines [largest diameter 4 mm.].
"The rays are composed (at least the backs and sides of them)
of four rows of plates [due to distortion, a careful analysis shows
five columns] which are so very, prominent that they appear to be
almost globular, and even pointed in their centers, the central
[radial] rows are the smallest [as in H. incomptus]', the first four
plates of the outer [supramarginal] row occupy three lines in length,
and of the inner [radial] rows nearly as many. Toward the point
of the arm all diminish rapidly in size.
"Beneath the outer [or supramarginal] rows two others can be
seen [the inframarginal], which are probably the outer marginal plates
of the under [or actinal side], corresponding to those of Petrasier
rigidus."
Locality and formation. Richmondian series, Charleton formation,
at Charleton Point, Anticosti Island. Two specimens collected by
J. Richardson are now in the Victoria Memorial Museum at Ottawa,
Canada (No. 1999).
Remarks. The same description was republished in 1858 with
the addition of three figures. These, and a direct examination of
the two type-specimens, show that this species clearly belongs to
Hudsonaster and that it is closely related to H. incomptus. The
smaller free specimen preserves a portion of the disk and parts of
two rays and shows clearly the presence of a single large axillary
plate. H. rugosus, however, attained a considerably larger growth,
has a greater number of plates in each column and these are described
as "almost globular," while in H. incomptus the abactinal plates are
very convex but not globular. Then the basal supramarginal and
REVISION OF PALEOZOIC STELLEROIDEA. 65
radial plates are much more decidedly ornate. Each of the supra-
and inframarginal columns has 14 plates against 9 in H. incomptus,
and of adambulacrals there are not less than 24 in a column against
18 to 22 in the same form. All of the ossicles are as strongly papillose
but not more so than in H. incomptus. The detail of the disk is not
determinable.
HUDSONASTER BATHERI, new species.
Plate 3, fig. 3.
Tetraster wyville-thomsoni NICHOLSON and ETHERIDGE (part), Mon. Silurian
Foss. Girvan Dist., Ayrshire, fasc. 3, 1880, p. 324, pi. 21, figs. 1, 2 (not the
other figs.).
A small Hudsonaster. R = 6 mm., r = 2.7 mm. The largest
specimen: R = 8 mm.
Actinal side only known. Inframarginal columns the largest,
with 8 or 9 distinctly tuberculate plates. Inside of these are
the columns of narrower and slightly shorter adambulacrals that
lie somewhat below the level of the inframarginals ; there are
10 or 11 of these plates bounding the ambulacral grooves, and the
pieces of adjacent columns are opposite or nearly opposite one
another. The ambulacral grooves are very narrow and deep and
no ambulacralia are discernible.
Formation and locality. Dr. F. A. Bather made wax squeezes
for the writer from three natural molds in the collection of Mrs.
Robert Gray, Edinburgh; the originals are from the Upper Ordovicic
of Thraive, Girvan District, Scotland. The holotype is the specimen
illustrated as above cited.
Remarks. Nicholson and Etheridge confused at least one of these
specimens with their Tetraster wyville-thomsoni , a species of totally
different construction, a cryptozonian, described elsewhere in this
work. Actinally the new species is in harmony with Hudsonaster
and although the abactinal side is unknown, it is thought that it
will be found to be like that in H. matutinus.
Cat. No. 60601, U.S.N.M.
Genus SILURASTER Jaekel.
Text fig. 6.
Siluraster JAEKEL, Zeits. geol. Ges., vol. 55, 1903, Protokol, p. 13 (106), figs. 2,
3, on p. 108. SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palaeontgr. Soc. for
1913), 1914, p. 18.
Description. "A typical asterid with opposite ambulacralia and
with well-developed marginalia, therefore a true form of Phanero-
zonia. The strongly folded madreporite lies dorsally in an inter-
radius."
66
BULLETIN
UNITED STATES NATIONAL MUSEUM.
l2
From the two excellent illustrations the following is made out:
Inframarginals conspicuous and as large as the adambulacrals.
Both columns finely tuberculate, with the ossicles of each alternating
with one another. Proximally the inframarginals of adjacent rays
rest on a somewhat larger axillary plate. The interbrachial areas
are therefore made up of single axillary ossicles (Jaekel's marginal
oral) and in front of these lies a pair of pointed adambulacrals, the
conspicuous pieces of the oral armature. These are perforated for
the continuation of the ring canal. The podial openings as usual
lie in the corners of four adjacent adambulacrals and ambulacrals,
but there is a de-
cided tendency to
restrict them to
only one of the am-
bulacral corners.
Abactinal area
unknown.
Remarks. This
genus appears to be
closely related to
Hudsonaster, but
FIG. 6.-SILT7RASTER PERFECTUS (JAEKEL'S FIGURE 3). Oj TO Og, AMBU- , -1 o, -7 ,
LACRALIA OF ONE SIDE; ABOVE THESE ddi TO ddg, THE ADAMBULACRALIA, l l6 ' 1
adi AS MOUTH CORNER PIECE; dfg, PODIAL GROOVES; dpff, PODIAL OPEN- DlOrC fully WOrked
INGS; m, INFRAMARGINALIA; mo, THE SINGLE AXILLARY INFRAMARGINAL; , , n
RC, RING-CANAL IN MOUTH CORNER PLATES. Ut lt} ^ ^maUl aS
an obscure genus.
Genoholotype and only species. S. perfectus Jaekel. Ordovicic
(D4), Zahorzan, Bohemia.
Family PAL^ASTERID^E Gregory (emend.).
Palaeasteridae (part) MILLER, N. Amer. Geol. Pal., 1889, p. 216. GREGORY, Geol
Mag., dec. 4, vol. 6, 1899, p. 344 (contains Palseasterinae, Xenasterinse, and
Lindstromasterinse); Treat. Zool., vol. 3, Echinoderma, 1900, p. 250.
Palsecisterinas SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 244.
Palxgoniasteridx (part) STURTZ, Palaeontographica, vol. 36, 1890, p. 246 (contains
Aspidosomella and Pal&aster).
Primitive, derived, five-rayed Phanerozonia with the ambulacrals
slightly alternate or opposite. Interbrachial arcs incipient or some-
what enlarged, and occupied by single axillary marginal plates. No
accessory interbrachial or axillary interbrachial plates present, but
there are developed accessory abactinal ray ossicles. Actinal plates
consisting of ambulacrals, adambulacrals, and inframarginals. Mad-
reporite abactinal.
Contains :
Palseaster Hall.
Australaster, new genus.
REVISION OF PALEOZOIC STELLEROIDEA. 67
Genus PALSEASTER Hall (emend.).
Plate 7, figs. 1-4.
Palseaster HALL, Nat. Hist. N. Y., Pal., vol. 2, 1852, p. 247; Twentieth Rep. N. Y.
State Cab. Nat. Hist., 1868, p. 282; rev. ed., 1868=1870, p. 324. ZITTEL,
Handb. Pal., vol. 1, 1879, p. 452.
Palseaster NICHOLSON and ETHERIDGE (part), Mon. Silurian Foss. Girvan Dist.,
Ayrshire, fasc. 3, 1880, p. 319.
(Not Palseaster SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 323= Uras-
terella.)
(Not Palseaster of most AUTHORS.)
Generic characters. Disk small and without distinct interbrachial
arcs. Rays five and slender.
Abactinal area of disk centrally occupied by a large plate, then a
ring of 5 separated, somewhat smaller, radially situated ossicles,
and inside the axillary pieces there are 5 subtriangular interradial
plates, against which lie 10 very large and conspicuous proximal
supramarginal plates. Between the central, radial, and interradial
plates are numerous small accessory pieces which separate the pri-
mary ossicles one from another. These accessory plates also continue
to crowd from the disk into the rays between the supramarginals,
and thus take the place of the radial columns, but are absent in the
distal third of the rays. Each ray has therefore 4 conspicuous
columns of plates, the 2 supramarginals situated inwardly of the
2 marginal inframarginals. The former columns terminate in 10
very large proximal plates situated in the axils of the disk, while the
inframarginals rest against the 5 very large actinal axillary margi-
nal pieces. All of the large plates are centrally smooth, bordered by a
pustulose area.
Madreporite abactinal, of medium size, circular in outline, and
radially striate. It lies on top of two proximal supramarginal plates
and the adjoining interradial plate.
Inframarginal plates thick and large, about as many in a column
(13) as in those of the supramarginals (13 or 14). The latter do
not completely overlie the inframarginals, since these bound the
rays both actinally and abactinally. The inframarginal plates, like
the plates of the abactinal area, have each a large, smooth, central
space encircled by a pustulose area. From their inner sides articu-
late short, thick, blunt spines, of which there are apparently not
more than 2 to each inframarginal plate.
Axillary marginal plates large, hexagonal, one in each axil and
common to both sides of the disk axils.
Adambulacral plates small, tumid, alternating in the main with
the inframarginal plates throughout the distal two-thirds of the rays,
but toward the mouth there are a few more adambulacrals than in-
framarginals. (In the type-specimen there are 13 of the latter in a
column to 16 of the former.) Oral armature consists of at least 5
pairs of drawn out and pointed basal adambulacrals. Along the inner
68 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
side of each adambulacral plate there is a linear brush of from 5
to 7 short, slender spines.
Ambulacral grooves very narrow, tapering but very slowly. Am-
bulacral plates obscured by the rock but apparently very short, one
to each adambulacral and placed opposite one another.
GenoJiolotype (type by monotypy) and only species. P. niagarensis
Hall. Siluric (Rochester shale).
Remarks. To this genus has been erroneously referred a multitude
of Paleozoic starfishes. It is to the starfishes what Terebratula and
Rhynchonella used to be among the brachiopods a general dumping
ground for Paleozoic forms. This lack of generic conception regarding
Palseaster lies somewhat in the fact that Hall's original diagnosis is
very meagre and that he did not describe nor illustrate the abactinal
surface, probably the most important area amongst Paleozoic aster-
ids for specific and generic differentiation. Of the twenty-four Amer-
ican species listed under Palseaster by S. A. Miller 1 only one now
appears to belong there, the genotype. All others are here referred
to Hudsonaster, Urasterella, Mesopalseaster, Promopalseaster, Anorth-
aster, Devonaster, and Neopalseaster.
No new material of this genus accessible to the writer appears
to have been found since Colonel Jewett discovered the original
two specimens of Palseaster niagarensis. The writer therefore had to
make the most of this material, and to determine the structure of
the abactinal area he worked away much of the blue shale from the
back of the one good specimen. The finer detailed structure was then
revealed by cleaning with caustic potash.
In 1858 Billings proposed the genus Stenaster 2 and writes- that
"as it has been suggested that the two species hereinafter described
should be referred to Palseaster, I give the following figure of the genus
in order to show the difference." This figure is a good reproduction
of Hall's figure 27. Billings then points out that "if the large plates
which border the grooves in Palseaster be adambulacral, then there
are only five oral plates, whereas in Stenaster there are ten. But if
they be not adambulacral but marginal plates, then Palseaster must
have both marginal and adambulacral [the correct view], while
Stenaster has only the latter."
Hall's generic description is very meager and his figures give
the impression that the rays bear but a single column of marginal
spiniferous plates. With this evidence one can understand why
Billings was in doubt as to whether these plates are inframarginals
or adambulacrals and therefore the uncertainty as to the nature of
the five large axillary marginal plates. If, however, he had read
Hall's accurate specific description, Billings would have seen that an
important discrepancy -existed between the description and figures.
1 North American Geology and Palaeontology, 1889, p. 265.
2 Geol. Surv. Canada, Canadian Organic Remains, dec. 3, 1858, pp. 77, 78.
REVISION OF PALEOZOIC STELLEROIDEA. 69
Hall writes: " Lower side of arms showing two ranges of plates on
each side of the avenue, the outer range composed of strong hex-
agonal plates, with an inner range of smaller ones alternating, the latter
usually covered by tufts of spines." The type-specimen clearly
shows columns of adambulacral and inframarginals bordering the
ambulacral grooves and that in each axil there is a single large axillary
marginal plate. This matter need not here be followed further than
to add that Stenaster originally included two generic types, both of
which are widely different from Palseaster. On the other hand, Hall 1
is in error in regarding all of Stenaster as synonymous with Uraster-
ella McCoy, a name never properly defined and finally abandoned by
its author. Billings originally referred to Stenaster, S. salteri, and
S. pulcTiellus. The latter is certainly congeneric with the type of
Urasterella as illustrated by Salter, but the former is quite different,
as is pointed out elsewhere in the remarks on Stenaster, which is a
good genus.
Palseaster in many respects reminds one of Hudsonaster, and the
actinal generic characters may be regarded as alike in both. The
abactinal area is also similar, but in Hudsonaster there are five columns
with decidedly fewer plates, while in Palseaster there are no radial
columns. The former genus is the more primitive one and the
progression in differentiation toward Palseaster appears to be as fol-
lows: In Hudsonaster, the rays throughout are composed of five
columns of plates, two inframarginals, two supramarginals, and one
radial. In Palseaster, there are but four columns in the outer third
of the rays, two infra- and two supramarginals, but in the proximal
two-thirds where the radials are also absent, their place is occupied
by numerous small, irregular accessory plates. This introduction of
numerous supplementary plates also takes place to a far greater
extent on the disk, where they are crowded in between the larger
central and the five basal radial plates.
The primitive disk structure of Palseaster is in large part again
repeated in Neopalseaster, but otherwise the two genera are different.
PALSEASTER NIAGARENSIS Hall.
Plate 7, figs. 1^.
Palseaster niagarensis HALL, Nat. Hist. N. Y., Pal., vol. 2, 1852, p. 247, pi.
51, figs. 21-23 (not p. 352, pi. 85, figs. 8-10, possibly an undescribed
species). BILLINGS, Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858, p.
78, fig. 1. HALL, Twentieth Hep. N. Y. State Cab. Nat. Hist., 1868, p. 282;
rev. ed., 1868=1870, p. 324. QUENSTEDT, Petrefactenkunde Deutschlands,
vol. 4, 1876, p. 73, pi. 92, figs. 32, 33. HUDSON, Bull. N. Y. State Mus.,
No. 164, 1913, pis. 9, 10, 12, 13.
Original description. "Body stellate; disk small; arms short, terete
with a deep avenue on the lower side, which is margined by strong
short spines; centre of plates (in the fossil) nearly smooth, margins
i Twentieth Rep. N. Y. State Cab. Nat. Hist., rev. ed., 1808=1870, p. 325.
70 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
strongly granulate; lower side of arms showing two ranges of plates
on each side of the avenue, the outer range composed of strong
hexagonal plates, with an inner range of smaller ones alternating,
the latter usually covered by tufts of spines; a large pentagonal plate
inserted at the base of the arms, on the lower side.
"In this species, the arms are about twice as long as the width of
the disk. There are about 15 or 16 plates in each range from the
base to the apex of the arms. The range of the plates margining
the avenue are usually not visible, the whole being covered by the
short spines, which also partially fill the avenue. "
In 1870 Hall added the following important observations:
"I have distinctly recognized the two ranges, marginal and
adambulacral plates; but the inner ones are not shown in the figure
as they should have been, while the large plate at the axil of the ray
(though the adjacent small oral plates of the inner range are not
seen) is evidently part of an incomplete series, and clearly belongs
to the marginal range.
"The genus Palseaster has two ranges of plates on each side of the
ambulacral groove; marginal and adambulacral plates on the lower
side, besides ambulacral or poral plates. The upper or dorsal side
has three or more ranges of plates."
Emended description. Measurements: R=1S mm., r=5 mm,
Width of ray at base 5 mm.
Rays long, slender, tapering slowly; abactinally strongly convex
and longitudinally ridged; flat actinally. Disk comparatively large,
strongly convex and probably abruptly elevated above the rays
abactinally. There are no interbrachial arcs, but the five axillary
marginal plates occupy the axils and are also partially visible
abactinally.
Abactinal area of rays consisting of 4 columns of large, thick,
convex plates. These are the supra- and inframarginal columns.
The supramarginal plates of each ray are distally in contact with
one another along the mid-line, but proximally they are forced more
and more apart by a series of small, irregular, accessory plates. In
the center of the disk there is a small, convex disk plate around which
are situated radially 5 other small ossicles, all of which are sepa-
rated from each other apparently by a considerable number of minute
intercalary pieces. Just inside of the axils are 5 large, sub triangular,
uiterradial plates and adjoining these directly are 10 large and con-
spicuous ones, the proximal plates of. the supramarginal columns.
In each column there are about 13 plates, not counting the proximal
piece. Between all of the larger plates of the disk and for two-
thirds the length of the rays between the supramarginals, there are
numerous small, convex, irregularly shaped, accessory plates. All
BEVISIOK OF PALEOZOIC STELLEROIDEA. 71
the larger ossicles of both sides are centrally smooth, surrounded by
a finely pustulose border.
Madreporite situated interradially and distally upon the three
large plates just inside an axillary marginal plate, two of which are the
proximal plates of supramarginal columns of adjoining rays. It is
circular in outline, flat, and radially striated.
Inframarginal plates occupying the outer edges of both the actinal
and ab actinal areas; from 13 to 15 plates hi each column. The
supramarginal plates are situated above and inside the inframarginals,
which inwardly bear short, thick, club-shaped spines. Otherwise
they are ornamented like the abactinal plates.
Adambulacral ossicles small, subquadrate, each one alternating
distally with an inframarginal plate, but proximally they are some-
what more numerous, there being from 17 to probably 20 in a column.
On these plates lie two series of spines the two short, thick, club-
shaped ones of each mframarginal plate while on the inner side of
each adambulacral there is inserted a linear brush of from five to seven
slender, short spines that are outwardly and laterally directed.
Ambulacral grooves very narrow, deep, and gradually tapering.
Ambulacral plates not well seen but apparently very short, one to
each adambulacral and directly opposite one another.
Axillary marginal plates five in number, very large and conspicuous
actmally, also showing slightly abactinally.
Locality and formation. The only good specimen was found many
years ago by Col. E. Jewett in the Siluric (Rochester shale) at Lock-
port, New York. A fragment of the ray of a much larger individual,
nearly twice the size of this type, was found by the same gentleman
at the same place. These specimens are now in the paleontological
museum of Cornell University (No. 7331) and were presented by
Hon. Ezra Cornell. Long after the study of the form was com-
pleted another specimen preserving two rays and showing the actinal
side was learned of in the University of Toronto (Walker collection,
No. 1008). It was found at Grimsby, Ontario.
Remarks. This species has never been well described nor figured,
and has accordingly led to endless generic confusion. Almost any
Paleozoic starfish with large marginal plates has been referred to
Pal&aster. This lack of generic conception was partially due, as
stated above, to the fact that the abactinal side remained unknown.
The specimen lies imbedded in blue shale showing the actinal area,
and the abactinal side was revealed by the writer by cutting away
the slab to the starfish and then developing the individual plates
with caustic potash.
In a general way Palseaster niagarensis resembles several species but
all are distinguished by generic characters. The nearest one is
Hudsonaster incomptus from the Cincinnatic strata, but it differs at
72 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
once in having five instead of four columns of large abactinal plates.
There are also a far less number of the accessory plates so numerous
on the disk and the proximal medial areas of the rays in Palseaster
niagarensis.
Hall 1 refers a fragment to this species which clearly is of quite
another species. It is very far removed from P. niagarensis and
for the present need not be considered. The specimen hus not been
seen by the writer.
AUSTRALASTER, new genus.
Plate 4, fig. 4.
Palseaster ( Monaster) ETHERIDGE, jr. (part), Mem. Geol. Surv. New South Wales,
Pal., No. 5, pt. 2, 1892, p. 74.
Monaster GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 345 (not of Etheridge, jr.).
Australaster SCHUCHERT, Fossiliiim Catalogus, Animalia, pt. 3, April, 1914, p. 12.
Austral=sout1n.eTn and aster. A Paleozoic starfish restricted to
Australia.
Generic diagnosis. Animal attaining a very large size, with five
long; slender, almost straight-sided rays, terminating bluntly.
Interbrachial areas comparatively small, widely concave, and occu-
pied by ponderous, single, axillary marginals and inframarginal
plates.
Abactinal area unknown (see A. (?) stutcfiburii) .
Actinally the animal is bounded by inframarginal plates which
however do not appear to be present in the distal third of the rays.
If such plates are present in these outer regions, they are either
very small or gradually pass from the actinal to the abactinal area,
the latter condition occurring ha related genera. The inframarginals
in the outer third of the rays are small, subquadrangular, and rounded,
thence they increase rapidly in size proximally, become decidedly
transverse, and in the axils the columns abut against a very large
axillary inf ramarginal. The largest plate is the axillary marginal, orad
to which occur the pairs of enlarged oral armature ossicles. The
inframarginals appear to be nearly smooth and devoid of spines.
The adambulacrals in the distal third of the rays are numerous,
very short, but quite wide that is, are decidedly transverse, and
decrease rather rapidly hi breadth toward the tips of the rays.
Proximally these plates also decrease quickly in width and rapidly
pass inside of the inframarginals. Inside of the axillary marginals
the plates are again prominent and especially the pairs of oral arma-
ture pieces. Each adambulacral plate bears prominent tubercles
for the articulation of stout but short spines. The columns have
the pieces arranged with a decided slant outward and distally.
i Palaeontology of New York, vol. 2, 1852, p. 352 and pi. 85, figs. 8-10.
REVISION OF PALEOZOIC STELLEEOIDEA. 73
Ambulacral furrows deep, slender and tapering slowly. Ambu-
lacra! plates unknown. Oculars not present.
Genoholotype. Palseaster (Monaster) giganteus Etheridge, jr. The
specific description and the bibliographic references are included in
the generic diagnosis.
Kes trie ted to the Lower Marine series of the " Permo-Carbon-
iferous" of New South Wales.
AUSTRALASTER(P) STUTCHBURH (Etheridge, jr.).
Palxaster (Monaster) stutchburii ETHERIDGE, jr., Mem. Geol. Surv. New South
Wales, Pal., No. 5, pt. 2, 1892, p. 73, pi. 13, fig. 1.
Monaster stutchburii GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 346.
This species is found associated with A. giganteus and its preserva-
tion is such that the form is very imperfectly known. In some
respects it resembles that species and may prove to be a smaller
specimen of it, but until the disk marginals are known the differ-
ences will remain obscure.
If A. (?) stutchburii has the large disk marginals of Australaster,
it then seemingly will have the essential structure of that genus.
Regarding this the writer wrote Professor Etheridge in 1900 and
he replied that " axillary plates are not visible in the specimen".
Since the abactinal area of A. giganteus is unknown, it is desirable
to give here the abactinal structure in A. (?) stutchlmrii, as it may
prove to be congeneric with A. giganteus.
" Abactinal surface moderately convex, bearing several (five or
six) rows of small convex polygonal plates, inclusive of the margi-
nal pieces, and becoming much crowded at the apices of the rays,
where they form oblique rows." If these plates throughout the
rays are as figured by Etheridge for the distal ends, the strongly
quincuncial arrangement of the polygonal, closely adjoining plates
is another good generic character for Australaster.
PROMOPAL^ASTERID^E, new family.
Progressive Phanerozonia with distinct columns of inframar-
ginal plates. Interbrachial areas more or less complex, composed
either of inframarginals, axillary interbrachials and ambulacrals,
or of these with the addition of interbrachial marginals and acces-
sory interbrachials, or entirely of adambulacral plates. Ambu-
lacrals as a rule opposite, but they may also be slightly alternating.
Podial openings through the sutures in the lateral corners of the
ambulacral plates, but proximally a few alternate pores may grad-
ually pass medially, when there are four columns of podial openings
in each ambulacral furrow.
Madrsporite abactinal.
Abactinal plates very numerous, generally small, either in dis-
tinct columns or rows or without either arrangement. The radial
74 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and supramarginal columns may be very distinct or may be ob-
scured as such. Accessory ossicles always more or less abundantly
developed.
Contains :
Mesopalaeasterinae.
Promopalseas fcerinse.
Anorthasterinse.
new subfamily.
Palxocwnidde GREGORY, Treat. Zool., vol. 3, Echinoderma, 1900, p. 254.
Primitive Promopalseasteridse with the interbrachial areas small,
composed of one pair of inframarginals, single axillary infcerbra-
chials, and the adambulacral plates.
Contains :
Mesopalszaster, new genus.
Spaniaster Schondorf.
Miomaster Schondorf.
Devonaster, new genus.
MESOPAL^EASTER, new genus.
Plates 7-11.
Palseaster (part) of AUTHORS.
fArgaster HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 287: rev.
ed., 1868=1870, p. 329.
Mesopalseaster SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914,
p. 24.
Meso = middle, and Palseaster. Not the largest of Palseasters of
authors.
Generic characters. Rays five, slender and tapering. Disk small,
with incipient interbrachial arcs.
Abactinal area with distinct columns of radial and supramar-
ginal plates, between which are inserted rows of accessory pieces,
one or two on each side of the medial columns. In mature specimens
the radial columns are less prominent proximally than distally.
The accessory columns proximally may also have here and there
inserted on each side a smaller piece, when the area between the
supramarginals has not only three or five columns of plates but also
other additional small plates. The inframarginal columns margin
the rays, while the supramarginals lie somewhat inside and above
the former. Between the two columns of marginals there may be
a row of accessory ambital plates. The arrangement of the plates
of the disk appears to be as follows: In the center there is a small
plate around which is a first ring of 7 pieces, followed by a sec-
ond ring having not less than 14 plates, and these are nearly of
one size. The medial ray columns terminate on the disk in 5
REVISION OF PALEOZOIC STELLEROIDEA. 75
large radial plates, but inside of the proximal supramarginals there
is no basal interradial plate as in Hudsonaster.
Madreporite unknown.
Inframarginal and adambulacral plates as in Hudsonaster, except-
ing that two of the former and a single axillary plate are in each
interbrachial area. Oral armature so far as preserved adambulacral,
consisting of five pairs of basal adambulacrals.
A single, pentagonal, axillary interbrachial plate is present in
each area, orad to the two axile inframarginal plates.
Ambulacral grooves deep and almost closed by the adambulacral
columns. Ambulacral plates carinate and apparently one to each
adambulacral ossicle. Their relation to one another is not definitely
determinable, but apparently they are slightly alternate or nearly
opposite.
Genoholotype. Palxaster shafferi Hall.
Distribution. The species of Mesopalseaster are restricted to
America in the upper portion of the Ordovicic and the lower portion
of the Siluric. A species doubtfully referred to this genus occurs
hi the Lower Devonic of Germany, and another in the Upper Devonic
of Maryland.
The following are the species of Mesopalseaster:
M. shafferi (Hall). Middle and upper beds of the Cincinnatic
series. Possibly also in the Utica formation of New York.
M. intermedius, new species. Maysville formation.
M. finei (Ulrich) . Eden formation.
M. (?) lanceolatus, new species. Utica formation.
M. proavitus, new species. Eden formation.
M. (?) wilberanus (Meek and Worthen). Richmond series.
M. (?) dubius (Miller and Dyer). Upper Trenton.
M . (?) antiquus (Troost) . Upper Trenton.
M. (?) parviusculus (Billings). Siluric (Lower Arisaig).
M. (?) cataractensis, new species. Siluric (Cataract formation).
M. granti (Spencer). Siluric (Cataract formation).
M. bellulus (Billings). Siluric (Rochester shale).
M. caractaci (Gregory). Caradoc of England.
M. (?) acuminatus (Simonovitsch) . Lower Devonic of Germany.
M. (?) clarki (Clarke and Swartz). Upper Devonic (Jennings
formation) .
Remarks. Mesopalseaster apparently developed directly out of
Hudsonaster in that the single axillary marginal plates are in the former
genus in nearly all the species pushed inward (interbrachial) and
their former place occupied by the proximal plates of adjoining
inframarginal columns. The same generic feature also distinguishes
Mesopalseaster from Palseaster. Further, in Hudsonaster there are
no abactinal accessory ray plates, but in Mesopalseaster one or two
more or less completely developed columns are inserted on each side
76 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
of the radial columns or, in other words, between the radials and
supramarginals. Between the inframarginal and supramarginal plates
in Mesopalseaster there are also either a few incipient accessory plates,
which in young specimens are wholly absent, or there is a complete
column of these ambital plates. The accessory and axillary inter-
brachial plates are additions to the generic structure of Hudsonaster
and Palseaster, and further distinguish Mesopalseaster from both.
This progression toward a greater number of columns of abactinal
plates attains its maximum in Promopalseaster and is most marked in
P. magnificus, which has not less than 28 columns at the base of a
ray and 5 interbrachial marginal plates in each interbrachial area.
The ambulacral furrows, which are very narrow in Hudsonaster, are
likewise so in Mesopalseaster, but are wide in Promopalseaster. Spi-
nosity, which is practically absent abactinally in Hudsonaster, is
developed in Mesopalseaster and very pronounced in some forms of
Promopalseaster (P. spinulosus and P. dyeri) . This is apparently also
true for the spines of the actinal area.
The youthful plate structure of recent species seems to be retained
to maturity in Hudsonaster and somewhat so also in the genotype
of Mesopaldeaster but in Promopalseaster the central area of the disk
is occupied by numerous very small plates and no definite arrange-
ment is discernible, though this is in large part due to their displace-
ment through fossilization. However, it is certain that the basal
radials and interradials do not increase nor maintain a relative size
as in Hudsonaster and in the genotype of Mesopalseaster (here
radials only), but must have diminished to that of the accessory
plates from which they are now not distinguishable. From Hudson-
aster through Mesopalseaster into Promopalseaster the animals are
constantly increasing in size, and this also continues in the species
of the latter genus, attaining culmination in P. magnijicus and P. dyeri.
While the generic abactinal differences between Hudsonaster and
Mesopalseaster are easily made out, this is not so readily accom-
plished between the latter and Promopalseaster. All of the abactinal
generic characters of Mesopalseaster are more pronounced in Pro-
mopalseaster excepting that in the former the ambital accessory
plates are almost always undeveloped as columns, while in the latter
these plates are always well developed in one or more columns.
When the actinal area is shown, Mesopalseaster is at once distin-
guished from Promopalseaster in that it has but one axillary inter-
brachial plate in each interbrachial area, while the latter has always
two or more up to five ( = interbrachial marginals) in each area.
Devonaster in some respects is closely related to Mesopalseaster
and may be in the line of descent from the latter. However, the disk
in Mesopalseaster has far fewer abactinal plates, and there are no
distinct interbrachial arcs with numerous small accessory plates
REVISION OF PALEOZOIC STELLEROIDEA. 77
as in Devonaster. The pieces of the radial columns in Mesopalseaster
adjoin one another and are not separated by intercalary plates, nor
are these nearly so numerous between the radial and supramarginal
columns as in Devonaster. These accessory plates in Devonaster are
also irregularly disposed and not in columns as in Mesopalseaster.
The name Argaster has not been denned nor is it mentioned in such
a way that anyone will recognize that Hall intended it for a new genus
or subgenus. In addition, its genotype (Asterias antiqua = Mesopalse-
aster (?) antiguus) is a poorly known species. Under these circum-
stances, and the additional one that the name Argaster never has had
any standing as a genus, it should not now be rehabilitated by
paleontologists.
MESOPALSEASTER SHAFFER! (Hall).
Plate 8, figs. 1, 2.
Palseaster sha/eri HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 284;
rev. ed., 1868=1870, p. 326, pi. 9, fig. 1. MEEK, Geol. Surv. Ohio, Pal., vol. 1,
1873, p. 66, pi. 4, fig. 1.
Original description. "Body of about medium size composed of
five tapering, acutely pointed rays, which, in the specimen examined,
measure seven-eights of an inch from the center of the disk to the
extremity. The lower side of the ray is formed of two ranges of plates
bordering the ambulacral groove. The marginal range consists of
moderately convex plates which gradually decrease in size from the
base to the extremity of the ray, 22 or 23 in number [not more than
20 exist in any ray of the type-specimen], besides a small terminal
one at the angle of the range; each plate of the marginal range is
marked on its outer surface by a comparatively large cicatrix for
the attachment of a strong spine. The inner range of plates (adam-
bulacral) are somewhat smaller, about the same in number, alternat-
ing with those of the marginal range; the basal pair (oral plates)
are elongate-triangular, and slightly constricted near the middle.
Ambulacral a,reas narrow, composed of a double range of poral plates,
which at the middle of the ray are about of equal length and breadth.
Pores not observed. [The writer could not make out these plates in
the type-specimen and believes those described to be the inner sides
of the actinal plates.] Upper surface of the ray composed of three
ranges of subnodose plates, the outer ranges bearing a strong spine
on each plate; the central range apparently destitute of spines. No
spines have been observed, but the plates are strongly carinate or
pointed."
Emended description. The largest specimen in the Harris collec-
tion measures: B, = 19 mm., r 5.5 mm., R = 3.5r. Another specimen
of the same collection: R = 16 mm., r = 5 mm., R = 3.1r. The
smallest known specimen in the Vaupel collection: R = 6.5 mm.,
78 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
r = 2.5 mm., R = 2.6r. The type-specimen, which is the largest knowii
example, measures: R = 21 mm., r = 6 mm., R = 3.5r. Width of ray
at base in various mature examples 5 mm. to 7 mm.
Rays more or less long, slender, abactinally slightly convex, with
marked longitudinal ridges, actinally flat or concave. Disk of medium
size, abactinally convex and without interbrachial arcs.
Abactinal area of rays occupied by a central range of radial plates,
on each side of which there is a range of accessory ossicles which
however do not attain the distal portion of the rays. In mature
rays there are in the proximal region also inserted a few plates on
each side of the intercalary columns. Outside of the latter are
the ranges of well-defined supramarginal plates which slightly overlap
the inframarginals. The marginal and radial plates are most promi-
nent, and all are highly carinate or pointed. There is a single large
spine on the apex of all radial and accessory plates and several
smaller ones are also arranged around it. The supramarginal plates
have numerous granules which bore articulating spines. The plates
of the disk can not be clearly made out, but in a small specimen
somewhat distorted there appears to be a small central plate sur-
rounded by a first ring of 7 ossicles followed by a second ring of
14 plates. These pieces are all nearly of a size. The medial ray
columns terminate in 5 large basal radial plates, while inside of
the proximal inframarginals there is no large interradial plate as in
Hudsonaster incomptus.
Ambital area very illy defined, but in the angles between the large
infra- and supramarginal plates there are here and there inserted
tiny accessory pieces. In a young individual there are none of these
accessory plates.
Madreporite unknown.
Inframarginal plates distinctly granulated, completely inclosing
all other abactinal plates and common to both the abactinal and
actinal areas. In mature specimens there are from 15 to 20 plates
on each side of a ray. Laterally each plate has a Well-marked small
pit, probably the place of attachment for rather strong spines, none
of which however have been observed.
Interbrachial areas with single, hexagonal, axillary interbrachial
plates.
Adambulacral plates slightly elongate-subquadrangular, of which
there are from 18 to 20 on each side of a ray; distally they appear to
pass over and to cover completely on the actinal side the inframar-
ginal plates. Between the adambulacral columns there is a more or
less wide ambulacral furrow. Each adambulacral plate bears on its
inner edge a rather thick but short, slightly striate spine and several
minute ones. Or ad to each axillary interbrachial there is a pair of
prominent elongate-triangular adambulacral plates bearing spines,
REVISION OF PALEOZOIC STELLEROIDEA. 79
one of the five pairs constituting a part of the oral armature. In other
words, the adambulacral columns of adjoining rays are continued
around the axillary interbrachials by two modified plates.
Ambulacral grooves deep, nearly closed by the adambulacralia.
Ambulacral plates not well shown but apparently carinate, one to
each adambulacral ossicle, and whether alternate or opposite can
not be stated.
Locality and formation. -In the middle and .upper stages of the
Cincinnatic group (Ordovicic). The type-specimen (No. 1195,
Amer. Mus. Nat. Hist.) was found by Mr. D. H. Shaffer at Cincin-
nati, Ohio, probably in the Maysville formation. Mr. Vaupel found
a young individual showing the abactinal area (No. 60605, U.S.N.M.);
another young one was secured by Mr. Faber (No. 9568, University of
Chicago), and Mr. George Oeh found an excellent matured individual
preserving the actinal area (now in the Yale Museum), all from the
Maysville formation on the hills back of Cincinnati. Two good speci-
mens and another small poor one are in the Harris collection of the
United States National Museum (No. 59391) and were found in the
Waynesville division of the Richmond formation near Waynesville,
Ohio.
A small asterid preserving two rays and a portion of the disk "is in
the United States National Museum collection (No. 23540) and was
found by Mr. W. P. Rust in the Utica shales one and a half miles east
of Rome, Oneida County, New York. It is associated with Trinucleus
concentricus, Dalmanella testudinaria multisecta, and other species.
The abactinal area only is shown, and appears to be that of Meso-
palseaster shafferi. It does not appear to be a Promopalseaster because
of the fewer columns of abactinal accessory plates, and the axillary
structure makes it almost certain that there is here but a single axil-
lary interbrachial plate. For the present it may be provisionally
referred to this species and attention is thus directed to it so that local
collectors may be on the lookout for other material.
Remarks. This species has actinally much the appearance of
Hudsonaster incomptus, but the latter is at once distinguished by not
having the five axillary interbrachial plates of Mesopal&aster shafferi.
M. (?) parviusculus (Billings) is smaller, with far less plates in all the
columns than in M. shafferi. M. (f) parviusculus and M. granti differ
also abactinally and ambitally in the better developed accessory
plates.
Cat. Nos. 60605, 59391, 23540, U.S.N.M.
MESOPAL^ASTER INTERMEDIUS, new species.
Plate 9, fig. 4.
This species is perplexing in its generic construction because of its
intermediate or transitional characters on the one hand (but least)
to Hudsonaster, and on the other (more decidedly) to Mesopalxaster
50601 Bull. 815 S
80 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and prophetic of Promopalseaster in the conspicuous ambulacral
furrows.
Rays long and slender, elongate-petaloid, disk small. R = about
14 mm., r=4 mm.
The interbrachial areas each have but a single, large, finely granu-
lated, axillary plate that margins the axils, and in this agree best with
Hudsonaster, but the rest of the known skeleton is that of Meso-
palaeaster. In the former genus the ambulacral furrows are always
very narrow and the ambulacralia are rarely seen, while in M.
intermedius the furrows are wide and the ossicles are rather like those
of Promopalseaster, though the same kind of ambulacralia is also
known in Mesopalseaster. Orad to the axillary plates there were two
small oral armature pieces that belong to the ad ambulacral columns.
These are not now present in the specimen, but the distinct and
large facet of the proximal side of the axillaries leaves no doubt on
this construction.
Inframarginal columns well-developed proximally, though never
very prominent, and becoming rapidly smaller distally. The four
proximal pieces are the largest, beyond which there are at least
14 other ossicles, and all are closely in contact and alternate with the
adambulacrals. Another feature of these inframarginals is that not
more than six can be seen from the actinal surface, while the rest pass
on to the sides and distally even somewhat over to the abactinal
area. All of the plates are finely granulated.
Adambulacral columns the most prominent of the actinal skeleton,
margining the distal half of the rays, and in the proximal portion
pass more and more inside of the inframarginals, diminishing slowly
in size and finally depressed beneath the axillaries. There are 22
plates in a column, all more coarsely granulated than the inframar-
ginals, largest and most transverse near mid-length of the rays, and
all are drawn out inwardly into blunt points that adjoin the high
ridges of the ambulacralia. Therefore the adambulacral and the
adjacent ambulacral columns have the ossicles directly opposite one
another, but the adjacent ambulacral rows are slightly alternating.
Ambulacral furrows conspicuous and wide, widest in the proximal
third but converging somewhat quickly hi the mouth area and far
more gradually distally. Therefore the furrows are elongate-petaloid
in shape. There are about 20 ambulacralia in a column. Proximally
the columns unite around and beneath the small adambulacral oral
armature pieces. Ambulacralia square to transversely rectangular,
each with a high L-shaped crest that laterally adjoins the points of
the adambulacrals and medially bends orally. Podial openings large
between the corners of pairs of ambulacrals and adambulacrals.
There is a well-developed groove down the center of the ambulacral
furrows.
REVISION OF PALEOZOIC STELLEEOIDEA. 81
Abactinal area unknown, but probably that of Mesopalseaster and
nearest to M. shafferi.
Formation and locality. The holotype was found by Faber in the
Maysville formation at Cincinnati, Ohio, about 350 feet above the
Ohio River. The type is in the University of Chicago Museum
(No. 9575).
Remarks. This interesting species need be compared only with M.
shafferi. It differs in having the axillaries in the axils of the rays
and not inside the basal inframarginals as in the latter form. Then
in M. shafferi the inframarginals are actinal in position throughout,
while in M. intermedius they are more on the sides. Further, in the
new form the ambulacral furrows are well developed with large
ambulacralia, while in M. shafferi the furrows are narrow and the
ossicles rarely seen.
MESOPAL^SASTER FINEI (Ulrich).
Plate 7, fig. 5; plate 9, fig. 5.
Palseaster finei ULRICH, Journ. Cincinnati Soc. Nat. Hist., vol. 2, 1879, p. 19, pi.
7, figs. 15-156.
Original description. "Small; rays five, of medium length, rather
broad, pointed and narrower where they are attached to the much
contracted body [probably due to distortion], than they are about
the center of their length.
" Dorsal side of rays composed of four [about seven, the supra- and
inframarginals, radials, and two intercalary columns] rows of pieces,
that are quite close [?] fitting, as wide as long, from 12 to 14 in each
row, and increase in size inward to the disk, which is composed of
irregularly shaped and prominent pieces some of which are smaller
and others larger than those composing the rays; the pieces in the
marginal rows [four columns infra- and supramarginals] are more
prominent than the two [there are three, radials and two intercalary]
rows between them, and have a small pit in the center, probably for
the articulation of a spine [all of the plates originally bore several
short slender spines]. Madreporiform body rather small, circular,
very prominent, and marked by strong striae, which become more
numerous toward the margin by intercalation.
"Marginal [inframarginal] pieces on the ventral surface, convex,
quite as long as wide, and numbering in different specimens on each
side from 11 to 12 [probably not more than 8 or 9]; the piece at the
junction of the rays is three times as large as any other of the series,
subcircular and very convex. [It appears that this large plate is an
axillary interbrachial since upon it proximally rest two basal plates
of the inframarginal series.]
82 BULLETIN" 88, UNITED STATES NATIONAL MUSEUM.
" Adambulacral plates more prominent [less prominent than the
inframarginal plates], slightly wider than long, and numbering, on
each side, from 9 to 10 [probably not less than 16 in each column].
Ambulacral pieces a little wider than long, not alternating with the
adambulacral plates, and each provided with a rather sharp ridge
across most of its width.
"There are 10 oral plates [oral armature] formed by the junction
of the adambulacral rows, which in form and size are scarcely dis-
tinguishable from the other plates of those series.
" Greatest breadth measuring between the opposite extremities of
the rays ; 0.7 inch; breadth of rays at their inner ends, 0.1 inch;
length of same, 0.3 inch; diameter of madreporiform piece, 0.02
inch."
Formation and locality. This small species appears to be common
at a very restricted horizon near the base of the Eden formation
(Fulton beds), exposed at low water mark in the Ohio Biver in the
eastern portion of Cincinnati, Ohio.
Remarks. Not one of the twenty examples seen is well preserved
and all the plates are more or less separated. The general structure
seems to be that of Mesopalseaster. The composition of the axillary
area is not positively ascertainable but one specimen shows what
appears to be a large axillary interbrachial plate (see pi. 7, fig. 5),
distally surmounted by two somewhat smaller inframarginals. This
is probably the true structure since it is so in all the smaller Meso-
palseasters, as M. lanceolatus, M. parviusculus, M. granti, and M.
proavitus.
The differences between M. finei and M. proavitus are not great.
The former is a smaller species with less plates in all of the ranges,
more prominent axillary inframarginal and axillary interbrachial
plates, and is also found at a lower geological horizon. M. finei
differs from the small species M. parviusculus of the earliest Siluric
in having more plates in all of the columns. M. granti is a larger
species and with smaller and more numerous plates than in M. finei.
M. lanceolatus is probably the most closely related to M. Jinei but is
distinguished abactinally by the diagonal arrangement of the plates
on each side of the radial columns.
Cat. No. 60604, U.S.N.M.
MESOPALJEASTER(P) LANCEOLATUS, new species.
Plate 4, fig. 3.
Measurements: R = 4.5 mm., r=1.3 mm., R = 3r ) .
Rays short, stout, distinctly lanceolate actinally and rapidly ta-
pering abactuially. Disk comparatively large, abactinally convex.
Interbrachial arcs distinct but small.
Abactuially the disk has a ring of large, strongly stellate plates
which are the basal plates of the radial and supramarginal columns.
REVISION OF PALEOZOIC STELLEEOIDEA. 83
Inside the ring the plates can not be made out, due to the small size
of the specimen. Rays each with a column of radials and two supra-
marginals between which are inserted columns of diagonally arranged
accessory plates. A few ambital accessory plates are probably also
present. The arrangement of the ossicles on the abactinal areas of
the rays is like that hi Promopalxaster magnificus.
Madreporite small, subconical, situated near the edge of the disk,
depressed between adjoining basal plates and marked by 10 or 11
sharp, somewhat spirally arranged ridges.
Inframarginal plates large, about six in each column proximal to
the very large single axillary interbrachial plate. These columns in
the distal half can not be seen actinally as they pass over on the dorsal
side.
Axillary interbrachial plates large, one in each axil, and not com-
pletely inclosed outwardly by the basal inframarginals. In this
character the form is still in the Hudsonaster stage.
Adambulacral plates small, subquadrangular, with 16 plates in each
column. The plates of the adambulacral oral armature are larger
and wedge-shaped, and are situated directly inside of the axillary
interbrachial plates.
Ambulacra! plates one to each adambulacral, apparently opposite
and with proximally converging ridges. Podial openings apparently
along the outer edge between the sutures of adjoining plates.
Formation and locality. Two specimens were found by the late
Prof. Charles E. Beecher near Rome, New York, in the Triarthrus
bed of the Utica formation that preserves the entire ventral anatomy
of these trilobites. The co types are in the Museum of Yale University.
Mr. W. S. Valiant in a letter states that they are common in a layer
2 to 3 inches above the Triarthrus bed.
Remarks. This species is most closely related to M. finei, from
which it is readily distinguished abactinally. Here there are more
and smaller plates than In M. lanceolatus, the interpolated plates are
arranged diagonally and not radially as in M. finei, while the basal
plates of the former are apparently much larger and are strongly stel-
late. Actinally there are in M. lanceolatus more adambulacral plates
in each column, although the specimens are only half the size of M.
finei.
MESOPAL^ASTER PROAVITITS, new species.
R=18 mm., r^about 4.5 mm. The specimens are distorted, however,
and no exact measurements can be given.
On the distal abactinal portion of the rays the infra- and supra-
marginal and radial plates closely adjoin. Proximally, however,
between the two marginal series there is inserted a column of inter-
calary plates gradually increasing in size and two other similar rows
of plates are placed between the supramarginal and radial columns.
84 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
There are therefore at the base of the rays no less than 11 columns
of plates. The radial columns proximally have a tendency to become
less distinct and indistinguishable from the accessory pieces. The
plates in general are highly convex, granular and abundantly spine-
bearing. There are not less than 20 plates in each supramarginal
series.
Madreporite highly convex, broadly oval in outline and radially
striated.
Inframarginal plates finely granular, highly convex, subquad-
rangular in outline and increasing in size slowly proximally. About
23 in each column.
Adambulacral plates like the inframarginals but not increasing
much in size proximally, with about 29 in each column. As in other
species of this genus, each plate bears three prominent spines, two
laterally and one ambulacrally.
Ambulacral plates one to each adambulacral plate, and with a
sharp ridge which medially bends abruptly orally. Podial openings
as in other forms of Mesopalseaster.
Interbrachial areas occupied by single axillary interbrachial plates
upon each of which rest proximally two axillary inframarginals.
The specimens are not figured, as an illustration can not readily
be made.
Formation and locality. Three more or less entire specimens and
fragments of four rays were found by Dr. E. O. Ulrich in the Eden
shale exposed back of Covington, Kentucky, at an horizon about
100 feet above low water in the Ohio River. All the specimens a^e
now in the Herzer collection, a part of the late Prof. James HalFs
private collection, recently purchased by the University of Chicago.
Remarks. M. proavitus and M. granti are closely related, since
both have 11 columns of abactinal plates, including the marginals,
and they are nearly alike actinally. However, the former is twice as
large and appears to have more conspicuous radial and supramarginal
columns than the latter. These differences and the further fact that
one occurs in the Utica and the other in the earliest Siluric will serve
to distinguish the two species for the present. M. proavitus may
prove to bo identical with M. (?) wilberanus, which see for further
remarks.
MESOPAL^ASTER (?) WILBERANUS (Meek and Worthen).
Petraster wilberianus MEEK and WORTHEN, Proc. Acad. Nat. Sci., Philadelphia,
for 1861, vol. 13, 1862, p. 142.
Palseaster wilberanus HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868,
p. 285; rev. ed., 1868=1870, p. 328.
Original description. "This beautiful starfish resembles rather
closely Petraster rigidus of Billings, 1 but is smaller, and has more
1 Decade 3, Org. Rem. Canada, pi. 9, fig. 3o.
REVISION OF PALEOZOIC STELLEROIDEA. 85
slender rays, with more angular spaces between. It also differs in
having but two ranges of plates on each side of the ambulacral
grooves, on the under side, instead of three. These pieces are about
of the same size in each row, and sometimes appear to alternate;
they are all rather prominent, and those of the outer range project
a little laterally in the form of small nodes. Toward the outer
extremities of the rays, however, the lateral ranges are contracted
behind the others, so as to be scarcely visible from below. About 23
of these pieces may be counted on each side of the ambulacral furrow
in each ray. The ambulacral furrows are very narrow, and indeed
seem to be closed toward the extremities of the rays, by the gradual
approximation of the inner rows of pieces on each side, which alter-
nate and appear to fit together.
"Our specimen only shows the under side, but along the outer
margins of two of the rays, there is some appearance either of the
overlapping of some of the dorsal parts by pressure, or of a slightly
developed disk. This part does not seem to have any distinct range
of marginal plates, but appears to be made up of small pieces, covered
with granules, or bases of small spines.
"We take pleasure in dedicating this interesting species to
Prof. C. D. Wilber, of the Illinois State Normal School, to whom we
are indebted for the use of the only specimen we have seen.
"Greatest diameter about 1 inch; smaller diameter 0.33 inch.
"Locality and position. Oswego, Kendall County, Illinois, in rocks
of the age of the Trenton or Hudson River Group of the New York
series." Probably equivalent to the Richmond beds of the highest
Ordovicic. The whereabouts of the type-specimen is not known.
Remarks. Hall pointed out that this species is not a Petraster
because P. wilberanus has but two columns of plates on each side of
the ambulacral groove, while Petraster has in addition "a few disk
plates on the ventral side." He therefore referred it to Pal&aster.
However, Meek and Worthen describe the abactinal area as having
no " distinct range of marginal plates, but appears to be made up of
small pieces, covered with granules, or bases of small spines." The
characters as far as mentioned appear to be those of Mesopal&aster.
Should there prove to be in P. wilberanus single axillary interbrachial
plates, then all of the characters will be in harmony with Mesopalse-
aster and M. proavitus may prove to be a synonym of it. This
can not be proven at present since we have not been able to locate
the type-specimen.
MESOPAL^ASTER (?) DUBITJS (Miller and Dyer).
Palseaster dubius MILLER and DYER, Cont. to Pal., No. 2, 1878, p. 5, pi. 4, fig. 8.
Original description. "Pentagonal; rays longer than the diameter
of the body, and uniformly tapering. The ambulacral groove is
sharply angular in the middle, formed by two series of plates having
86 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
their ends together in such manner as to make an angular gutter
[the plates of adjoining columns are arranged practically opposite].
The length of these plates is three times as great as the diameter.
The plates are placed with the length across the rays. There are
15 plates on each side of the groove in each ray, in the length of one
quarter of an inch [this is an error as there are about 20 plates in a
column]. The diameter of the body is three-tenths of an inch.
"This species is founded upon a single specimen, in Mr. Dyer's
collection, which shows only part of the ventral side. The ends of
the rays [most of the adambulacrals] and marginal plates arc destroyed
[each axillary area appears to bear one axillary interbrachial plate].
The parts preserved seem to distinguish it from any species hitherto
described."
Formation and locality. In the original description the locality
is given as Cincinnati, Ohio, but the character of the rock shows that
it is from the uppermost portion of the Trenton limestone, probably
opposite Cincinnati, in the river quarries at Ludlow, Kentucky.
The type (No. 25) is in the Museum of Comparative Zoology, Harvard
University.
Remarks. This specimen preserves well the very wide ambulacrals
but almost no other plates. The adambulacrals are nearly all removed
and but very little is to be seen of the marginals. In three of the
axillary areas there are single interbrachial plates, one of the essential
features of Mesopalseaster. It is referred to this genus provisionally,
as nothing is known of the abactinal side.
MESOPAL^ASTER (?) ANTIQUTJS (Troost).
Asterias antiqua TROOST (not Hisinger 1837), Trans. Geol. Soc. Penn., vol. 1, 1835,
p. 232, pi. 10, fig. 9; Proc. Amer. Assoc. Adv. Sci., vol. 2, 1850, p. 59 (cat. name).
Petraster (?) antiqua SHUMARD, Trans. St. Louis Acad. Sci., vol. 2, 1866, p. 386
(catalogue name).
Palxaster (Argaster) antiqua HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist.,
1868, p. 287; rev. ed., 1868=1870, p. 329.
Palxaster antiquus MILLER, N. Amer. Geol. Pal., 1889, p. 265 (cat. name).
Argaster antiqua GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 345 (gen. ref.).
Pal&aster antiqua WOOD, Bull. U. S. Nat. Mus., No. 64, 1909, p. 105, pi. 8, fig. 1.
Although this is the first recorded American fossil starfish, very
little is known about it and that little is mostly of a misleading nature.
The specimen lies on a limestone slab and is very badly weather-
worn or it may have been treated with hydrochloric acid so that now
it is nothing more than a polished section of a starfish. An illustra-
tion that will show its actual characters can not be made.
Professor Hall errs in stating that Asterias antiqua has "ambu-
lacral grooves occupied by a single row of subquadrate ossicula, which
extend across and alternate with the adambulacral plates of each mar-
gin. * * * It is possible that this character may prove to be
of generic importance." It may have been this character on which
REVISION" OF PALEOZOIC STELLEROIDEA. 87
Hall thought of basing the genus Argaster, but he does not character-
ize it nor even mention that the name in parenthesis is intended
as a new term with Asterias antiqua Troost as the genotype. This
species, like all other Paleozoic starfishes, has double columns of
ambulacral plates. This the specimen clearly demonstrates on
the edge of the slab where the distal parts of the rays are broken
away. Argaster should therefore be regarded as a nomen nudum, and
should A. antiqua prove to be a Mesopal&aster, it should not be
made to displace this genus.
Asterias antiqua has about 15 inframarginal plates in each column
and about 32 in each adambulacral column. Two of the latter plates
meet as usual in a pair of triangular oral armature pieces.
Each axil is occupied by two large, quadrangular, basa] inframar-
marginal plates. Between these proximally there is a large, widely
triangular, interbrachial plate the apex of which may or may not
attain the margin. Proximal to each axillary interbrachial plate
and between the four or five pairs of axillary adambulacral plates,
there is in the specimen an open space in each of the five areas.
What additional plates, if any, occupied this area is not de terminable.
It may be that the axillary interbrachial plates occupied the entire
interbrachial areas and that the present hiatus is due to the worn
condition of the specimen. This appears to be the most natural
interpretation as it is the normal inter brachial structure of Mesopa-
Iseaster. In Promopalseaster there are always two, three, five, or seven
interbrachial marginal plates hi each area, a fact which excludes Aste-
rias antiqua from that genus.
The abactinal area is not visible, but many of these plates are
squeezed beyond the inframarginals, showing the presence of num-
erous small plates recalling Mesopalseaster and Promopalseaster.
Formation and locality. Troost's label reads: " Lower limestone
on Harpeth Kiver, Davidson County, Tennessee." His manuscript
reads: "It was found * * * on Harpeth Kiver, Davidson
County, Tennessee. Associated with Spirifer lynx [PlatystropJiia
l}iforata] } Cyathop7i[y]Ua [ Streptelasma] } Orthis [testudinaria], etc."
This is apparently the same horizon as that about the city reservoir
in Nashville, which is now regarded as of Upper Trenton (Catheys)
age. The specimen is in the United States National Museum.
Cat. No. 39914, U.S.N.M.
MESOPAL^EASTER(?) PARVIUSCULUS (Billings).
Plate 9, fig. 1.
Palseaster parviusculus BILLINGS, Canadian Nat. and Geol., vol. 5, 1860, p. 69,
figure. DAWSON, Acadian Geology, 2d ed., 1868, p. 594, fig. 197. HUDSON,
Bull. N. Y. State Mus., No. 164, 1913, pis. 1-4.
Original description. ' l The specimen is about six lines in diameter.
The rays are two lines in length and one and a half in width at the
88 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
base, tapering at an angle of a little less than 45. The five oral
plates are subpentagonal, about half a line in width. The first
adambulacral plates of each pair of adjacent rays are in contact
with each other outside of the oral plates, and not completely sep-
arated as they are in P. niagarensis. There are six or seven adam-
bulacral plates on each side of the ambulacral groove in each ray,
and they gradually decrease in size from the oral plate outward to
the point of the ray. The width of the ambulacral groove is equal
to one-third the width of the ray and consequently the adambu-
lacral rows of plates are also each equal to one-third the whole width
of the ray. In each groove there are two rows of small and appar-
ently nearly square ambulacral plates, 12 or 14 in each row, and they
seem to be continued round on the inner margin of the oral plates;
the mouth is about one line wide."
Emended description. Measurements: R = 6 mm., r = 2.5 mm.,
R = 2.4r. Width of ray at base 3 mm.
Rays short, stout, tapering rapidly. Disk comparatively large,
and without interbrachial arcs.
Abactinal area unknown.
Inframarginal plates at the base of the rays very large and thick,
highly convex, duninishing rapidly distally, usually quadrangular
in form and seven on each side of a ray.
Just within each axil, inside of the basal inframarginals, there is
a single, large, very convex, pentagonal, axillary interbrachial plate.
Adambulacral plates gradually sinking below the inframarginals
proximally, convex and quadrangular in form. There are 13 plates
in each column adjoining the very narrow ambulacral furrow, or
nearly two plates to each inframarginal ossicle. Two of these
plates (oral armature) are situated in front of each axillary inter-
brachial, uniting the adambulacral columns of adjoining rays.
Ambulacral plates unknown.
Locality and formation. The type-specimen, a natural mold, was
found by Rev. D. Honeyman in the Lower Arisaig of the Siluric
( = Clinton) rocks at Arisaig, Nova Scotia. The holotype is in
Redpath Museum of McGill University, Montreal, Canada, and the
illustration is taken from a gutta-percha squeeze kindly made for
the United States National Museum by Sir William Dawson.
Remarks. The general actinal appearance of this little asterid
reminds one much of Palseaster and Hudsonaster. It differs, however,
at once in a little detail of considerable importance, in that each
axillary area has three plates, while in Palseaster and Hudsonaster
there is but one. In other words, the large axillary plate of those
genera has in M. (?) parviusculus been pushed orally and is here an
interbrachial plate, while the axils of the rays are occupied by two
basal inframarginal plates. For further remarks on this species see
M. granti (Spencer).
REVISION OF PALEOZOIC STELLEROIDEA. 89
So long as the abactinal area of M. (?) parviusculus remains
unknown its generic position will be doubtful. For the present its
relations appear to be nearest to Mesopalseaster and to M. (?) cata-
ractensis.
Cat. No. 60620, U.S.N.M.
MESOPAL.EASTER (?) CATARACTENSIS, new species.
Plate 9, fig. 2.
The specimen of this species has been confused with M. (?) gmnti,
from which it differs in being smaller and in having far fewer plates
in the columns. The actinal side of M. (f) cataractensis alone is
known, while of M. (?) granti only the abactinal, but even so the
above comparison can still be made because the size of plates is
relatively about the same in species of Mesopalseaster.
Measurements: K = 9 mm., r=3 mm., R = 3r.
The species is most closely related to M. (f) parviusculus, but
differs in being larger with more plates in the columns. Of M. (?)
cataractensis only the actinal side is known, with 12 to 13 plates
in each inframarginal column (7 in M. (?) parviusculus) and
about 19 in the adambulacrals (13 in M. (?) parviusculus). Then
the inframarginals increase very rapidly in size proximally, while
in M. (?) cataractensis there is but little enlargement.
A single small, pentagonal, axillary interbrachial plate occurs
in each area immediately beneath the basal inframarginals. These
are much smaller relatively than those in M, (?) parviusculus.
Ambulacral plates unknown. Ambulacral furrows deep, nearly
completely closed by the adambulacral columns.
Formation and locality. in the base of the Siluric (Cataract for-
mation) at Hamilton, Ontario. The holotype was found by Mr.
Elliott near the city reservoir and is in the collection of the Hamilton
Natural History Society. It was loaned to the writer by the late
Col. Charles Coote Grant.
MESOPALJEASTER GRANTI (Spencer).
Plate 9, fig. 3.
Palseaster granti SPENCER, Bull. Mus. Univ. Missouri, No. 1, 1884, p. 53, pi. 7,
fig. 1.
Original description. "Body stellate and small with short arms,
about 2 centimeters across; disk less than 1 centimeter wide, and
apparently formed by the junction of the rays; rays 5 millimeters
wide at base, tapering slowly, and terminating in rounded extrem-
ities, at about 8 millimeters from junction of their base with the
disk; upper surface of rays composed of 5 [not less than 9 in mature
specimens at mid-length of rays] ranges of highly convex or tuberculi-
form plates (the marginal [supramarginal] and ventral [inframarginal]
90 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
rows being the most conspicuous) and separated from each other by
minute plates (becoming fewer on approaching the extremities of
the rays) .
"The ellipsoid [supra] marginal tuberculiform plates number
about 12 [20 in mature examples] for each complete side of the rays,
and the central range [radial] is composed of a similar number, but
in form these plates are more circular. The disk between the ter-
minal central [radial] row of plates of the rays is crushed and struct-
ureless, except a slight elevation in the center. The terminal [proxi-
mal] plates of the marginal series are larger than the others of the
range. The madreporiform tubercle is of a spherical form and rela-
tively large, being nearly 2 millimeters in diameter, and is situated
at the axil of two rays. Both the tuberculiform plates and the
madreporiform tubercle have a granulated surface.
"The ventral side is unknown.
Emended description. A mature specimen measures: R=16 mm.,
r=5 mm. ; R = 3.3r.
Rays short, stout, tapering rapidly, and with very small inter-
brachial arcs.
Abactinal area of disk unknown. Along the center of each ray
there is a radial column of small, highly convex plates bounded on
each side, at about the mid-length of the rays, by two columns of
smaller, strongly tumid, accessory plates. The radial column is,
however, not distinctly differentiated from those adjoining. Out-
side of these are the supramarginal columns, each with about 20
plates which are larger and more easily distinguished than the radial
plates. Dis tally the rays have only the ossicles of the infra- and
supramarginal and radial columns, with none of the accessory plates
as yet developed.
Ambital area with one column of very small accessory plates
like the marginal plates on each side of them. These accessory pieces
disappear before attaining the apex of the rays.
Madreporite spherical in form and relatively large, nearly 2 milli-
meters in diameter, and situated at the .axil of two rays between
two adjoining supramarginal columns.
Formation and locality. From the Cataract formation, the basal
deposits of the Siluric system, near the reservoir in the city of
Hamilton, Ontario. Col. Charles C. Grant, after whom the spe-
cies is named, discovered most of the specimens, which are six in
number, two in the Spencer collection, now destroyed by fire, one
in the Redpath Museum of McGill University, Montreal, another
in the Geological Survey of Canada at Ottawa, and two in the
Grant collection, one of which has generously been donated to Yale
University.
Remarks. This species is apparently closely related to M. bel-
lulus, which see for comparison.
REVISION OF PALEOZOIC STELLEEOIDEA.
91
MESOPAL^EASTER BELLULUS (Billings).
Text fig. 7.
Petraster bellulus BILLINGS, Geol. Surv. Canada, Pal. Foss., vol. 1, 1865, p. 393,
fig. 368. MILLER, N. Amer. Geol. Pal., 1889, p. 269, fig. 389.
Original description. "Deeply stellate, about 18 lines across; disk
5 lines wide; width of rays, at the base, half the width of the disk,
uniformly tapering to their extremities; ambulacral grooves, narrow
and deep, with about 30 adambulacral plates on each side. These
plates are strongly convex and nearly square [distally, but proximally
are wider than long]. Outside of these there is a row of [about 21
infra] marginal plates, which appear to [actually do] extend to the
extremities of the rays, but on this point there is some doubt, as the
specimen is not perfect. [The four proximal pieces increase rapidly
in size so that they are considerably larger than the others. All of
the actinal plates are granular.] There
appear to be one or two [there is but one
axillary] small disk plates between the
[basal adjoining infra] marginal and ad-
ambulacral plates just outside of the oral
angles.
"Locality and formation. Township of
Grimsby [Ontario]; in the Niagara shale
[ = Rochester shale].
1 i Collector. J ohnson Pettlt, Esq. , Fi G. ?. THE ORIGINAL FIGURE OF ME-
Grimsby." The holotype is in the Vic-
toria Memorial Museum, Ottawa, Canada
(No. 2665).
Remarks. The holotype and only
known specimen of this species was seen
by the writer at Ottawa, and has now been freed from the rock so
that the abac tin al side can also be studied. The generic charac-
ters are those of Mesopalseaster and one is impressed at once by the
almost specific identity of the abactinal side of the specimen with
that of M. granti. However, as the specimens show slight differ-
ences and come from different Siluric formations, it is thought best
to recognize them as distinct species. Moreover, the actinal side
of M. granti is still unknown.
Abactinally all of the skeletal pieces are of about the same size,
small, each ossicle irregular in outline, and more or less stellate.
There are about 17 in the length of one-half inch. This skeleton is
more or less disturbed and it is impossible to clean off all the adhering
shale, so that it is difficult to make out the detailed arrangement.
The following, however, can be made out: There is no true disk as
in Petraster , and the parts that appear as such are either adhering
shale or displaced ossicles due to distortion during sedimentation,
SOPAL.EASTER BELLULUS, AFTER BlL-
LINGS. NATURAL SIZE. THE SPECI-
MEN HAS STILL TOO MUCH SHALE
ADHERING TO IT AND BESIDES IS CON-
SIDERABLY DISTORTED, PREVENTING
ITS BEING SATISFACTORILY PHOTO-
GRAPHED.
92 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Distally the rays have columns of small radial pieces, and outside of
these are the two supramarginal rows. Proximally between these
primary columns there appear accessory pieces, one column of which
wedge in between the radials and supramarginals and from one to
three between the latter and the inframarginals. In this way finally
as many as 13 columns can be counted at the base of a ray.
The madreporite is situated near the margin between adjoining
supramarginal columns, and is conspicuously protruded and large
when compared with the small adjacent ossicles. It is a large,
radially striate, irregularly circular plate about six times larger than
the adjoining pieces.
MESOPALJ2ASTER CARACTACI (Gregory).
Plate 9, fig. 6; plate 11, fig. 1.
Palseaster caractaci SALTER, Cat. Foss. Mus. Practical Geology, 1865, p. 30 (nomen
nudum). NICHOLSON and ETHERIDGE, Mon. Silurian Foss. Girvan Disk,
Ayrshire, fasc. 3, 1880, p. 321 (no description). GREGORY, Geol. Mag.,
dec. 4, vol. 6, 1899, p. 344. SCHONDORF, Jahrb. nassauisch. Ver. Naturk.,
Wiesbaden, vol. 63, 1910, p. 227.
Protopalxaster caractaci SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr.
Soc. for 1913), 1914, pp. 21, 30, pi. 1, fig. 5.
Remarks. The writer was greatly pleased to receive from Dr. F.
A. Bather of the British Museum wax squeezes of the type-specimens^
because they clearly show that the genus Mesopalseaster is present in
Europe. M. caractaci has its closest relationship in M. sJiafferi, the
genotype of Mesopald&aster. The differences are easily to be seen on
the abactinal side, as illustrated in this work.
Rays long and slender, abactinally flattened. R = 12.5 mm.,
r3.S mm., R, = 3.3r. Radialia small and no larger than the pieces
of the accessory columns on either side of them ; the former alternate
in position with the pieces of the latter and all of them are more or
less spicular in outline. Basal radials slightly larger than the others.
The ossicles of the supramarginal columns large and solid, the most
prominent of the abactinal side, very large proximally and exceedingly
small distally; there are about 14 in a column. Basal inframarginals
most conspicuous of all abactinal plates, with each pair occupying
the axils, and together with the smaller basal radialia form a ring
bounding the disk. The disk has fallen in but still shows a few
small pointed plates.
" Madreporite is large, and placed close by one of the interradial
angles" (Gregory).
Inframarginal ossicles almost smooth, large and prominent proxi-
mally, decreasing rapidly in size so that not more than 9 can be
seen actinally, where the tiny pieces pass somewhat over to the
abactinal surface, or better the lateral sides; Gregory states that
there are 12 in a column. The basal pieces of adjacent columns do
not as a rule touch one another in the axils.
BEVISION OF PALEOZOIC STELLEROIDEA. 93
Interbrachial areas with single, large, hexagonal, axillary inter-
brachial plates.
Adambulacral pieces quadrangular, fairly uniform in size for two-
thirds the length of the rays and then become gradually smaller
distally; there are about 15 in a column. A peculiarity , of this
species is that the ambulacralia do not continue around the axillary
interbrachials, but cease with the basal inframarginals. There are,
therefore, in M. caractaci no adambulacral oral armature pieces.
Ambulacral furrows moderately wide, with as many rectangular
ambulacralia as there are adambulacrals, plus probably four more
in each column, and these continue around the axillary interbrachial.
Therefore, the oral armature consists entirely of ambulacralia. All
of these ossicles are directly opposite one another and have high
medial crests that are continuous across the ambulacral furrow.
Podial openings large laterally between the plates and the adambu-
lacrals.
Locality and formation. Caradoc sandstone, Soudley quarry,
Church Stretton, England. The holotype, a split nodule, is in the
British Museum (Natural History), No. 48206; wax squeezes from
which the photographs have been made are in the United States
National Museum. Other specimens, Bather states, are in the Museum
of Practical Geology, London; these are from Marshbrook and occur
at a somewhat higher geological level.
Remarks. Gregory wrote in 1899 that "the nearest ally of this
species is P. matutina (Hall)," but the illustrations here presented
wiU show that the relationship is with Mesopaldeaster shafferi and not
with Hudsonaster matutinus.
Cat. No. 60606, U.S.N.M.
MESOPAL^EASTER (?) ACUMINATUS (Simonovltsch).
Asterias acuminatus SIMONOVITSCH, Sitzb. d. mat.-naturw. Classe Akad. Wiss.,
Wien, vol. 64, Abt. 1, 1871, p. 100, pi. 3, figs. 2-2d.
Asterias acwninata SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 72, 109,
pi. 11, figs. 7, 8. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 62, 1909, p. 32, pi. 5, figs. 4, 5.
Roemeraster (?) acuminattis STURTZ, N. Jahrb. fiir Min., etc., 1886, vol. 2, p. 154.
Actinally this small and slender species is nearly in harmony with the
generic characters of Paldeaster and Hudsonaster, differing only in that
the axillary interbrachial plates barely extend to the margin of the
disk, while in those genera they are prominent and marginal in the
axils. In Asterias acuminatus they are as yet not crowded so far orally
as in most species of Mesopalseaster. It is a difference of degree, not
of kind; therefore this species has more the actinal character of
Mesopalseaster than Hudsonaster. Ambulacral plates unknown.
Abactinally Simonovitsch's species is said to have six columns of
tumid plates, all alike and persisting throughout the rays. If the
94 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
outermost columns of each ray are regarded as the inframarginals
and the ones immediately inside as supramarginals, two medial
columns remain to be homologized with those in Mesopalseaster. If
Simonovitsch is correct, then by this interpretation there are two
columns between the supramarginals. Both can not be radials and to
regard one as such will leave a unilateral and apparently an unnatural
development. If five or seven columns of abactinal plates were present,
this species would be hi harmony with Mesopalxaster. Since Simono-
vitsch's work is of the best, we have at present the only alternative of
supposing that the radials in Asterias acuminatus are suppressed and
that its two medial columns are homologous with the intercalary
abactinal plates of other starfishes. Among Paleozoic asterids this
development is rare and is found only in Encrinaster and Palseaster.
In that event, A. acuminatus is not a Mesopalseaster.
A single example of this species was found in the Lower Devonic
(probably Upper Coblenzian) near Braubach, Germany.
In his revision of the Lower Devonic starfishes of Germany,
Schondorf was not able to discover the whereabouts of the type-speci-
men and therefore had to leave this species as determined by Simono-
vitsch. The former states that actinally the structure reminds one
of Spaniaster latiscutatus, the genotype of Spaniaster, and closely
related to Mesopalseaster. For further remarks see Spaniaster.
MESOPAL^ASTER (?) CLARKI (Clarke and Swartz).
Plate 10, figs. 1, 2.
Pals&aster clarki CLARKE and SWARTZ, Maryland Geol. Surv., Upper Devonian,
1913, p. 543, pi. 46, figs. 3,4.
Original description. "This species is represented in the collec-
tions by a single specimen affording a pretty sharp cast of both sides
of a very regular and complete individual. In general structure and
appearance the species is quite similar, especially on the oral surface,
to Palseaster eucharis Hall of the sandy Hamilton shales of central
New York, but the latter is a much larger form.
"The disk is small, rays long and slender, thecal plates all promi-
nently developed. The ambulacral surfaces are represented only by
a narrow linear depression beneath which the ambulacral plates are
concealed. These depressions are bordered by thickened and somewhat
elevated quadrangular or pentangular ad ambulacra. The marginal
plates are in single rows, much thickened, with generally quadrangular
outline and convex surface, each projecting on the margin of the ray.
At each axilla is a single pear-shaped plate with its apex outward,
these plates being the largest in the individual. The abactinal surface
is tessellated by rows of strong convex plates of similar size to the mar-
ginal plates. Of these there are three rows, a median row of narrow
oval ones between the ends of which are interlocked the edges of the
much larger plates of the lateral rows which are highly convex and
REVISION OF PALEOZOIC STELLEROIDEA.
95
FIG. 8. SPANIASTER LATISCUTATUS, AFTER
SCHONDORF. SCHEMATIC ARRANGEMENT OF
THE ABACTINAL PLATES. C, PRESUMABLE
POSITION OF ANUS; Ce, CENTRAL PLATE;
Jn, BASAL INTERRADIALIA; mdp, PROBABLE
POSITION OF MADREPORITE; mo, SUPRAMAR-
GINALIA; RI, BASAL RADIALIA.
thickened in the center and greatly depressed to the sutures . Thus each
plate has a cushioned surface. Between the ends of each plate of the
middle row are two minute accessory plates lying in the angles at
which the lateral plates enter. At the base of each ray and upon the
disk is a single large plate whose surface rises into a high clavate node.
Between each two of these is one of
less height.. The central portion of
the aboral area is destroyed and no
trace of madrepore is seen.
"The width of this specimen from
tip to tip is 33 mm.
"Occurrence: Jennings formation,
Chemung member. Yellow sandstone
on the road northeast of Oakland,
Garrett County [Maryland], where it
is associated with Spirifer disjunctus.
"Collection: Maryland Geological
Survey."
Remarks. This clearly determined
species is a late survival of early
Paleozoic primitive asterids. The
characters so far as determinable in
the natural mold are those of Meso-
palseaster, but as the disk skeleton is not preserved, it is very prob-
able that when this feature is known the form will be seen to belong
to a new genus.
Genus SPANIASTER Sehondorf.
Plate 12, figs. 1, 2; text fig. 8.
Calaster SANDBERGER (not Agassiz 1835), Verst. d. rheinischen Schicht. Nassau,
1855, p. 381.
Spaniaster SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 60,
1907, p. 176; vol. 62, 1909, p. 30; Palaeontographica, vol. 56, 1909, pp. 73, 109.
Remarks. This genus with its single very small species has its
nearest relations with Mesopalseaster, in that it has a single axillary
interbrachial plate in each actinal axillary area, but differs from it
in that there are many more adambulacral and ambulacral plates
than there are inframarginals. The marked and generic difference,
however, is on the abactinal side, where there are but three columns
of plates, one radial and two suuramarginal, of large and thick ossicles
arranged in parallel rows, the pieces of which do not alternate with
one another; further, the supramarginals are almost completely
superposed upon the inframarginals, the two columns together, but
more particularly the inframarginals, bounding the rays and not the
inframarginals alone, and not so pronouncedly as in Mesopalseaster.
The disk is also more primitive in its construction, in that the central
50601 Bull. 8815 7
96 BULLETIN
disk plate is encircled by a ring of but 5 small accessory disk pieces,
separating the former from the next ring of 10 much larger plates
that are the primary pieces of the radial and supramarginal columns.
Genoliolotype and only species. Ccelaster latiscutatus Sandberger.
Restricted to the Lower Devonic of Germany.
SPANIASTER LATISCUTATUS (Sandberger).
Plate 12, figs. 1, 2; text fig. 8.
Ccelaster latiscutatus SANDBERGER, Verst. d. rheinischen Schicht. Nassau, 1855,
p. 381, pi. 35, figs. 1, la.
Xenaster simplex SIMONOVITSCH, Sitz. d. mat.-naturw. Classe Akad. Wiss., Wien,
vol. 64, Abt. I, 1871, p. 97, pi. 3, figs. 1, la.
Spaniaster latiscutatus SCHONDORF, Palseontographica, vol. 56, 1909, pp. 73-82,
109, pi. 8, fig. 2; pi. 10, figs. 2, 9, 10; pi. 11, fig. 9 (complete synonymy given
here); Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 31, pi.
5, figs. 1-3, text fig. 2.
The Sandberger specimen, showing only the abactinal side, was
found in the Lower Devonic Spiriferensandstein at Unkel, near
Bonn, Germany. Another specimen from the Lower Coblenzian
quartzite has been recently found at Bienhorntale, near Coblenz;
this shows both sides. Still another is from the Lower Coblenzian
at Oberstadtfeld in the Eifel. The holotype of X. simplex is from the
Upper Coblenzian at Niederlahnstein on the Rhine.
Genus MIOMASTER Sehondorf.
Plate 8, fig. 3.
Miomaster drevermanni SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 62, 1909, p. 38, pi. 3, fig. 4; pi. 4, fig. 3.
Remarks. This genus is most closely related to Spaniaster, in
that both have the single axillary interbrachial plates and the infra-
marginals and supramarginals are directly superposed. They differ
from one another mainly on the abactinal side, though this area is so
poorly preserved in Miomaster that almost nothing definite can be said.
Sehondorf illustrates a section through .the rays that shows the
presence of single columns of very small accessory pieces between
the radialia and supramarginalia. In the description, however, he
states: "One can not positively determine whether the supra-
marginal plates are separated from the median pieces by small
accessory ossicles, or whether the three dorsal columns were closely
adjoining." The central disk area is also too much disturbed to make
out the skeletal arrangement. There appear to have been small
interbrachial abactinal areas. Under these circumstances, the genus
Miomaster can not be said to be well established, and its final dispo-
sition is dependent on finding better preserved material.
Genoholotype and only species. M. drevermanni Sehondorf (as
cited above). A single example was found by Drevermann in the
Upper Coblenzian at Miellen-on-the-Lahn, Germany. The holotype
is in the Senckenberg Museum, Frankfort-on-the-Main.
REVISION OF PALEOZOIC STELLEROIDEA. 97
DEVONASTER, new genus.
Plates 11 and 12.
Palseaster (part) of AUTHORS.
Xenaster (?) or new genus GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 346.
Devonaster SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914, p. 14.
A characteristic starfish of the American Devonic.
Generic characters. Rays five, stout, tapering rapidly. Disk
large, with small interbrachial arcs.
Abactinal area of rays with prominent columns of radial supra-
marginals and less prominent inframarginal plates, all of which
bear numerous small pustules. The radial columns are completely
separated from the supramarginals by a great abundance of very
small, conical, irregularly arranged, accessory plates which in the
proximal region may also force apart single plates of these columns.
The supramarginal columns of adjoining rays nearly meet in the
axillary areas a short distance inside of the margin, at which level
the radial columns also cease. Inside of the area bounded by the
proximal plates of the radial and supramarginal columns, the disk
has numerous small accessory plates like those of the rays. In the
center of the disk is a plate somewhat more prominent than those
surrounding it, and just inside of the adjoining proximal supramar-
ginals is a small but distinct plate interradial in position.
Ambital accessory plates abundantly developed between the
marginal ossicles in the axillary areas. These plates are like the
other accessory plates and do not extend beyond the third or fourth
proximal inframarginal pieces, where the ambital areas cease and
the supramarginal ossicles come to lie more and more completely
over the inframarginals.
Madreporite flat or concave, situated between two proximal
supramarginal ossicles and outside of one of the small interradially
situated plates.
Inframarginal pieces lajge and thick, increasing rapidly in width
proximally, so that the tw.o wedge-shaped proximal plates are nearly
three times as wide as long. The plates are pustulose, and the proxi-
mal ones bear several slender spines.
Adambulacral plates more numerous than the inframarginals,
continuing around the axillary interbrachial plates, in front of
which the columns meet in two triangular and enlarged adambu-
lacral oral armature plates. Each piece bears two short, obtusely
pointed spines.
Ambulacral grooves narrow and deep, either straight or very slightly
petaloid. Ambulacral plates equaling in number the adambulacrals
and arranged in directly opposed, closely adjoining columns. Podial
openings in two columns through the sutures in the lateral part of
the furrows.
98 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Axillary interbrachial plates large, a single one situated in each
area between the adambulacral and inframarginal columns. Each
bears short, thick, blunt spines.
Mouth plates (tori) minute, one situated in front of each pair of
oral armature plates.
Genoholotype. Palseaster eucharis Hall.
Distribution. Restricted to the Hamilton and Chemung of the
American Devonic. The species are:
D. eucharis (Hall). Hamilton.
D. chemungensis, new species. Chemung.
Remarks. This genus is readily distinguished from Hudsonaster
hi having well-developed radial and ambital accessory plates, in
the numerous very small plates of the disk and in the presence of
interbrachial axillary plates.
Devonaster differs from Palseaster in having radial columns of
plates and in the presence of three plates in each interbrachial area
against one in Palseaster. There are also other differences. From
Neopalseaster it differs in having ambital accessory plates and no
ocular plates; while the proximal supramarginals and radials are
very large in the former, they are small in Devonaster, particularly
the supramarginals, which are not readily distinguished from the
accessory disk plates. In Mesopalseaster and Promopalseaster the
accessory plates are always arranged in columns or rows and the
ambital accessory plates are not restricted to the axillary areas as
hi Devonaster. Further, in those genera (excepting in the smaller
forms of Mesopalzeaster) there are always from two to five inter-
brachial marginal plates, while Devonaster has but one axillary
marginal in each interbrachial area.
Devonaster greatly resembles abactinally Xenaster of the Lower
Devonic of Germany, but is readily distinguished actinally in that
the latter has more large interbrachial plates, and especially in the ad-
dition of numerous accessory interbrachial pieces. These differences
are pointed out in more detail in the remarks on Xenaster.
DEVONASTER EUCHARIS (Hall).
Plate 12, figs. 3-5.
Palseaster eucharis HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p.
287, pi. 9, figs. 3-3a (74); rev. ed., 1868=1870, p. 330, pi. 9, figs. 3-3a (74).
QUENSTEDT, Petrefactenkunde Deutschlands, vol. 4, 1876, p. 71, pi. 92,
fig. 29. ZITTEL, Handb. Pal., vol. 1, 1879, p. 452, fig. 323. COLE, Bull.
Geol. Soc. America, vol. 3, 1892, p. 512, pi. 15. CLARKE, Bull. N. Y. State
Mus., No. 158, 1912, pp. 44-45, 6 pis.
Xenaster eucharis SCHONDORP, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol.
66, 1913, pp. 87-96, pi. 3, figs. 1, 2, text figs. 1-3.
Original description. " Body rather large; the largest individual
being one inch and seven-eighths from the center of the body to the
extremities of the rays; the whole having a robust aspect; rays
REVISION OF PALEOZOIC STELLEROIDEA. 99
acutely pointed at the extremity. Upper [abactinal] surface of
rays composed of three [radial and two supramarginal] ranges of
large, highly convex or tuberculiform plates which are nearly cir-
cular at the bases of the rays, becoming quadrate and widened to-
wards the extremities; separated from each other in the lower part
by numerous minute [accessory] plates or granules, which become
fewer near the middle of the ray, and before reaching the extremity.
The central portion of the disk is occupied by an elevated pentagon,
the angles of which are formed by the abrupt termination of the
central row of plates of each ray: the whole composed of very mi-
nute, highly convex plates, which vary in size, the larger ones pen-
tagonally arranged. The angles between the rays have a few [nu-
merous] small [accessory ambital] plates outside of the outer ranges
of tuberculose [supramarginal] plates on the upper side, uniting
with the inframarginal plates below [and not extending beyond
the third or fourth inframarginal proximal plate]. Madreporiform
tubercle distinct, situated lateraUy at the bases of the outer [supra-
marginal] range of large plates of two adjacent rays. Ventral sur-
face having deep ambulacral grooves, bordered by two ranges of
strongly tuberculose plates; the outer [infra] marginal range con-
sisting of 27 or 28 plates, besides a large, round [or elliptical], ter-
minal or axillary [interbrachial] plate; the others are wider than
long in the basal portion of the ray, becoming gradually shorter
towards the extremity where they are rounded. All the [infra]
marginal plates are visible from the upper [abactinal] side, and
usually appear as an additional range of plates on each margin of
the ray, making five with the three properly belonging to the upper
surface. Those of the inner range bordering the ambulacra (adam-
bulacral plates) are smaller than the marginal plates, about 38 to
40 in number; the basal or oral plates [oral armature] are triangular,
those of the adjacent rays uniting by their longer margins; and with
a single minute plate situated at these points. The plates of the
exterior surface, both upper and lower, present a granulose or stri-
ato-granulose surface which appears to have been produced by short
setae or spines; and at the angles of the rays the marginal plates
are armed by a few spines, which are as long or longer than the
transverse diameter of the plates. Ambulacra composed of a double
range of short, broad poral plates (ossicula), equal in number to
the adambulacral plates ; their outer ends excavated on the posterior
border, forming a comparatively large pore, just within its junction
with the adambulacral plate. There appears to have been [is] but
one range of pores in each set of ossicula, but these are large, dis-
tinct, and pass between [through and not between] the plates."
The following additional description is that of Mr. Cole. The
surface of the axillary interbrachial plate "is granulose and bears
100 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
three short, thick, blunt pointed spines. The [infra] marginal plates
bordering each reentrant angle bear similar but more slender spines,
which are not 'as long as the transverse diameter of the plates.'
The spines are arranged in a row near the distal margin of the plates
and number five on the plates at the angle, the number and size
decreasing until they disappear at the sixth or seventh plate from
the angle. All the [infra] marginal plates are nearly smooth on the
free margin and become gradually more granulose toward the line
of junction with the adambulacral plates. * * *
"The adambulacral plates are apparently less numerous than
stated in the original description, and l the single minute plate
[mouth plates] ' at the points of the pairs of the oral plates [oral
armature] is visible in this specimen and is armed with two relatively
long, slender spines which are apparently but a part of the full
armature. The adambulacral plates, including the triangular oral
[armature] plates bear well-defined spines, which are shorter than
the diameter of the plates to which they are attached. Each plate
bears two spines so near to the distal margin that the impressions
of the short and obtusely pointed spines frequently bridge the
well-defined groove between the adjacent adambulacral plates and
terminate near the proximal margin of the next plate. The spines
decrease in size toward the end of the ray and a few plates show
only one spine. The plates of this range are thick, equaling two-
thirds to three-fourths the depth of the groove. The vertical angle
of the faces forming the lateral walls of the groove are beveled, so
that lateral extensions of the groove are formed between each two
plates on the same side. These lateral expansions are narrow and
shallow at the oral surface, deeper and wider inward; so that the
faces of the adambulacral plates near their junction with the poral
[ambulacral] plates are reduced to a narrow edge which projects
inward and nearly touches the corresponding plate on the other
side of the groove. The general appearance of the fossil as well as
the outline of the rays at the points where the broken block presents
a transverse section of them indicates that the plates have their
normal position, not having suffered distortion by pressure.
"The ambulacral plates are shown by a well-defined mold of their
under or external surface. The soft matrix which filled the ambula-
cral furrow pressed upon the membranes connecting the ambulacral
plates and occupying their pores, and as these membranes decayed
it was forced by gentle pressure into the pores and between the
edges of the plates. The mold of the groove is less than one-eighth
of an inch in width in a ray measuring five-eighths at its base. The
upper surface of the mold bears a narrow longitudinal median ridge
which marks the junction of the two ranges of ambulacral plates.
Similar transverse ridges, which are continuous with the lines mark-
REVISION OF PALEOZOIC STELLEROIDEA. 101
ing the junction of the inner faces of the adambulacral plates, mark
the proximal and distal margins of the ambulacral plates. These
ridges do not cross at right angles to the median line, but include
between their proximal sides an angle of about 125. These ridges
indicate that the ambulacral and adambulacral plates were equal
in number, and that the former were united in pairs along a straight
median line rather than in an alternate right and left arrangement
along a zigzag line, as is shown in Dr. Hall's figures. The pores
described as being ' excavated in the posterior border of the ambu-
lacral plates and just within their junction with the adambulacral
plates' are not clearly shown on this specimen, although there are
irregular and inconstant markings at some of the points of the molds
of the lateral extensions of the groove. A series of pores near the
median line is indicated by a series of small rounded prominences
on each side of the median ridge and very close to it. These promi-
nences are opposite the lateral expansions of the groove, and one is
found on the mold of each ambulacral plate. The pores appear to
have perforations very near the edges of the plates, or excavations
in their margins."
Abnormal development. Among the 400 specimens of this species
recently found near Saugerties, New York, there are a few indi-
viduals each with but four rays, though otherwise they appear to
be of normal development. This is the first discovery of a four-
rayed starfish in the Paleozoic.
Formation and locality. In the Hamilton of the Middle Devonic
near Hamilton, Summit and Cooperstown, New York, Two sepa-
rated rays of apparently this species were found by the writer in the
lower third of the Hamilton near Bartletts Mills, south of Thedford,
Ontario. Two other fragments from the same locality are in the
University of Toronto (Walker collection, No. 1610H). The type
and other specimens are in the American Museum of Natural History,
one* is in Colgate University, another at Wesleyan University, and two
in Yale University Museum.
Recently the New York State Survey collected in the Hamilton
sandstone at Mount Marion, near Saugerties, New York, over 400
examples of this fine starfish. They occur in a limited area and
are often found in association with Grammysia and in such manner
that Doctor Clarke believes the starfishes were feeding on the bivalves.
This is probably the most remarkable find of Paleozoic starfishes, and
is certainly so for America.
DEVONASTER CHEMUNGENSIS, new species.
Plate 11, fig. 2.
Of this species only the actinal side is known, and its general
structure so far as can be made out is that of D. eucharis (Hall).
102 BULLETIN
It differs from the latter in that each axillary interbrachial plate
bears one stout spine, while D. eucharis has three. Further, in
D. eucharis six or seven of the proximal inframarginal plates of
each column bear on their outer side a number of small, slender
spines. In D. chemungensis these spines may also be present but
in addition each plate bears centrally a prominent tubercle for the
articulation of a large spine.
Formation and locality. The type is in the Museum of Columbia
University, New York City (No. 6228G). It is from the Spirifer
disjunctus sandstone of the Chemung of " Central Pennsylvania."
NEW GENUS.
NEW SPECIES.
In the University of Chicago collection there is a specimen (No.
14397) collected by Professor Weller in the St. Louis limestone on
Fountain Creek, at Waterloo, Illinois. This five-rayed asterid is
small, shows only the abactinal side, and has very long but narrow
marginals and not over six or possibly seven in a column; the basal
plates of adjoining columns make the axils. Otherwise the abactinal
skeleton of the rays consists of many small, loosely adjoining pieces
whose arrangement can not be made out. Nor can the disk struc-
ture be determined.
The relationship of this specimen seems to be with the Promo-
palseasteridse, with possible nearest affinity to the genus Mesopa-
Ixaster.
Measurements: R = not less than 10 mm., r = 4 mm.
I'liODVJIOI'-A.ILi^E^STElRIlSr^B, new subfamily.
Progressive Promopalseasteridse with the interbrachial areas com-
plex and composed of the single axillary interbrachials, more than
one pair of interbrachial marginals, and adambulacral plates. Podial
openings in each ambulacral furrow mainly in two columns, Jbut
proximally there may be two additional ones in a length never
more than one-third the rays.
Contains :
Promopalseaster, new genus.
PROMOPAL^EASTER, new genus.
Plates 13 to 23, 25.
Palxaster (part) of AUTHORS.
Promopal&aster SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April,
1914, p. 24.
Promos, chief, and aster. The largest and chief starfishes of the
American Ordovic.
Generic characters. Disk above medium size, with distinct and
angular interbrachial arcs. Rays five, slender to stout, more or less
long, and tapering. Some species attain a large growth.
BEVISION OF PALEOZOIC STELLEROIDEA. 103
Abactinal area of rays with numerous columns of small, more or
less tumid, closely adjoining or reticulated, spine-bearing plates. A
medial radial and two lateral or supramarginal columns of plates are
usually more conspicuous than the accessory columns; however, the
radial plates may be also inconspicuous. The longitudinal arrange-
ment in columns is usually most pronounced, but sometimes the
plates on each side of the radial column may have a decided diagonal
arrangement. Small accessory plates are usually inserted between
the columns of ossicles or between the individual pieces both distally
and proximally. Disk with numerous small accessory plates like
those of the rays, with no apparent definite arrangement except
the supramarginal columns, which continue over the disk and unite
angularly in the axillary areas. The larger plates bear several
short articulating spines of which the one on the apex is the largest.
Ambital areas well developed, there being one or more columns
of small plates like the radial accessory columns both in form and
arrangement.
Madreporite usually conspicuous, radially striated, and situated
near the margin of the disk between the adjoining columns of supra-
marginal plates.
Inframarginal plates small and numerous distally like the adambu-
lacrals, but proximally they usually increase rapidly in width and
assist in forming the small interbrachial arcs. Each ossicle has
numerous small granules which probably bore short, smooth, articu-
lating, blunt spines.
Adambulacral plates distally like the inframarginals, increasing in
width proximally and continuing around the plates of the inter-
brachial areas. The two proximal plates of adjoining columns are
the paii^s of plates in the oral armature. Each adambulacral plate
has a more or less well-developed narrow ambiilacral extension which
unites with the carina of the ambulacral plates. These extensions
belong to the adambulacral plates, apparently always so distally, but
proximally they are often separated from the adambulacral plates
by sutures ; throughout the greater portion of the rays they are situ-
ated medially, but in the proximal region they are either on the distal
or proximal edge, each alternate plate having the extension from
the same corner a little longer. Each plate has more or less numerous
spine-bearing granules in addition to the three larger articulating
spines which are inserted one on the ambulacral and two on the lateral
edge.
Ambulacral furrows generally very wide and regularly tapering.
In each ray there are two columns of these plates which are slightly
alternating or opposite. The plates are as numerous as those of the
adambulacral columns, highly carinated, about as wide as long in the
extreme distal portion of a ray, increasing rapidly in width proximally
104 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and at the base of the rays may be four times as wide as long. The
podial openings, one to each plate and in single ranges, are at the
extreme lateral edges excavated between the sutures of adjoining
plates and beneath the adambulacrals. Proximally, however, every
other podial opening progresses inwardly with the joined carinae and
issues where these begin to fork. Here there are, therefore, four
columns of podial openings in each ambulacral furrow. This change
is indicated by the changed position of the adambulacral plate exten-
sions and also b} 7 the pairs of forked carinse of the ambulacral plates.
Throughout the greater portion of a ray the carinse are regular and
alike on each plate, but toward the mouth they change rapidly in
direction and soon they are arranged in forked parrs, one curving
distally, the other proximally, with the lateral portions of each pair
hi contact and uniting with the extensions of the adambulacral
plates. The most proximal plate of each ambulacral column is
usually considerably modified, longer than wide, and more or less
triangular in outline, between which there is sometimes inserted a
small quadrangular ossicle. These pieces belong to the oral
armature.
Interbrachial areas of medium size, with the interbrachial marginal
plates usually arranged in pairs but in some forms the series may be
terminated by single ossicles. The number of these plates in an area
varies in different species, there being two, three, or five inside the
marginal inframarginals, and all seem to be derived from the
inframarginal series by inward crowding.
Genoholotype. Palseaster granulosus Meek (not Hall = P. speciosus
Meek).
Distribution. Restricted to the Middle and Upper Ordovicic of
America, chiefly within a radius of 50 miles about Cincinnati, Ohio.
The following are the known species:
P. wilsoni (Raymond). Black River (Lowville).
P. prenuntius, new species. Trenton.
P. sp. undet. ?Young of P. speciosus Meek. Maysvillian.
P. speciosus (Meek). Maysvillian.
P. granulosus (Hall). Lower Richmond.
P. bettulus, new species. Richmond.
P. spinulosus (Miller and Dyer). Richmond.
P. exculptus (Miller). Richmond.
P. wylcqffi (Miller and Gurley). Richmond.
P. dyeri (Meek). Maysvillian.
P. magnificus (Miller). Maysvillian and Richmond.
Remarks. Eight of the ten species here referred to Promopalseaster
have been described as Palxaster. They have little direct relationship
with the latter genus in that the species are much larger, have well
developed ambital areas, numerous accessory plates, well defined
REVISION OF PALEOZOIC STELLEEOIDEA. 105
interbrachial arcs, and from two to five interbrachial marginal plates
in each interbrachial area. Promopalseaster is a direct development
from Mesopalseaster, in that it preserves the generic characters of the
latter, which is in turn a descendant of Hudsonaster. In Hudsonaster
there are no interbrachial plates, accessory or ambit al columns. In
Mesopalseaster the single axillan^ marginal plate of Hudsonaster has
been crowded orally and is no longer a marginal but an axillary
interbrachial. Further, the radial and both marginal columns are
more or less completely separated from one another by accessory
columns of smaller plates. In Promopalseaster this addition of
columns of small plates is carried to its greatest development. The
character, however, which is relied upon to distinguish it from Meso-
palseaster is that Promopalseaster always has two or more interbrachial
marginal plates in each interbrachial area, while the former genus
has one, the axillary interbrachial, and Hudsonaster has none. The
position and number of axillary and interbrachial marginal plates
will readily distinguish these genera, but if the abactinal area of a
new form alone is seen it will be difficult to determine whether it is a
Mesopalseaster or Promopalseaster. Small size and few accessory
columns, especially ambital, will help somewhat to distinguish
Mesopalseaster from Promopalseaster.
Accepting Promopalseaster as a descendant of Mesopalseaster, it is
seen that the axillary interbrachial plates of the latter have in P.
exculptus been followed by two proximal pieces of adjoining infra-
marginal columns. The same is probably true of P. spinulosus, but
here no axillary interbrachial has been observed. In P. speciosus
and P. bellulus, another pair of proximal inframarginals have become
interbrachial plates, but the axillary interbrachial is also unknown
in this species. In P. dyeri and P. magnificus the axillary inter-
brachial is present along with two pairs of interbrachial marginals.
In other words, there is in Promopalseaster a constant increase in the
size of the interbrachial area produced by the crowding of pairs of
proximal inframarginal plates.
P. speciosus and P. bellulus are closely related species, and so far
as can be determined have but two interbrachial marginal plates,
while our theoretic development requires three. There is, however,
in each area orad to the interbrachial plates a small hiatus which in
some cases appears to be filled by extensions from the adambulacral
plates. It seems therefore probable that the axillary interbrachial
plate may be retained, but is not now discernible on account of its
small size or adhering clay; it may also have dropped out or have been
absorbed and its place occupied by the adjoining adambulacral plates.
In a general way it can be stated that there is also a regular increase
in the number of accessory plates from Mesopalseaster to Promopalse-
aster through P. spinulosus, P. speciosus, P. bellulus, P. dyeri into
106 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
P. magnijicus. It seems probable from the mature material studied
that the most differentiated species of Promopalseaster, P. magnificus,
passed through ontogenetic stages comparable to Hudsonaster, Meso-
palseaster granti, and Promopalxaster bellulus.
P. magnificus at present stands alone in its beautiful diagonal
and longitudinal arrangement of abactinal plates.
PROMOPAL^ASTER WILSONI (Raymond).
Plate 13, figs. 1, 2.
Palseasterf wilsoni RAYMOND, Ottawa Naturalist, vol. 26, 1912, p, 77, pi. 5, figs.
1-1.
Original description. " The specimen is exposed from the abactinal
side, and preserves the greater part of one arm, the disk, and the
stumps of the other four arms. The diameter of the specimen,
when complete, must have been about 75 mm. (3 inches), and the
diameter of the disk is 20 mm. This is large for a starfish from the
lower Ordovician. The arms are quite convex, with a gentle taper,
reminding one somewhat of the common recent starfish, Asterias
vulgaris, and as in that species, the arms were probably somewhat
flexible. The greater part of the abactinal side of the disk and arms
is covered with small convex, overlapping, V-shaped plates, which
are arranged with the point of the V directed toward the margins.
Along the crest of each arm there is a single row of larger plates
(radial column). These plates are quite large and hexagonal in out-
line near the disk, but become smaller, triangular, and alternate in
position farther out on the arm. There are two rows of marginals
[supra and infra] these plates being larger and flatter than the other
plates, and covered with minute tubercles, which may be spine bases.
Close to the disk, the supramarginals and [infra] marginals seem to be
of the same size, both rectangular, and the plates' of the supra-
marginal row directly over those of the marginal series. Farther
out on the arms, the plates are pentagonal, those of the two rows
alternating in position, and dove-tailing, and the supramarginals
are smaller than the marginals. One of the marginals, about half-
way out on the arm, is 1.25 mm. high and of about the same breadth.
The smaller triangular plates which cover the greater part of the
arm average about 0.5 to 0.6 mm. in height. On one of the arms,
the small triangular plates seem to be arranged in rows parallel to the
axis of the arm, but the plates on the longer arm seem to be more
irregular, although a general arrangement in rows can be seen. On
this arm there are a number of very small plates scattered about,
especially on the top of the arm, thus adding to the irregularity. The
triangular shape of these plates gives the arm a neat pattern, the plates
making diagonal rows backward and forward from the row of large
plates along the top of the arm. The madreporite, which is nearly
circular in outline, and 2 mm. in diameter, is in position, but slightly
BEVTSION OF PALEOZOIC STELLEROIDEA. 107
tipped down at the inner side, in an interradius, and not far from the
center of the abactinal side of the disk. The surface is probably worn,
for it appears perfectly smooth."
In places the abactinal ossicles are removed, "disclosing the ambu-
lacral plates. These plates, which are long and rather thick, seem
to be alternate in position. Two of the plates seem to be pierced by
pores near their proximate ends, two pores piercing each plate ver-
tically. Near the outer end of the more perfect arm there is a space
where a few of the small triangular plates are missing, and here also
the ambulacral plates can be seen from the upper side. Each plate
has a narrow keel on that side. Other details of the plates of the
actinal side are unknown."
Formation and locality. In the Lowville limestone at City View,
near Ottawa, Canada, where this interesting asterid was found by
Miss A. E. Wilson, in whose collection the holotype now is.
Remarks. It is interesting to note that this fully developed
Promopalseaster, one closely related to P. magniiicus of the highest
Ordovicic, occurs in much older rocks; that is, in the Black River
formation. This occurrence again brings out the fact often noted
by the writer, that Paleozoic asterids are slow to change in characters
accessible to the paleontologist.
P. magniiicus, like P. wilsoni, Raymond states, "has two rows
of large marginals and a row of large plates along the top of each
arm, while the greater part of the surface is covered with small
convex triangular plates. Miller found the plates to be spine-bearing,
a point which can not be definitely decided in the present species.
From the photograph (submitted by Schuchert) it appears that there
are spaces between the small triangular plates on the abactinal sur-
face in Pals&aster f magniiicus, while in P. ? wilsoni these plates actually
overlap each other."
PROMOPAL^EASTER PRENUNTIUS, new species.
Plate 13, fig. 3; plate 15, fig. 5.
Remarks. This species is represented by a good specimen showing
the actinal side, the other adhering to the limestone. R = somewhat
less than 30 mm., r = 7 mm. Width of ray at base about 9 mm.
P. prenuntius is closely related to P. speciosus, but is distinguished
therefrom by the smaller size, less pustulose ornamentation of the
plates, and the lower position in the geological column. The plates
of the axial area are also more regular in size and there are in the
proximal region nearly always two adambulacrals to each inframar-
ginal; at about mid-length of the rays the plates of each of the two
columns are about equal in number, but every now and then there
is some duplication among the adambulacrals. P. prenuntius is
equally as spinose as P. speciosus and the former is undoubtedly the
ancestor of the latter.
108 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Formation and locality. The holotype was collected by Mr. Wendt
near Frankfort, Kentucky, apparently in the Lower Trenton. The
writer's attention was called to it by Prof. Arthur M. Miller, who
has it in his charge at the State University of Kentucky at Lexington.
PROMOPAL^ASTER, species undetermined.
Plate 14, figs. 1, 2.
Asterias primordialis ANONYMOUS, two lithographic figures sent out by the Western
Academy of Sciences previous to 1872 without description or known author.
Nothing more is known of this species than the two good original
illustrations reproduced here. These figures were sent to F. B. Meek
on May 18, 1872, by Mr. D. H. Shaffer, of Cincinnati, Ohio, with the
following comment (they were found in the Meek correspondence
kept in the United States National Museum): "The lithograph in-
closed of this Asterias was found here, and in the possession of
Joseph Clark (deceased). I saw it, and this is a faithful picture of it.
I think the fossil is either in the possession of his nephew or in the
cabinet of Maxwell's Female Seminary, which now is the owner of
Mr. Clark's cabinet. I think it is worthy a place with Palseaster
shafferi in the illustrations of the Ohio Geological Survey."
From the illustration one sees that this form had on the abactinal
side of the rays about nine columns of ossicles. Of these the radial,
supra- and inframarginals are largest. Between the marginals there
is a single column of tiny ambital pieces, while on each side of the
radials are two columns of small accessory pieces. The structure of
the disk pieces can not be made out.
The interbrachial areas are small and do not appear to have more
than two inwardly crowded inframarginals, but there may also be
present single very small axillary pieces.
The adambulacral and inframarginal columns appear to have not
more than 20 ossicles hi a column. Of ambulacrals there are some-
what more, about 25, and these alternate slightly with one another.
"Asterias primordialis" appears to be the young of Promopalse-
aster speciosus (Meek), and the differences can all be explained as due
to incomplete development; that is, the former has fewer ossicles and
fewer secondary columns of plates.
Formation and locality. From the hills of Cincinnati, in the Mays-
ville formation. The present whereabouts of this fine specimen is
unknown. A still smaller and younger specimen is in the Faber
collection of the University of Chicago (No. 9567), and was collected
at 350 feet above the Ohio River at Cincinnati. It has but a single
axillary plate in each interbrachial area, and about 15 adambulacrals
in a column. It is interesting to note that in these young individuals
of Promopalxaster, the smaller they are the more they approach
Mesopalseaster and suggest Hudsonaster.
EEVISIOST OF PALEOZOIC STELLEKOIDEA. 109
PROMOPAL^ASTER SPECIOSUS (Meek).
Plate 14, figs. 3, 4; plate 15, figs. 1-4.
Asterias antiquata LOCKE (not A. antiqua Troost), Proc. Acad. Nat. Sci., Phila-
delphia, for 1846-47, vol. 3, 1848, p. 32, fig. on p. 33.
Palseaster antiquata HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868,
p. 286.
Palssaster antiquatus HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., rev. ed.,
1868=1870, p. 328.
Palseaster granulosus MEEK (not Hall 1868), Amer. Journ. Sci., ser. 3, vol. 4, 1872,
p. 276; Geol. Surv. Ohio, Pal., vol. 1, 1873, p. 60, pi. 4, figs. 3a-c.
Palseaster speciosus MEEK, Amer. Journ. Sci., eer. 3, vol. 4, 1872, p. 277 (name at
end of description); Geol. Surv. Ohio, Pal., vol. 1, 1873, p. 61 (name at end
of description).
Also see Promopalxaster, sp. undet., page 108.
Original description by Meek (1873). " Attaining a large size;
rays (as a little depressed by accident) slightly more than twice as
long as their breadth at their inner ends ; and tapering regularly from
the disk to their free ends, which are rather acutely angular; breadth
of the disk a little less than that of the inner ends of the rays. Mar-
ginal pieces convex [bearing numerous small spines], nearly or quite
as long as wide, numbering on each side, in the whole length of each
ray (1.40 inch), 31. Adambulacral pieces, a size smaller, as long as
wide, convex, and numbering on each side 39 to 40; like those of the
[infra] marginal series, each bearing a small spine [several spines],
the largest of which are each about 0.14 inch in length and 0.02 inch
in thickness [between each two adambulacral plates just above the
podial opening is inserted a large and thick spine], Ambulacral
pieces very short, or nearly three times as wide as long, apparently
not alternating with the adambulacral pieces, and each provided
with a rather sharp ridge across nearly its entire breadth. Dorsal side
of disk and rays composed of small tuberculiform pieces, of which
about 16 rows may be counted at about half way between the inner
and outer ends of each ray, along the middle of which those of two
[always two] or three of the rows are a little larger than the others;
each dorsal piece bearing a minute, very short spine [they are nume-
rous and like those of the marginal plates]. Surface of all the pieces
minutely granular, one of the central granules always being somewhat
larger than the others for the articulation of a spine [all of these
granules bear spines]. Oral pieces and madreporiform body un-
known."
Emended description. Rays regularly tapering. R = 44 mm.,
r= 11 mm., R = 4r. Width of ray at base 14 mm.
Abactinal area of rays covered by small, highly convex, tuber-
culated plates, each bearing numerous short slender spines. These
ossicles are arranged in longitudinal columns, of which there a?e
15 on each ray at about its mid-length. The plates of one or
110 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
two median columns are larger and more convex than the three
columns immediately on each side, while the pieces of the fourth
column are again more pronounced than those on each side of it, and
probably represent the supramarginals. Between the supramarginals
and the inframarginals are from one to three columns of ambital
plates, and between these are inserted here and there some small
extra pieces. The extra pieces are mainly developed beside the supra-
marginal columns. Plates of the disk like those of the rays, but their
arrangement is unknown.
Madreporite unknown.
Inframarginal plates very convex, about as wide as long except-
ing near the base of the columns where they are about twice as wide
as long. There are about 31 of these ossicles in a column, and they
have numerous small, slender, sharp spines articulating on well-
developed tubercles.
Adambulacral plates greatly resembling the inframarginals.
They are largest near the mid-length of each ray, diminish in size
distally, but proximally they hold their width and decrease in length
and therefore are more numerous. There are 43 plates in each
column besides those in the oral armature. Ambulacrally each
adambulacral plate has a short attenuate extension which articulates
with the crests of the ambulacral plates. Between all the adam-
bulacral ossicles there is inserted in the suture facing the ambulacral
groove a single large, thick spine, excepting for a short distance
proximally where such are inserted in every other suture. The
lateral surface of the adambulacral plates in the distal half of the
columns bears numerous small spines like those of the inframarginal
series, but proximally these spines are gradually displaced by two
or possibly more larger ones like those on the ambulacral face.
Ambulacral furrows wide and deep. Ambulacral plates of adjoin-
ing columns opposite, two or three times as wide as long, and slightly
overlapping proximally. Upon each plate there is a high median
ridge which arises at the inner end of the plate and abuts against
the short extension of the adambulacral plate. The podial open-
ings are between the plates and beneath the sutures of the adam-
bulacral plates, excepting near the base of the rays, where they
occur in every other suture. The ridges here are sigmoid and not
straight, as they are more distally. There appear never to be more
than two rows of podia in an ambulacrum.
In each axillary area inside of two large inframarginal plates and
between the converging adambulacral columns are inserted two
pairs of wedge-shaped interbrachial marginal plates ornamented
like the inframarginals.
Formation and locality. The holotype and only known specimen
was collected by Mr. Charles B. Dyer in the Maysvillian at Gin-
REVISION OF PALEOZOIC STELLEROIDEA. Ill
cinnati, Ohio. It is now in the Dyer collection, Museum of Com-
parative Zoology (No. 22), Harvard University. The original of
Astcrias antiquata Locke was found at Cincinnati, but the specimen
appears to be lost.
Remarks. Meek (1873) in describing this specimen, provisionally
referred it to Palseaster granulosus Hall and commented as follows:
"Although this fine Star-fish agrees pretty well in most of its
characters with the description of Palseaster granulosus of Hall (which
has not yet been figured) , I am far from being entirely satisfied that it
is really the same, as it seems to differ in some important respects.
For instance, the rays of P. granulosus are described as being 'obtusely
rounded at the extremities'; while in the form under consideration
they are rather acutely rounded, if not angular. Again, instead of
having 25 of the marginal pieces on each side, in a space of one inch
and a quarter from the apex of each ray, and 42 or 43 of the adam-
bulacral, it shows in this space 28 marginal and only 32 adambula-
cral pieces. The number of the latter being about 10 less on each
side in the same space a rather decided difference, showing the
inner row to consist of proportionally larger pieces. Its ambulacral
ossicula seem to have the same proportional breadth and length as in
the type of P. granulosus and also have each a similar ridge across
the middle; but these ridges do not show the zigzag arrangement
mentioned in the description of P. granulosus. Prof. Hall does not
say how many rows of pieces are seen on the dorsal sides of the rays
of his species; but he states that it is probably the same species that
the Western Academy of Sciences sent out lithographs of under
the name Asterias primordialis. One of these lithographs, now
before me [reproduced here on plate 14, figs. 1, 2], represents from
8 to 10 rows of these dorsal pieces, which is 6 or 8 less than may be
counted near the middle of the rays of our specimen.
"In noticing this form in the Am. Jour. Sci., vol. 3 (3d series),
p. 277, I proposed, in case it should be distinct from P. granulosus,
to call it P. speciosus. J)
With such marked differences between P. granulosus and P. speci-
osus, and since the type-specimen of the former can not be located
to determine the interbrachial structure, it is deemed advisable to
adopt Meek's name P. speciosus for the specimen described above.
In the upper portion of the Ordovicic in the Richmond formation
occurs another closely related species, P. ~bellulus, which is dis-
tinguished from P. speciosus in having more slender rays and usually
a greater number of plates in the actinal columns. The greatest
difference, however, is that it has seven pairs of adjoining adam-
bulacral plates proximal to the interbrachial plates, while in P.
speciosus there are but two or three pairs of adambulacrals. In
other words, the adambulacral oral extensions are far longer in
P. bellulus than in P. speciosus.
50601 Bull. 8815 8
112 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
The illustration of Asterias antiquata Locke referred to above by
Meek is very poor, and yet it seems to be the same species which
the latter described as P. granulosus Hall = P. speciosus Meek.
This, however, can not now be established since the whereabouts
of Locke's specimen is unknown. Both specimens are from Cin-
cinnati and are of about the same size. Hall comments on this
species (1870) as follows:
"This species was noticed by Dr. Locke, as cited above, but with-
out specific description, and expressing a doubt whether it was or
was not identical with the Asterias antiqua of Troost. The figure
would indicate a distinct species from that of Dr. Troost."
PROMOPAL^EASTER GRANULOSUS (Hall).
Palxaster granulosa HALL Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868,
p. 285.
Palseaster granulosus HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., rev. ed.,
1868=1870, p. 327.
Original description. "Body of medium size, five rayed; rays a
little more than twice as long as their breadth at base; obtusely
rounded at the extremities. Upper surface of rays composed of
numerous very small tuberculose or subspinose plates; the madre-
poric tubercle large, quite distinct, situated laterally at the base of
two of the rays. Under surface of rays composed of a [infra]marginal
range of small tuberculose plates, about 25 on each, side in a ray
measuring one inch and a quarter from base to apex; and an inner
(adambukicral) range of smaller plates, of which about 42 or 43 can
be counted on the same ray; the terminal or oral [armature] plates
are small, elongate, subtriangular, in pairs at the base of the adjacent
rays. Ambulacral areas composed of a double series of short, broad,
slightly curved poral plates (ossicula) , each plate marked by a sharply
elevated ridge along its entire breadth, commencing on the one plate
at the outer posterior angle and terminating on the anterior inner
angle, and running in the opposite direction on the adjacent plate.
When the outer ridged surface of the poral plate is ground away,
the narrow openings or pores are visible between the plates, apparently
in two rows in each series, making four ranges of pores in each ambu-
lacral area.' (The marginal ranges of pores are obscure, and may
only be apparent [there is probably an error here in regarding the
inner openings as podial openings].) On the under surface, near the
bases of the rays, the tubercles bear short spines some of which are
still in place."
' * Some figures of a Palssaster, closely allied to or identical with this
one, from Cincinnati, Ohio, have been circulated by the Natural
History Society of that place, under the name of Asterias primordialis;
but no description of it has ever been published, so far as I know, nor
EEVISION OF PALEOZOIC STELLEKOIDEA.
113
do I find it at all recognized in the catalogues." These original
figures are here reproduced as Promopalseaster, sp. undet. (pi. 14,
figs. 1, 2). The specimen is probably an immature individual of
P. speciosus.
Formation and locality. The holotype was found by Mr. J. Kelly
O'Neall in the Lower Richmond formation near Lebanon, Ohio, and
was loaned to Hall for description. The whereabouts of the specimen
can not now be located. Some years ago the writer asked the late
Professor Whitfield about it and he answered that Halt had returned
it to O'Neall, but the latter in turn wrote (Aug. 27, 1896) that he
had not received it.
Remarks. This species seems to be distinguished by the smaller
number of plates in the inframarginal columns, as the following table
shows. However, until the nature of the interbrachial areas is known,
the relationship of P. granulosus to other Paleozoic forms can not
be stated.
Inframar-
ginals.
Adambu-
lacrals.
Inter-
brachials.
Adambu-
lacrals
orad to
inter-
brachials.
P granulosus (Hall)
25
42-43
P speciosus (Meek) . *
31
43
4
4
P bellulus new species .
31-38
48-55
4
8
P. magnificus (Miller) .
45
50-52
5
5-6
PROMOPAL^ASTER BELLULUS, new species.
Plate 15, figs. 6-8; plate 16, fig. 1; plate 18, figs. 4, 5.
R = 36 mm., r= 10 mm., R = 3.6r. Width of ray at base, 11.5 mm.
A first view of this species gives the impression of P. speciosus
(Meek). Of the latter but a single specimen is known and it is now
lost, while of P. bellulus there are numerous examples, all of which
have the same specific characters. The specimens preserved in lime-
stone have the rays more slender, while those from soft shale are as
stout as in P. speciosus.
In the holotype of P. bellulus from limestone there are 55 adam-
bulacral and 38 inframarginal plates in each column, while in
P. speciosus there are 43 and 31 plates, respectively, in the same
columns. Shale specimens of P. lellulus have less plates in a column.
One of these, a somewhat smaller specimen than the holotype, has
31 inframarginal plates, the same number as in P. speciosus, and
about 48 adambulacrals in each column. P. bellulus has therefore
from 31 to 38 inframarginal plates and from 48 to 55 adambulacral
plates in each column.
114 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
A more important distinguishing character exists in the inter-
brachial areas. In P. lellulus and P. speciosus there are two pairs
or four inter brachial marginal plates, while the number of adjoining
adambulacral plates is greater. In the latter there are four of these
on each side, while in the former there are eight. In other words,
the interbrachial areas in P. lellulus are much more elongated orally
than in P. speciosus and the interbrachial arcs are also greater.
Abactinally on the rays the arrangement and number of columns
of plates appear to be very much like those in P. speciosus. The
individual plates are, however, less convex, cruciform, more like those
of P. spinulosuSj and have stronger and more abundant spines. The
spines in P. bellulus are very numerous, short, and slender, and are
inserted one on the apex of each plate and a number around the sides
in the angles between the radial extensions. In P. speciosus the spines
are grouped over the convex area of the plate and around the larger
central spine. The arrangement of the plates of the disk appears to
have no marked pattern and consists of very numerous small plates
like those of the rays.
The adambulacral plates each bear three spines as in P. speciosus
and increase in size toward the base of the rays. One is inserted on
the inner or ambulacral face of the plate and the two closely adjoining
ones immediately above and laterally on the rounded surface.
Ambulacral plates as in P. speciosus.
Madreporite of medium size, subcircular in outline, with numerous
straight, sharp, radiating ridges. It is inserted proximally between
two adjoining columns of supramarginal plates.
A development occurring only in this species of Promopalseaster
is a single plate oral to the proximal adambulacrals. This plate
has been seen in four axillary areas. It may be a modified ambu-
lacral and may correspond to a similar plate, which is an ambulacral,
hi P. exculptus.
Formation and locality. Of this species quite a number of speci-
mens were secured by Mr. Harris in the Waynesville beds of the
Richmondian. They are all in the United States National Museum.
One excellent specimen (the holotype) showing the actinal area,
and two smaller poorer specimens are attached to limestone, while
eleven fair specimens and fragments of eight other individuals are
from a soft blue shale associated with Zygospira modesta and Ptilo-
dictya sliafferi (upper form). All were secured in the vicinity of
Waynesville, Ohio. The shale specimens were exhumed from one
small area, showing that probably many starfishes could be similarly
secured by collectors, since these soft specimens in weathering out
are broken up and lost; more asterids will be found when they are
dug or quarried for. Another good specimen is in the Gurley col-
lection of the University of Chicago (No. 10983) and was found at
Versailles, Indiana.
REVISION OF PALEOZOIC STELLEEOIDEA. 115
Remarks. The similarity and difference between P. bellulus and P.
speciosus are pointed out above. Another related species is P. spinu-
losus, which has the same kind of abactinal plates and madreporite
as P. bellulus. In the former the rays are far longer and narrower,
with fewer ambital plates and columns between the supramarginals.
Actinally P. spinulosus is at once distinguished from P. bellulus
in the much smaller interbrachial areas, in that it does not have
more than two interbrachial marginal plates while P. bellulus has
four. In the latter species there are also far more adambulacral
plates around the interbrachial pieces.
It will always be difficult to distinguish fragments of P. bellulus,
P. speciosus, and P. spinulosus from one another, even with good
material, since at least two specimens of P. bellulus were identified
as Palseaster granulosus Hall = Promopalseaster speciosus (Meek) .
Cat. No. 40879, U.SJST.M.
PROMOPAL^EASTER SPINULOSUS (Miller and Dyer).
Plate 16, fig. 2; plate 17, figs. 1, 2; plate 18, figs. 1-3.
Compare with Promopalseaster exculptus (Miller).
Palseaster spinulosus MILLER and DYER, Journ. Cincinnati Soc. Nat. Hist., vol.
1, 1878, p. 32, pi. 2, figs. 12-126.
Palxaster longibrachiatus MILDER, Journ. Cincinnati Soc. Nat. Hist., vol. 1, 1878,
p. 102, pi. 3, fig. 4.
Original description (of Palseaster spinulosus). "Pentagonal;
rays longer than the diameter of the body; marginal plates globular
near the apices of the rays, but lengthened toward the junction
with the body. Six of these plates measure from the body on a ray
three- tenths of an inch. Two wedge-shaped plates form the junc-
tion of the marginal plates with the body.
" Adambulacral plates a little smaller [in width] than the margi-
nal pieces, and possessed of two or three [four or more] spines to
each plate. These spines taper to a fine point, and are longer than
the diameter of the plates. The oral pieces are not determined
[they are like those in P. exculptus]. The ambulacral plates have
their greatest length across the rays, and are possessed of a sharp
ridge in the middle which seems to be [is] connected at one end with
the adambulacral pieces. [The entire number of plates in each
range can not be given, since the rays are imperfect.]-
"The dorsal side is covered with plates united by angular exten-
sions. They are of unequal size, and strongly tuberculated or spi-
nous.
"The madreporiform tubercle on its upper face is an oblate sphe-
roid, much depressed, and marked by fine radiating stride, which
become more numerous by intercalation, without bifurcation.
[This plate is interpolated between the proximal plates of two ad-
116 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
joining columns of supramarginal ossicles and about medially be-
tween the margin and the center of the disk.]"
Supplementary description. II unknown, r=7 mm. Width of
ray at base 9 mm.
Abactinal area of rays margined by inframarginals, above and
inside of which are columns of large, bipyriform, multispinous, su-
pramarginal plates of which there are five in 5 mm. at the base of a
ray. The columns of adjoining rays unite in the axillary areas.
Inside of the supramarginal columns the rays and disk have numer-
ous smaller accessory plates, usually triangular in outline or vari-
ously stellate. They lie upon or against each other, each ossicle
bearing at least one small spine, and leave between them numerous
abactinal openings. Outside of the supramarginal plates in the
ambital areas are spicular ambit al plates which cover the abactinal
side of the inframarginals.
Description of the type of Palseaster longibrachiatus. 11 = 38 mm.,
r = 7 mm., R = 5.4r. Width of ray at base 9 mm.
Rays large, slender, tapering slowly; actinally somewhat convex
at their outer ends, but elsewhere concave.
Abactinal area unknown.
Inframarginal plates granular, large, decreasing gradually in size
dis tally, highly convex, in outline tetragonal or pentagonal and
common to both the abactinal and actinal areas. From 28 to 30
plates in a column on each side of a ray.
Axillary areas occupied by the proximal plates of adjoining infra-
marginal columns, and in the interbrachial areas there are two sub-
quadrangular or sub triangular interbrachial marginal plates. There
may be an additional small single plate in each area.
Adambulacral plates greatly resembling the inframarginal ossicles.
Distally they progressively overlap the inframarginal plates con-
siderably, while proximally they gradually increase in size and are
entirely inside and depressed beneath the plane of the marginal
columns. Orally the columns of adjoining rays unite in two wedge-
shaped modified adambulacrals (oral armature pieces) .
Ambulacral furrows deep and gradually tapering. Ambulacral
plates unknown.
Formation and locality. The type of Palseaster spinulosus (No. 16,
Mus. Comp. Zool.) is said to have been found at Cincinnati, Ohio.
This is probably an error, since il/s color and preservation indicate
the Richmond formation and that it comes from some locality
considerably to the north of Cincinnati. The only other known
specimen, the type of P. longibrachiatus, was found in the Richmond
formation near Clarksville, Ohio, and is in the Harris collection,
United States National Museum.
Remarks. The type-specimen of Palseaster spinulosus, which is
free, preserves but the disk and a small proximal portion of the rays.
REVISION OF PALEOZOIC STELLEROIDEA. 117
Comparing it with the actinal area of P. longibrachiatus Miller, it
is evident that both forms are identical. In the latter the rays are
complete, very long and slender, and there is nothing to show that
they were dissimilar in P. spinulosus Miller and Dyer. Both types
are nearly of the same size and the actinal plates are identical in form
and arrangement. The abactinal area in P. longibrachiatus is not
known, but since its actinal structure is that of Promopalseaster
spinulosus it is safe to assume that the abactinal structure is alike
in both.
P. spinulosus actinally is also closely related to P. exculptus (Miller).
The latter appears to have shorter and more rapidly tapering rays
and in general seems to be a more robust species. Another character
by which these species can be separated is that the interbrachial
areas in P. spinulosus have two, while P. exculptus has three inter-
brachial marginal plates. It is possible that the former species
also has three interbrachial marginal plates, but as yet the small
single piece of each area has not been observed, while in P. exculplus
it is distinctly present in each of the five areas. Should P. spin-
ulosus also prove to have three interbrachial plates, it would be diffi-
cult to distinguish it actinally from P. exculptus.
P. dyeri and P. magnificus are far larger species and have five
interbrachial marginal plates in each area.
Cat. No. 40881, U.S.N.M.
PROMOPAL-EASTER EXCULPTUS (Miller).
Plate 18, fig. 7; plate 20, fig. 2.
Compare with Promopalasaster spinulosus (Miller and Dyer).
Palseaster exculptus MILLER, Journ. Cincinnati Soc. Nat. Hist., vol.4, 1881, p. 69,
pi. 1, fig. 1.
Original description. " Pentagonal; rays a little longer than the
diameter of the body; diameter of the body, about 0.93 inch; length
of ray measuring to the center of the body or disk, about 1J inches;
breadth of a ray at the junction with the body, about 0.57 inch; rays
obtusely pointed.
"The [infra-] marginal range consists of somewhat quadrangular
plates, having a width a little greater than the length; the first 8
of these have a length of one-half inch, and there are about 18 in
the length of an inch, and not far from 25 in each range, though the
specimen does not permit us to make the count with certainty. The
surface is strongly tubercular, and was probably spinous [originally
covered with numerous small spines].
"The adambulacral range consists of about 28 [25] plates, on each
side of a ray; they are narrower than the marginal plates, but have
about the same length. Each plate [has a short ambulacral exten-
sion conforming with the ridge of the ambulacral plates and] bore
118 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
strong spines, and some of them, preserved on our specimen, have
a length greater than the length of a plate. * * *
"The ambulacral plates have their greatest length across the rays,
thus providing a wide ambulacral furrow. Each plate is furnished
with a sharp ridge in the middle, that curves slightly outward, from
the center toward the adambulacral range, increasing in height
until it approaches or abuts against the adambulacral plate. * * *
The appearance of having been carved out, which is presented by
the ambulacral plates, suggested the specific name. [The plates of
adjoining columns are opposite one another or slightly alternate.]
"The dorsal side and madreporiform tubercle unknown."
Supplementary description. R = 37 mm., *r=12 mm., R = 3.1r i .
The specimen is very much flattened and the exact width of a ray
can not be given but appears to have been about 10 mm. at the base.
In the axils of the rays the proximal inframarginal plates of adjoin-
ing rays rest against each other, inside of which are two wedge-shaped
interbrachial marginal plates. Orad to these and adjoining the pairs
of proximal adambulacrals (oral armature pieces) is situated a single
subquadrate ossicle, making three interbrachial marginal plates in
each interbrachial area.
The podial openings in the ambulacral grooves are between the
sutures of adjoining adambulacral and ambulacral plates along the
line where these two columns adjoin. The proximal ambulacral
plates orad to the axial adambulacrals are considerably modified
and are to be considered as parts of the oral armature. A single
rather large plate is here partially wedged between the pair of termi-
nating adambulacrals, and on each end of it abuts a single broadly
triangular plate, a modified ambulacral, completing the oral ends of
the ambulacral columns (see drawing, pi. 20, fig. 2).
Formation and locality. In the Richmond formation near Waynes-
ville, Ohio. The holotype and only specimen is in the Harris collec-
tion of the United States National Museum.
Remarks. This species is near P. spinulosus, but apparently had
wider, shorter, and more rapidly tapering rays. There appear to be
but two interbrachial marginal plates in P. s-pinulosus, while P.
exculptus has three in each area. Should the former species also prove
to have three plates, a possibility, P. exculptus will be actinally
identical with P. spinulosus. The wider and more rapidly tapering
rays of P. exculptus can then be explained by the fact that the type-
specimen is considerably distorted, since the adambulacrals and infra-
marginals now lie wholly outside of the ambulacrals, which are com-
pletely exposed, an unnatural condition. So long as the abacti-
nal area of P. exculptus remains unknown the species had best be
retained.
Cat. No. 60608, U.S.N.M.
REVISION OF PALEOZOIC STELLEROIDEA. 119
PROMOPAL^ASTER WYKOFFI (Miller and Gurley).
Plate 18, fig. 6; plate 19, fig. 2.
Palxaster wykoffi MILLER and GURLEY, Bull. No. 12, Illinois State Mus. Nat.
Hist., 1897, p. 46, pi. 3, fig. 27.
Original description. " Species medium size [R = 26 mm., r = 8 mm.
Width of ray at base 10 mm.] Rays pateliform; length about one
and a half tunes the diameter of the body; breadth of a ray at the
place of junction with the body, about two-thirds the diameter of
the body; obtusely pointed.
"Marginal [inframarginal] plates wider than long and numbering
about 15 in the length of half an inch from the body [or about 28
in a complete column]. The [infra] marginal plates curve regularly
around the tips of the rays. The adambulacral range curves around
the ends of the rays within the marginal plates and consists of sub-
quadrate plates, wider than long. They are a little shorter than the
marginal plates, so that there are about 18 in the length of half an
inch [about 35 in a complete column]. There are 10 oral plates [oral
armature] at the junction of the adambulacral rows, which present
triangular extensions toward the center of the ventral cavity. A
single irregular axillary plate rests between the terminal marginal
plates and the angle formed at the junction of the adambulacral
plates [this description is inadequate, see beyond for the detail].
The ambulacral plates have their greatest length across the rays,
thus providing a wide ambulacral furrow. Each plate is furnished
with a sharp ridge in the middle, extending from the middle furrow
to the adambulacral plates.
"Our specimen exposes only the ventral side and no spines are
preserved. It is a well-marked and beautiful species."
Formation and locality. Found in the Richmond formation "near
Madison, Indiana, by Charles W. Wykoff, in whose honor we have
proposed the specific name." The holotype is now in the Gurley
coUection of the University of Chicago (No. 6066).
Remarks. Each interbrachial area begins orally with a pair of
elongate triangular basal adambulacrals (the oral armature pieces)
dis tally to which there are two pairs of adjoining adambulacrals
followed by a single axillary ossicle and usually one but also more
or less of a second pair of interbrachial marginal plates. P. wylcoffi
is therefore distinguished from P. exculptus in that it has orally to
the single interbrachial plate two pairs of proximal adambulacrals
instead of one as in the last-named species. These differences are
not decided, and it may eventually be seen when more well-preserved
material is at hand that they are either individual variations or due
to differences in age.
120 BULLETIN" 88, UNITED STATES NATIONAL MUSEUM.
P. wykoffi also greatly resembles Anorfhaster miamiensis, but the
interbrachial areas in the latter are entirely composed of adambu-
lacral plates, there being no interbrachial marginals present. While
both species attained a similar size, another -difference is that the
former has a greater number of plates in the adambulacral and
inframarginal columns.
PROMOPAL^EASTER DYERI (Meek).
Plate 18, fig. 8; plate 20, figs. 3-6; plate 25, fig. 1.
Palseaster dyeri MEEK, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 257; Geol. Surv.
Ohio, Pal., vol. 1, 1873, p. 58, pi. 4, figs. 2a-2/.
Original description (1873). "Among the specimens loaned to me
for study and description by Mr. Dyer, there is a very imperfect
example of one of the largest known species of Silurian Starfishes.
When entire, it could scarcely have measured less than 5 inches in
diameter [R = probably 3 inches, or 75 mm.] across from end to end
of the rays on opposite sides; and it presents a breadth of disk (as
flattened by pressure) of about 2 inches [r = probably seven-eighths
inch, or 22 mm.]. Its state of preservation is, unfortunately, such as
not to admit of systematic description, but I think enough of its
character can be given to enable the student to identify it, by the
additional aid of figures.
"The dorsal side of both disk and rays is composed of numerous
small pieces [with stellate extensions], with the pores between them
apparently so large that these pieces only seem to touch at three to
four salient points of each, so as to form a kind of reticulated struc-
ture; while they each bear a little central tubercle, with a minute pit in
its top for the articulation of small, short [blunt] spines, generally
about 0.07 to 0.09 inch in length, and about 0.02 to 0.03 inch in
thickness. [Each plate bears but one of these spines and there appear
to be no other smaller spines.]
"In one of the axilla between two of the rays the so-called madri-
poriform body can be seen near the margin of the disk. It is nearly
flat, of an obtusely subtrilobate form [probably cruciform], with a
breadth or transverse diameter of 0.30 inch, and a diameter at right
angles to the same of 0.24 inch [not less than 10 mm.]. Its lobed
edge is directed inward toward the middle of the disk, and its middle
lobe is largest, and shows the little divisions between the slits or
furrows, diverging and bifurcating inward like the nervation in the
pinnules of some kind of ferns; while in the smaller lateral lobes these
markings diverge outward.
"The ventral side is also much obscured by the adhering matrix,
and numerous detached and confusedly mingled spines. A row of
comparatively small, tumid, nearly square [infra] marginal pieces,
REVISION OF PALEOZOIC STELLEROIDEA. 121
however, can be seen in places, on each side of the ambulacral
furrows, and alternating with a row of similar but slightly smaller
adambulacral pieces, the number of pieces in each side of these being
about the same. [In the proximal portion of a ray there are 19
marginal plates in 40 mm.]. The marginal pieces seem each to
connect with the adjacent range of dorsal pieces, at places in the
rays, by a little central salient point only, while those of both series
are roughened by numerous comparatively coarse granules [all of
which bear small spines], and each piece (especially [only] of the
marginal row) also shows a small central pit for the articulation of
a spine. These spines are larger than those connected with the
dorsal pieces, being generally about 0.22 inch in length and 0.05 inch
in thickness; they are smooth, straight, rounded, slightly thickened at
the attached end, and tapering at the free end to a slightly blunted
point. [The adambulacral plates also possess these large spines,
which are inserted on the distal inner edge of each plate.]"
In each interbrachial area there are two pairs of wedge-shaped
pieces and a single diamond-shaped interbrachial marginal plate.
There are at least 4 (and may be as many as 10) pairs of adam-
bulacral plates orally or in the actinal interbrachial areas, the terminal
pair being of the oral armature.
Ambulacral plates much wider than long, increasing rapidly in
width toward the mouth. Each plate is carinated as in P. sped-
osus and proximally as in P. magnificus, with the podial openings
confined to the extreme outer edge of the ambulacral furrows.
Formation and locality. Found at Cincinnati, Ohio, in the Mays-
ville beds, about 200 feet below the top of the hills. The ,holo type
is in the Dyer collection of the Museum of Comparative Zoology,
Harvard University (No. 13). Another specimen from near the top
of the Maysville at Cincinnati, preserving only the disk and showing
best the actinal area, is in the Gurley collection of the University
of Chicago (No. 10989). Originally this individual appears to have
been larger than the holotype.
Remarks. The relationship of this species is with P. spinulosus,
but the larger size, greater development of both dorsal and ventral
spines, absence of columnar arrangement of the abactinal plates,
and the presence of five instead of two or three interbrachial marginal
plates will readily separate P. dyeri from the latter species. Even
though this form appears to have attained a larger growth than
P. magnificus, which it resembles in some characters, it differs in
being more ponderous in its skeleton and in the far stouter and
longer actinal spines.
Regarding the generic position of Palseaster dyeri, Meek wrote:
" Possibly I should have called it Petraster dyeri, for if the apparent
presence of a few disk plates on the ventral side between the mar-
122 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
ginal pieces and the adambulacral, is not deceptive, it would, in
that respect, conform to the structure of that group, and differ
from Palseaster, as now understood; though I am inclined to think
this appearance due to the accidental displacement of the parts at
the point where there are some indications of a few disk pieces."
These pieces are not to be correlated with the accessory interbrachial
plates of Petraster, for in this genus accessory plates consist of two
short columns situated between the adambulacral and marginal
plates and uniting in the axillary areas. In P. dyeri, however, the
four interbrachial marginal plates are continuous with the infra-
marginal columns and evidently were derived from that series,
and are not accessory interbrachials. It is true that all interbrachial
plates increase the size of the disk, but in one they are derived from
the inframarginal series, by crowding and in the other they are
interpolated, newly developed plates that force apart the columns
of plates in the rays sometimes almost to the distal ends.
PROMOPAL.EASTER MAGNIFICUS (Miller).
Plate 21, fig. 1; plate 22, fig. 1; plate 23, figs. 1-3.
Palseaster magnificus MILLER, Journ. Cincinnati Soc. Nai. Hist., vol. 7, 1884,
p. 16, pi. 4, figs. 3, 3a.
Original description. "The diameter or breadth of the disk is
one and one-fourth niches, and the distance from the point of one
ray to the point of the opposite one, if the rays were wholly preserved
in the specimen under examination, would be fully 6 inches. * * *
"The plates upon the dorsal side are very convex, and part of
them, at least, were spine-bearing, though it would seem that there
was not more than one spine upon any single plate. The arrange-
ment of the plates on the dorsal side of the rays is very ornamental.
A single series of highly convex or conical plates, larger than the
others, and each evidently bearing a central spine, occupies the
middle of each ray; on either side near the margin of each ray there
is a similar series, and the two intervening spaces are filled with
smaller, convex plates arranged in rows which are directed diagonally
forward from the plates of the side series to the plates of the central
series, forming angles with each plate in the central series occupying
an angle. This disposition of the plates on the dorsal side of the
rays will, so far as known, serve to distinguish this species from any
hitherto described.
"The plates covering the dorsal side of the body or disk have been
so much disturbed in our specimen that one can not correctly define
them.
"The ambulacral furrows are wide. The marginal plates are
hexagonal, about the size of the larger plates on the dorsal side of
KEVISION OF PALEOZOIC STELLEKOIDEA. 123
the rays, and separated from the side series by intervening smaller
plates. Each bore several small spines, as shown by the small pits
for their articulation. * * *
"The adambulacral plates are hexagonal and much wider than
long. They are more numerous than the marginal plates near the
disk, but toward the apices of the rays they interlock alternately
with the marginal plates. The number on each side of a furrow hi
a complete ray would be 50 or more. The pits for the articulation
of the spines are as numerous as they are on the marginal plates.
These two series on the ventral side of the species, with numerous
spines upon each plate, are in striking contrast with the plates on
the dorsal side of the rays, where no plate bears more than a single
spine.
"The ambulacral plates have their greatest length across the
rays, and seem to be about as numerous as the adambulacral ones.
An angular depression marks the center of each ambulacral furrow,
upon each side of which a sharp ridge arises upon each ambulacral
plate, and curving forward and outward abuts against an adambula-
cral plate. "
Emended description. The largest specimen, the holotype, meas-
ures R = (about) 67 mm., r = 17 mm., R = nearly 4r. Another speci-
men: 11 = 58 mm., 3* =15 mm., R= nearly 4r.
Abactinal surface very ornamental. Medially along each ray there
is a radial column of highly convex plates, and two columns of similar
plates, the supramarginals, are situated near the margin. Between
the inframarginal, supramarginal, and radial columns are very nu-
merous diagonal rows of smaller conical or angular plates, from 2 at
the apex of the ray to 14 at base of same in each row on each side of
the radial column. This may be stated in another way. Between
the radial and supramarginal columns and again in the ambital area,
or between the supramarginals and inframarginals, the diagonal rows
have from one to seven plates in each area. These rows are directed
diagonally and distally across the rays from the sides to the median
column, the apex of the angle thus formed being occupied by the
larger plates of the radial column. All of these smaller plates appar-
ently bore a single short spine. Between all of the diagonal rows
are inserted both proximally and distally numerous minute, conical,
irregular-shaped plates. The supramarginal columns continue across
the disk and those of adjoining rays unite in the axillary areas. The
radial columns also extend over the disk but do not attain the center,
and the diagonal arrangement of the small plates ceases near the
proximal portion of the rays. The pattern arrangement of the disk
plates can not be made out since their original position is disturbed.
Madreporite quite large, quadri-lobate in outline, depressed, convex
and very finely radiately striate. It is situated in one of the axillary
124 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
areas, less than 4 mm. from the margin. Against it rest two columns
of supramarginal plates.
The inframarginal plates, of which there are at least 45 in each
column, are distally small and subquadrate but proximally increase
rapidly in size and at the base of the columns are three to four times
as wide as long. These plates bear numerous pitted pustules upon
which articulated short, sharp, smooth spines. The columns of ad-
joining rays meet in the axils and continue into the interbrachial
areas.
The adambulacral plates are like the inframarginals but do not
increase so rapidly in width as those plates and are somewhat more
numerous, there being in a complete ray about 52 in a column.
Proximally some of the large plates are broken into two or three
often very convex ossicles. In addition to the small spines like those
of the inframarginals, each plate bears on its ambulacral edge a prom-
inent socket in which articulated a long, thick, smooth spine. The
adambulacral plates in the proximal third of the rays each have a
small, more or less wedge-shaped, carinated, accessory adambulacral
plate. In this region these ossicles separate the adambulacrals from
the ambulacrals and beneath the accessory adambulacral plates are
the podial openings. Five or six adambulacral plates of each column
abut against the interbrachial marginal plates, while five or six pairs
continue orally, the terminating pieces being of the oral armature.
Ambulacral furrows wide, increasing in width proximally. An
angular median sinus marks each ambulacral furrow. Ambulacral
plates short but wide, slightly overlapping proximally, and one to
each adambulacral plate. All of the plates are strongly carinated,
the carina in the distal two-thirds converging medially and proximally.
In the proximal third the carinse gradually are changed in form and
are there arranged in forked pairs. In other words, the carinae adjoin
the accessory adambulacrals, are arranged in pairs, continuing ad-
joining and straight for a short distance, and then diverge in broad
curves, terminating near the center of the ambulacral groove. This
arrangement produces in the proximal portion of the ambulacral
furrows four columns of ovoid pits, all terminating laterally into
podial openings. Every other podial opening belongs to the same
column, one series beneath the adambulacrals, the other gradually
passing medially in the ambulacral grooves. In the distal two-thirds,
however, the podial openings are arranged in single columns, one to
each ambulacral plate and issuing from underneath the adambulacrals.
Interbrachial areas with seven plates, each composed of two prox-
imal inframarginals, beneath which are two pairs of interbrachial
marginals and a single diamond-shaped axillary marginal plate. All
bear similar spines and ornamentation on the inframarginals. Some-
times one of the distal interbrachial marginals may be divided and
REVISION OF PALEOZOIC STELLEEOIDEA. 125
the arrangement of the interbrachial plates made irregular. In a
specimen from Jefferson County, Indiana, there are 3 pairs of inter-
brachial marginals instead of 2 as in the Ohio individuals. There-
fore there are also more adambulacrals here, 14 pairs against 10 in
the typical specimens.
Formation and locality. Two free specimens of this magnificent
starfish were found in the Richmond formation near Waynesville,
Ohio. Originally pieces of these two individuals were glued together
as one specimen, which is the cause of our figures being less perfect
than Mr. Miller's. Fragments of seven other individuals were found
in the vicinity of Waynesville. In the University of Chicago col-
lection (No. 10981) there is a specimen from Jefferson County,
Indiana, apparently of this species. Mr. Vaupel secured the distal
portion of a ray of an apparently young example of this species in
the Maysville beds on Rons Hill, Cincinnati, Ohio, and Mr. Ulrich
has fragments of four individuals from the Maysville strata about
Cincinnati, and Covington, Kentucky.
Remarks. This species is readily separated from the other species
of Promopalxaster by the marked diagonal rows of abactinal plates.
Actinally P. magnificus is readily distinguished from P. exculptus and
P. spinosus in having five instead of three interbrachial marginal
plates in each area, in the rapidly increasing size of the plates in the
inframarginal and adambulacral columns, and in the proximal am-
bulacral plates. P. speciosus differs at once from P. magnificus in that
its abactinal plates are arranged in regular longitudinal columns,
are far less numerous, larger and are all nearly of a size.
This is one of the largest and best preserved of American Paleozoic
starfishes. P. dyeri may be a larger species but is distinguished abac-
tinally not only by the arrangement and difference in the forms of
the plates but also by the short, thick, blunt spines which now ap-
pear to have no definite arrangement, while P. magnificus has all of
its very fine spines arranged in diagonal rows like the plates.
Cat. Nos. 40883, 60621, 60622, U.S.N.M.
new subfamily.
Aberrant Promopalaeasteridae with the axillary and interbrachial
areas composed entirely of adambulacral pieces.
Contains :
Anorihaster.
ANORTHASTER, new genus.
Plate 13, fig. 4; plate 20, fig. 1.
Anorthaster SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914, p. 11.
A + orihos + aster = out of the regular, in reference to the completely
adambulacral na-ture of the interbrachial areas.
126 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Generic characters. Disk small, with apparently small interbrachial
arcs. Rays five, of moderate length and tapering at first slowly
and then rapidly distally.
Abactinai area unknown.
Inframarginal plates small, the columns not attaining the axils,
but resting upon the second enlarged axillary adambulacral plate.
Adambulacral plates distally like the inframarginals, increasing
in size slowly proximally until near the axillary region, where four
plates suddenly attain great width, terminating in the axils of the
rays and here suppressing the inframarginal plates. Eight inter-
brachial adambulacrals of varying form are situated orally to the two
large axillary adambulacral plates, and at first give one the impression
of slender interbrachial areas of the same nature as in Promopalszaster.
These plates, however, are not interbrachial marginals, as in other
genera, and derived by the oral crowding of proximal inframarginals.
They are distinctly adambulacral ossicles, forming slight interbrachial
areas, and, with the axillary adambulacrals, form small arcs. For the
shape and position of these plates see figure 1, plate 20.
Ambulacral plates alternating, slightly carinated, and very short
but wide. In the distal region they are as long as the adambulacral
plates, but proximally decrease in length where there are nearly two
of them to one adambulacral ossicle. Podial openings between every
other plate through the sutures of adjoining pieces at their outer
edge and immediately beneath the inner edge of the adambulacral
ossicles. The proximal plates of each column are modified into oral
armature pieces that are slightly tumid, as long as wide, and sub-
triangular in outline. Between the proximal ambulacrals and orad
to the proximal interbrachial adambulacrals are situated additional
single, very small, quadrangular oral armature plates.
Genoholotype and only species. Palseaster miamiensis Miller.
Distribution. Ordovicic, Richmond formation, near Waynesville,
Ohio.
Remarks. The suppression of the inframarginal plates in the axils
of the rays by two of the adambulacral plates of each column and
the oral continuation beneath the latter of four pairs of adambulacral
plates holding an interbrachial position distinguish this genus
fromPromopalseaster, to which it otherwise seems very closely related.
This character also distinguishes it from all other associated genera.
The interbrachial structure of Anorthaster appears abnormal. In
Mesopalseaster and Promopalseaster the proximal inframarginal plates
are crowded orally and form interbrachial areas; in Anorthaster their
formation is effected in the opposite way, not by the inframarginals,
but by the adambulacrals. This character is seen in a single speci-
men, but it is so regularly developed and well preserved as to indicate
a normal and not a pathologic or abnormal condition.
KEVISION OF PALEOZOIC STELLEROIDEA. 127
ANORTHASTER MIAMIENSIS (Miller).
Plate 13, fig. 4; plate 20, fig. 1.
Palseaster miamiemis MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 3, 1880,
p. 143, pi. 4, fig. 3.
Original description. " Pentagonal; rays about one and a half
times the diameter of the body, or about nine-tenths of an inch;
diameter of the body about six-tenths of an inch; breadth of a ray
at the point of junction with the body a little more than half the
diameter of the body, or about seven-twentieths of an inch; rays
obtusely pointed.
"Marginal plates wider than long, and numbering about 12 in
the length of half an inch from the body. Two marginal plates
form the junction of the rays. Ambulacral furrow wide, the plates
being more than twice as long as wide. There are about 18 ambu-
lacral plates in a length of one-half inch, and each one is provided
with an angular ridge taperingfrom the marginal plates to the furrow."
Emended description. R = 23 mm., r = 7 mm., R = 3.2r. Width of
rays at base 8 mm.
Rays tapering very slowly in the proximal half, but more rapidly
distally.
Abactinal area unknown.
Inframarginal plates about 16 in each column and subquadr angular
in outline. These ossicles do not continue into the axils, but rest
upon the second large axillary adambulacral plate.
Adambulacral plates about 18 in each column, like the infra-
marginal pieces, increasing in size proximally, while the two axillary
ones are suddenly enlarged, extend to the margin of the disk, and have
the position of marginal ossicles. Orally to the latter the adambu-
lacral plates continue as interbrachial ossicles. The distal pair is
large and both are wedge-shaped, followed by two pairs of quite
small quadrangular plates and a final pair of long and narrow plates;
in all there are, therefore, 22 adambulacral plates in each column.
The interbrachial adambulacral plates are terminated by a single
quadrangular oral plate wedged in between the proximal ambu-
a ILbulacral furrows very wide, broadly gutter-shaped. Ambu-
lacral plates about 30 in a column, slightly alternating, decidedly
wider than long proximally, increasing in length distally, and finally
becoming about as wide as long near the apex of a ray. Each plate
is slightly carinated, the ridges crossing the plates from the lateral
distal edge to the medial proximal edge. The proximal ambulacral
plates are modified, conspicuous, and subtriangular in outline
Podial openings between two adjoining plates in alternate sutures at
50601 Bull. 8815 9
128 BULLETIN- 88, UNITED STATES NATIONAL MUSEUM.
their extreme outer edges and immediately inside of the adambu-
lacral plates.
Formation and locality. In the Richmond formation, near Waynes-
ville, in Montgomery County, Ohio. The holotype is in the Harris
collection of the United States National Museum.
Eemarks. -This species has a general resemblance to some of the
Promopalseasters, particularly P. ~bellulus and P. wylcoffi, but the
peculiar arrangement of the axillary and interbrachial adambulacral
plates will distinguish it not only from them but from all other
Ordovicic asterids.
Cat. No. 40880, U.S.N.M.
Family XENASTERID^E Schondorf.
Xenasteridx GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 346; Treat. Zool., vol. 3,
Echinoderma, 1900, p. 250.
Palsegoniasteridse STURTZ, Palaeontographica, vol. 36, 1890, p. 247 (not opp. 246).
Xenasteridse SCHONDORF, Palaeontographica, vol. 56, 1909, p. 105; Jahrb. nassau-
isch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 25; vol. 63, 1910, pp. 244, 250.
Progressive Phanerozonia having the general characters of the
Promopalseasterinse, with the addition of accessory interbrachials.
The ambulacral and adambulacral plates are directly opposite one
another and do not waver between the slightly alternate and opposite
arrangement in the Promopalaeasteringe. Podial openings in two
columns in each ambulacral furrow, the apertures lying between
each four adjoining plates, that is, between two adambulacrals and
two ambulacrals.
Contains :
Xenaster Simonovitsch.
Agalmaster Schondorf.
Khenaster Schondorf.
Eifelaster Schondorf.
Trimeraster Schondorf.
Schondorf defines the family as follows (leaving out much of his
detail) :
Lower Devonic starfishes with well-developed supramarginals and
usually still larger inf ramarginals ; the latter are somewhat more numer-
ous than the former, causing them to be unequally superposed. Ambu-
lacra opposite one another and opposite the adambulacrals; podial
openings in two rows. Actinally each interradius has either three or
five interbrachial plates; abactinally this area is filled with small
accessory plates and margined only by inframarginals. Disk skeleton
consisting of small separated pieces, a central disk plate, five primary
radialia, five primary interradialia, and more or less of accessory disk
KEVISION OF PALEOZOIC STELLEEOIDEA. 129
ossicles. Eays with three columns of plates, one radial and two
supramarginal, that may or may not be separated from one another
by small accessory pieces. Madreporite dorsal and interradial.
Genus XENASTER Simonovitsch emend. Sehondorf.
Plate 24.
Archseasterias MULLER, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 12, 1855, pp.
6, 8 (Archxasterias founded in error).
Xenaster SIMONOVITSCH, Sitzb. mat.-naturw. Classe Akad. Wiss., Wien. vol. 64,
Abt. 1, 1871, p. 88. SCHONDORF, Palseontographica, vol. 56, 1909, pp. 42,
105 (complete synonymy given here); Jahrb.nassauisch. Ver. Naturk., Wies-
baden, vol. 62, 1909, p. 26.
Generic characters. Xenaster has its nearest relationship in
Devonaster. No detailed description need be given, since the splendid
illustrations of Sehondorf give all the necessary characters. It
differs from Devonaster as follows:
In Devonaster there is but a single interbrachial plate in each
actinal axil, the axillary interbrachial, while in Xenaster there are
two pairs of interbrachial marginals and a single axillary inter-
brachial plate; in other words, five interbrachial plates instead of
one (1+2+2). In addition to these plates in Xenaster, the interbra-
chials are more or less surrounded by small accessory pieces, and then
other similar plates separate the inf ramarginal and adambulacral col-
umns to near the outer third of the rays. The inf ramarginal plates in
Xenaster do not increase so rapidly in width proximally as do these
plates in Devonaster, and while the interbrachial areas are of about
the same size in both genera, the space occupied by the additional
interbrachial marginal and accessory interbrachial ossicles in the
former is taken up in the latter genus by the increased size of the
inf ramarginal pieces.
Abactinally Xenaster has the generic characters of Devonaster.
The plates in all the columns of the former genus are more prominent
and rounder, particularly the accessory plates, of which there appear
to be less than in Xenaster. The center of the disk in both genera
is occupied by a small central disk plate, but there are many more
accessory plates around it in Devonaster than in Xenaster. In both
genera the supramarginal columns converge proximally, producing
small disk areas between the infra- and supramarginal plates occu-
pied by ambital accessory plates.
Madreporite large and striate in Xenaster.
Genotype. X. margaritatus Simonovitsch.
130 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Distribution. Restricted to the Lower Devonic of Germany.
The species are:
X. margaritatus Shnonovitsch.
X. dispar Schondorf.
X. elegans Schondorf.
X. (?) rhenanus (Miiller).
Remarks. Miiller in his description of Asterias rtienana, 1 the geno-
type of Archseasterias , notes that the adambulacral plates of adjoining
columns alternate and that the ambulacral columns ' ' have the appear-
ance also of alternating/' which if established would be very remark-
able, and would justify the creation of a distinct genus Archseasterias
for this asterid of the Rhine grauwacke. Schondorf shows that
these statements of Miiller' s are very faulty and further that the
genus Archseasterias has not been denned so as to be recognizable.
Under these circumstances it was best for Simonovitsch to disre-
gard Archseasterias and to establish a new generic name for his species
margaritatus. There is no Paleozoic genus of starfishes better
described or illustrated than this one by Simonovitsch, and should
it ever prove that Asterias rhenana Miiller is identical with Xenaster
margaritatus, a possibility, the latter should not be made to give
way to the former on the ground of priority. It is an injustice to
extend the law of priority indiscriminately to all work, and to reject
Archseasterias is no reflection on the work of Miiller, as his material
was very poor and his work of early date in paleontology. He pub-
lished at a time when it was very desirable to learn anything what-
ever about Paleozoic starfishes.
As pointed out above, Xenaster has much of the structure of
Devonaster, yet it differs in the important character of having
numerous accessory interbrachial plates. This feature seemingly
relates it with forms like Palasterina which have a great abundance
of accessory interbrachials, yet in no interbrachial area of this genus
are there axillary interbrachials or interbrachial marginals. In other
words, the Palasterina type of disk is made by the intercalation of
accessory plates between the inframarginals and adambulacrals,
whereas hi Devonaster it is accomplished by the greater width of the
inframarginals and by the oral crowding of a plate which in primitive
forms is an axillary marginal. Devonaster although from younger
strata is more primitive than Xenaster because of its simpler inter-
brachial actinal structure. If it were not for the accessory inter-
brachials, Xenaster would be closely related to Promopalseaster which
also has a number of proximal inframarginals crowded into the inter-
brachial areas. It is the Promopalseaster stock out of which Xenaster
probably developed, while Devonaster apparently came through
Mesopalseaster.
* Verb, naturh. Ver. preuss. Rheinl., etc., vol. 12, 1855, pp. 6, 8.
REVISION" OF PALEOZOIC STELLEROIDEA. 131
XENASTER MARGARITATUS Simonovitsch emend. Schondorf.
Plate 24, figs. 1,2.
Xenaster margaritatus (part) SIMONOVITSCH, Sitzb. mat.-naturw. Classe Akad.
Wiss., Wien, vol. 64, Abt. 1, 1871, p. 88, pis. 1, 2. SCHONDORF, Palseon-
tographica, vol. 56, 1909, pp. 43, 106, pi. 7, figs. 1, 3, 4; pi. 8, fig. 1; pi. 11,
figs. 2, 3, 10, 11 (complete synonymy given here); Jahrb. nassauisch. Ver.
Naturk., Wiesbaden, vol. 62, 1909, p. 26, pi. 2, figs. 1, 2. SPENCER, Mon.
Brit. Pal. Asterozoa, pt. 1 (Palaeontogr. Soc. for 1913), 1914, p. 30.
This species is one of the best described and illustrated of Paleozoic
starfishes and does credit to its author, and to its subsequent reviser,
Schondorf. It occurs rarely in the Lower Devonic (Upper Coblen-
zian) as natural molds at Niederlahnstein on the Rhine, Germany.
XENASTER DISPAR Schondorf.
Xenaster margaritatus (part) SIMONOVITSCH, Sitzb. mat.-naturw. Classe Akad.
Wiss., Wien, vol. 64, 1871, Abt. 1, pi. 2, figs. 2, 2a, 26.
Xenaster dispar SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 50, 84, 106,
pi. 9, fig. 1; pi. 11, fig. 13; Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol.
62, 1909, p. 28, pi. 3, fig. 3.
From the Upper Coblenzian at Niederlahnstein, and Miellen,
Germany.
XENASTER ELEGANS Schondorf.
Xenaster margaritatus FOLLMANN (not Simonovitsch) , Jahrb. naturh. Ver. preuss.
Rheinl., etc., vol. 48, 1891, p. 150.
Xenaster elegans SCHONDORF, Palseontographica, vol. 56, 1909, pp. 53, 107, pi. 9,
figs. 2, 3; pi. 11, figs. 4, 12; Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol.
62, 1909, p. 40, pi. 4, figs. 6, 7.
From the Lower Devonic (Coblenz quartzite) at Konigstuhl, near
Rhens, Germany.
XENASTER (?) RHENANUS (Muller).
Asterias (Archseasterias} rhenana MULLER, Verh. naturh. Ver. preuss. Rheinl.,
etc., vol. 12, 1855, p. 6, pi. 1, figs. 4-6.
Xenaster (?) rhenana SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 55, 107,
text fig. 2 (complete synonymy given here); Jahrb. nassauisch. Ver. Naturk.,
Wiesbaden, vol. 62, 1909, p. 28.
This species is based upon incomplete material and can not be
made out fully. It seems probable that it is a species of Xenaster
and closely related to X. margaritatus Simonovitsch. The examples
are in the Berlin Museum and were found in the Lower Devonic at
Winningen, near Coblenz, and Kemmenau, near Ems, Germany.
Genus AGALMASTER Schondorf.
Text fig. 9.
Agalmaster SCHONDORF, Palseontographica, vol. 56, 1909, pp. 58, 107; Jahrb.
nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 29.
Original remarks. Agalmaster differs from Xenaster in the following
characters: Forms larger, broader rayed, with flat actinal but highly
arched abactinal plates. Supramarginals much stronger than the
large inframarginals and projecting considerably beyond these.
132 BULLETIN 88, UNITED STATES NATIONAL MUSEUM..
\
Abactinal interbrachial area small, and occupied in the main by two
axillary inf ramarginals ; actinally this area has the same structure as
Xenaster. Madreporite dorsal.
Genoholotype (the writer here selects the first species). A. miel-
lensis Schondorf. Other species are A. grandis Schondorf and A.
intermedius Schondorf, both from the Lower Devonic of Germany.
/ \ Remarks. The only
/ \ character that may be
of generic value and
that at present distin-
guishes Agalmaster from
Xenaster is that in the
former genus the supra-
marginals margin the
rays instead of the in-
f ramarginals, as is the
case in the latter genus.
In a second species (A.
grandis) the small ac-
cessory interbrachials
crowd in between all the
pairs of interbrachial
FIG. 9. AGALMASTER MIELLENSIS, AFTER SCHONDORF. SCHEMATIC marginals and further
ARRANGEMENT OF ABACTINAL PLATES. R* AND -R 3 , SECOND AND , . '
THIRD PROXIMAL RADIAUA; Jn, PRIMARY INTERRADIALIAJ Cdp, tllCre &TQ J&T ID.OTG OI
CENTRAL PLATE; mdp, MADREPORITE; mo, SUPRAMARGINALIA; ^ggQ pieces between
mu, INFRAMARGINALIA; Pifm, PRIMARY INTERRADIALIA.
the inframargmals and
adambulacrals; in fact, there are three to four columns of these in-
stead of one (as in A. mieUensis).
AGALMASTER MIELLENSIS Schdndorf.
Text fig. 9.
Agalmaster mieUensis SCHONDORF, Palseontographica, vol. 56, 1909, pp. 58, 107,
text fig. 3; pi. 8. fig. 4; pi. 9, fig. 4; pi. 11, figs. 1, 5; Jahrb. nassauisch. Ver.'
Naturk., Wiesbaden, vol. 62, 1909, p. 29, pi. 3, fig. 2; pi. 4, figs. 1, 2.
A single specimen with the rays about 43 mm. long was found in the
Lower Devonic (Upper Coblenzian) at Miellen-on-the-Lahn, Germany.
AGALMASTER GRANDIS Schdndorf.
Xenaster margaritatus SIMONOVITSCH (part), Sitzb. mat.-naturw. Classe Akad.
Wiss., Wien, vol. 64, Abt. 1, 1871, pi. 2, figs. 3, 3a.
Agalmaster grandis SCHONDORF, Palseontographica, vol. 56, 1909, pp. 62, 108,
pi. 9, fig. 5; Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 29,
pi. 3, fig. 1.
One specimen (Xenaster margaritatus Simonovitsch, part) from
the Lower Devonic (Upper Coblenzian) at Niederlahnstein on the
Rhine, Germany.
REVISION OF PALEOZOIC STELLEROIDEA.
133
AGALMASTER INTERMEDIUS Schflndorf.
Agalmaster intermedius SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 82,
108, pi. 11, fig. 14; Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62,
1909, p. 30.
From the Upper Coblenzian at Miellen-on-the-Lahn, Germany.
Genus RHENASTER Sehondorf.
Text fig. 10.
Rhenaster SCHONDORF, Palseontographica, vol. 56, 1909, pp. 65, 108, pi. 8, fig. 3;
pi. 10, figs. 1, 4.
Remarks. Rhenaster is a small form and differs from Xenaster,
Agalmaster, and Devonaster in the complete absence of all abactinal
accessory plates, allow-
ing the radial and supra-
marginal pieces to ad-
join one another closely.
Dorsal axillary disk ~7=TP?^O g V\C\
T i I /I )s~*\-/ sr?~^ ^-' /\S. \V
areas not completely
but almost absent,
supramarginals in the
main lying wholly upon
the inframarginals and
the two columns of
plates together margin-
ing the rays. There
also are no accessory
plates of any kind on
the actinal side. Other-
wise the generic struc-
ture is that of Xenaster, except that in Rhenaster the first pair of
interbrachial marginals are not yet pushed so much orally and they
almost take part in margining the axils. Madreporite unknown.
Genoholotype and only species (one specimen). R. schwerdi Schon-
dorf, from the Lower Devonic (Upper Coblenzian) at Dorrbachtal,
near Coblenz, Germany.
Genus EIFELASTER Sehondorf.
Etfelaster SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 70, 108, pi. 10, figs.
7,8.
Remarks. This genus is very closely related to Rhenaster and
differs only in that there are small dorsal interbrachial areas between
the adjoining proximal portions of the supramarginal columns, and
these are filled with a small number of accessory ambital pieces.
The outer portion of these ambital areas is made up of the two basal
inframarginals that occupy the axils of the ventral side.
FIG. 10. REENASTER SCHWEKDI, AFTEK SCHONDORF. SCHEMATIC
ARRANGEMENT OF ABACTINAL PLATES. SYMBOLS AS IN FIG. 9.
134 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
The genoholotype and only species, E. fottmanni, occurs in the
Lower Devonic (Lower Coblenzian) at Landscheid, near Wittlich,
in the Eifel, Germany. Plastotypes of the holotype were made for
the writer by Professor Jaekel, of Greifswald, and these are now in
the Yale University collection.
Genus TRIMERASTER Schondorf.
Plate 19, fig. 1.
Trimeraster SCHONDORF, Palaeontographica, vol. 56, 1909, pp. 68, 108, pi. 7,
fig. 2; pi. 10, figs. 5, 6.
Remarks. Trimeraster is a small form, very closely related to
Xenaster, and can be distinguished generically only in that the former
has inside of the inframarginals but one. pair of interbrachial margi-
nals and a single axillary interbrachial in each interbrachial field
instead of the five plates of the latter genus.
Genoholotype and only species (one specimen). T. parvulus Schon-
dorf, from the Lower Devonic (Coblenz quartzite) at Konigstuhl,
near Khens, Rhine Valley, Germany.
NEOPAL^ASTERIDJE, new family.
Progressive Phanerozonia without interbrachial arcs. The columns
of supra- and inframarginal plates wholly superposed. Madreporite
abactinal. Well-developed ocular plates present. Ambulacrals alter-
nating. Disk with a ring of large plates of basal, radial, and inter-
radial ossicles.
Contains :
Neopalseaster, new genus.
The origin of this family is uncertain, but it seems to be related to
the Promopalseasteridse, since its essential characters are those of the
Mesopalaeasterinse. In the latter the supramarginals do not com-
pletely overlie the inframarginals making one consolidated column as
in Neopalseaster. It is, so far as known, the oldest family developing
ocular plates. This fact is of considerable importance for it means
that the oculars were originally basal radialia, and that they have
progressively wandered from the disk and then remained at the tip
of the rays.
NEOPALSEASTER, new genus.
Plate 13, fig. 5; pi. 23, fig. 4.
Neopalseaster SCHUCHERT, Fossilium Catalogue, Animalia, pt. 3, April, 1914, p. 26.
Generic characters. Disk of medium size, without interbrachial
arcs. Eays five, short and tapering rapidly.
Abactinally the disk has a small central plate surrounded by
numerous smaller plates. Bounding the smaller accessory disk
KEVISION OF PALEOZOIC STELLEEOIDEA. 135
plates is a ring of very large, thick, tumid, variously shaped, radial
and interradial plates. Of the latter there may be one plate inside
of the basal supramarginals of adjoining columns, or this plate may
be divided, when the pieces appear as inwardly crowded supramar-
ginals. Laterally and distally upon the interradials are situated the
large basal radial plates. The rays are bounded laterally by columns
of large, thick, subquadrate, tubercular supramarginals terminated dis-
tally by single ocular plates. These plates appear to be common to
both the actinal and abactinal areas and must therefore be the united
infra- and supramarginal columns. Between the supramarginals dis-
tally appear single small isolated plates which proximally become
larger and continuous, forming an inconspicuous column of radial
ossicles. On each side of this column are inserted a number of appar-
ently irregularly arranged small accessory plates which are like those
of the disk. All of the plates of the abactinal area are finely tuber-
culose, these points being for the attachment of very fine, short spines.
Madreporite small, rounded, not very convex, very finely radially
striated and resting directly upon two basal supramarginal plates of
adjoining rays.
Ocular plates small but distinct, one terminating each ray and rest-
ing against the distal marginal plates.
Supra- and inframarginal plates large and conspicuous, apparently
firmly united and indistinguishable laterally as separate columns.
Abactinally the supramarginal plates are wider and overlie more or
less the adambulacrals.
Adambulacrals smaller than the inframarginals but otherwise
resemble the latter.
Ambulacral grooves narrow. Ambulacra! plates alternating, about
as numerous as the adambulacrals, 1 -shaped, with the podial open-
ings between adjoining plates.
All other actinal characters unknown,
Genoholotype and only Jcnown species. Palseaster crawfordsvillensis
Miller. From the Keokuk crinid beds near Crawfordsville, Indiana.
Remarks. Neopalseaster retains a very primitive abactinal plate
structure and calls to mind Hudsonaster, Palseaster, and typical
Mesopalseaster. It is distinguished from these early Paleozoic genera
by the complete consolidation of the supra- and inframarginal plates
into one column, and the presence of ocular plates. The inter-
brachial area is unknown in Neopalseaster, but its abactinal structure
seems to indicate a single interbrachial plate in each area bounded
distally by two basal inframarginal plates. If this interpretation is
correct, then this character will further distinguish Neopalseaster from
Hudsonaster and Palseaster.
136 BULLETIN" 88, UNITED STATES NATIONAL MUSEUM.
NEOPAL^EASTER CRAWFORD SVILLENSIS (Miller).
Plate 13, fig. 5; plate 23, fig. 4.
Palseaster crawfordsmllensis MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 2,
1880, p. 256, pi. 15, fig. 3; N. Amer. Geol. Pal., 1889, p. 265, fig. 379.
Original description. "This species is founded upon the dorsal
view of a single specimen. The rays are longer than the diameter of
the body, and not of uniform size. They are flattened or depressed
in the middle, as is also the central part of the body. Many of the
plates possess a central tubercle or small spine, and probably all of
them did.
"The marginal plates are large ; somewhat elliptical in outline, and
have their shorter diameters in the direction of the length of the
rays. There are about 12 plates on each side of a ray, and they
come together at about the eighth plate from the body, though in
the ray opposite the madreporiform tubercle they come together at
the seventh. The space between the marginal plates of each ray
is filled with smaller plates; three of these unite the larger plates at
the body, but they diminish in number toward the apex of the ray,
and cease at the eighth plate. In addition to the two large plates
which form the junction of the rays with the body, a few large plates
cover the outer part while the central part is covered by smaller
plates. The madreporiform tubercle is supported by three plates,
two of them are large marginal plates, which form a junction between
two rays, and the other is a large plate within, forming part of the
covering of the body."
Emended description. The type-specimen measures: R, = 22 mm.,
r=7 mm., K, = 3.1r > . Width of rays at base 7 mm. Another speci-
men in the Museum of Comparative Zoology has rays 3 mm. longer.
Rays short, tapering rapidly. Disk rather large for a small species.
Abactinally the disk has a central circular plate surrounded by
numerous small, conical, irregularly disposed, accessory plates, among
which are a few larger pieces usually adjacent to the large interradial
plates. Outside of the small disk plates is a ring of very large, irregu-
larly shaped, radial and interradial plates. Five of these are inter-
radial in position, and are situated just within the basal plates of
adjoining supramarginal columns. These plates may remain normal
as a single plate or may be divided into two plates, when they some-
what resemble inwardly crowded supramarginals. Upon these
laterally and distally rest five large radial pieces, the basal plates
of the radial columns. It is very probable that the radial columns
are continuous from the basal plate to near the distal portion of the
ray, where a few of these plates are much reduced in size and are
EEVISIOIST OF PALEOZOIC STELLEEOIDEA. 137
separated and wedged in between the adjoining four supramarginal
ossicles. There appear to be 10 or 11 of these in a column. On each
side of the radial columns in the proximal half of each ray are a number
of small accessory plates arranged in irregular columns. All of the
abactinal plates are more or less convex and bear numerous small
pustules clustered around a larger central one, all of which bore small
spines.
Supramarginal plates very conspicuous, subquadr angular in out-
line, generally a little wider than long, with numerous small pustules,
among which toward each lateral edge is a large one upon which
probably articulated prominent spines. There are from 11 to 13 of
these plates in each column. In the distal third of a ray both
columns approximate and adjoin each other closely. Each supra-
marginal plate lies wholly and directly over an inframarginal and the
pieces appear as one consolidated plate with the abactinal side convex
and the actinal nearly flat. The supramarginals are the widest and
extend in part over the adambulacrals.
Madreporite small, circular in outline, slightly convex and very
finely radially striated. It lies directly upon two basal supramar-
ginal plates of adjoining rays and directly outside of one of the large
interradial plates.
An ocular plate terminates each ray.
Inframarginals apparently like the supramarginals excepting
distally, where they are nearly twice as long as wide.
Adambulacral plates smaller than the inframarginals, nine in
the space occupied by five of the latter, somewhat wider than long,
convex, and covered with numerous small pustules.
Ambulacral grooves very narrow in the distal region. The ambu-
lacral plates appear to be as numerous as the adambulacrals, and are
h- -shaped, with the podial openings between the plates in about the
mid-width of the column.
All other actinal characters are unknown, since the two specimens
expose the abactinal area, though one has portions of the plates
removed, revealing the actinal characters described.
Formation and locality. The type-specimen is in the Harris
collection of the United States National Museum, and another
specimen is in the Dyer collection of the Museum of Comparative
Zoology (No. 4). Both were found in the crinid beds of the Keokuk
group of the Lower Carboniferous at Crawfordsyille, Indiana.
Remarks. There is no Carboniferous asterid with which Neopalse-
aster crawfordsvillensis can be confounded.
Cat. No. 60607, U.S.N.M.
138 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Family PALASTERINID^E Gregory (emend.).
Pateasterinidse GREGORY (part), Geol. Mag., dec. 4, vol. 6, 1899, p. 348 (contains
Palasterina, Schaenaster, and Schuchertia)} Treat. Zool., vol. 3, Echinoderma,
1900, p. 250.
Lindstromasterinss GREGORY (part), Geol. Mag., dec. 4, vol. 6, 1899, p. 346 (contains
Lindstromaster and Uranaster).
Palseasterinidse STURTZ, Palaeontographica, vol. 36, 1890, p. 246.
Progressive Phanerozonia developing large interbrachial arcs.
Ambulacral plates more or less alternate. Madreporite abactinal.
Disk large, pentagonal, and the rays separated by well-developed
interbrachial arcs. The inframarginal plates bound the animal and
are separated from the adambulacrals more or less completely by a
varying number of interbrachial plates.
Abactinal surface with longitudinal columns of radial, supra-
marginal and accessory columns of plates, or the radial columns
may be reduced or even replaced by transverse rows of small accessory
pieces. The central region of the disk may retain a ring of larger
basal radial and supramarginal pieces.
Contains:
Pet raster Billings.
Lindstromaster Gregory.
Palasterina McCoy.
Uranaster Gregory.
Palxostella Stiirtz.
Pseudopalasterina Stiirtz.
Genus PETRASTER Billings.
Plates 23, 26, 27.
Petraster BILLINGS, Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858, p. 79.
HALL, Twentieth Rep. N. Y. State Cab.. Nat. Hist., 1868, pp. 283, 294; rev.
ed., 1868=1870, pp. 325, 337.
Palseaster (part) ZITTEL, Handb. Pal., vol. 1, 1879, p. 452.
PalsRosterina of some AUTHORS.
Generic characters. Disk and interbrachial arches more or less
large. General shape pentagonal. Rays five, short and slender or
stout and tapering rapidly.
Abactinal area of rays distally with one medial radial, two supra-
marginal and two inframarginal columns of plates. Between the
radial and supramarginal columns proximally are inserted columns
of accessory plates' In P. speciosus there are five on each side of
the radial column. Between the supramarginal and iinVaniarginal
columns in the axillary and basal ray regions are inserted columns
of ambital plates. In P. speciosus there are three. The plates
are more or less stellate, with numerous subcircular openings in the
angles between adjoining plates.
REVISION OF PALEOZOIC STELLEKOIDEA. 139
Inframarginal plates bounding the entire outer edge of the animal,
conspicuous, closely adjoining, not stellate and common to both the
actinal and abactinal areas.
Madreporite radially striate, abactinal.
Adambulacrals closely adjoining the inframarginals in the distal
portion of the rays but proximally the former are separated from the
latter by more or less numerous, small, accessory interbrachial plates. '
The adambulacrals may be of nearly the same size, or considerably
wider than long in the mid-length of the column. The adambula-
crals of adjoining columns meet orally in a pair of larger triangular
plates, the oral armature.
Interbrachial areas more or less large and the space between the
inframarginal and adambulacral columns filled with a variable
number of accessory interbrachial plates.
Ambulacrals slightly alternating, about as numerous as the adam-
bulacrals, each with an L-shaped, proximally converging carina.
Podial openings through the outer proximal corner of the plate and
proximal to the carina. Grooves narrow, regularly tapering.
GenoJiolotype (type by monotypy). Petraster rigidus Billings. The
above diagnosis is based on the genotype and Pal&asterina spedosa
Miller and Dyer.
Distribution. Ordovicic of North America, Siluric of Australia.
The species are:
P. rigidus (Billings). Trenton.
P. speciosus (Miller and Dyer). Maysville, Kichmond.
P. (?) americanus (D'Orbigny). Maysville.
P. smythi McCoy. Siluric of Australia.
'Remarks. While Billings has correctly pointed out the diagnostic
characters of Petraster, yet the genus has been very imperfectly
understood. This may partly be accounted for in that he later also
included a specimen of Hudsonaster matutinus (Hall), which of course
fails to have' the essential generic character of Petraster, namely, the
interbrachial accessory plates. The following is Billings' original
description:
"This genus has both marginal and adambulacral plates, with a
few disk-plates on the ventral side. The general form is deeply
stellate, and the rays long and uniformly tapering. A single specimen
has been collected, and as it shows the underside only, the characters
of the dorsal surface can not be given. The structure of the mouth
is also unknown. Generic name from petra, a stone.
"It differs from Palasterina by the presence of large marginal
plates outside of the disk-plates [that genus has small marginals],
and still more from Stenaster, which has neither disk nor marginal
plates. It is allied to Astropecten."
140 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Professor Hall at first recognized the distinctness of Petraster, since
he wrote as follows (1870):
"The character of Petrasier here described, and illustrated in the
figure (3a, pi. 9), gives an intercalated partial range of disk-plates,
between the adambulacral and marginal plates, which will separate
these forms from any of the Palaeasters in my collection.' 7
On a later page, after an examination of the genotype, he added
a "Note on the genus Petraster" in which he concludes that the
specimen was injured during life, as shown by the unequal distribu-
tion of the accessory interbrachial plates, and further that one ray
is "without intercalated plates on either side.' 7 He adds: "This
yiew is sustained by the fact that the other parts have the ordinary
structure of Palseaster, and in all other respects the specimen agrees
with the typical Palseaster matutina." The genotype was studied
at Ottawa by the present writer and found to be as described by
Billings. The accessory interbrachial plates, though somewhat
irregular in development in different areas in the type species of
Petraster, are normal in position and a similar but more complete
development also appears in Lindstromaster and Palasterina.
Petraster is distinguished from Palasterina as follows: The most
important feature is that the inframarginal plates of the former
are prominent, closely adjoining and short but wide, while in Palas-
terina they are small, not conspicuous, and globular, or subquadrate.
The next important difference is on the abactinal disk, where in
Palasterina there is an irregular ring of prominent basal radial,
and supramarginal plates. These are absent in Petraster. The acces-
sory interbrachial ossicles in the latter genus never attain the distal
region of the rays as in Palasterina where these pieces are also more
numerous. Palasterina is also more abundantly spinose than
Petraster.
Schuchertia has no inframarginals as marginal plates, which at
once distinguishes it from both Petraster and Palasterina. It is
true that in Schuchertia inframarginals are also present but they are
usually not well developed and remain adjoining the adambulacrals.
These two columns are therefore not separated from one another by
accessory interbrachial plates as in Petraster and Palasterina.
The primary structure of Petraster is that of Hudsonaster. So many
of the generic characters are common to both that the former genus
appears almost certainly to have been derived through the latter.
It is true that the central abactinal area of the disk in Petraster is
devoid of the large basal plates, a marked character in Hudsonaster,
but the other generic characters are all present in the former. Petras-
ter adds more or less numerous columns of radial accessory ambital
and accessory interbrachial plates, none of which are present in
Hudsonaster. It is these accessory ossicles which differentiate these
OF PALEOZOIC STELLEKOIDEA. 141
genera and show the line of development from the deeply stellate
primary form to the pentagonal genera with well-developed inter-
brachial arcs.
PETRASTER RIGIDUS (Billings).
Plate 27, fig. 5.
Palasterina rigidus BILLINGS, Geol. Surv. Canada, Rep. of Progress, 1853-1856,
1857, p. 291.
Petraster rigidus BILLINGS, Geol. Surv. Canada, Can. Org. Hem., dec. 3, 1858,
p. 80, pi. 10, fig. 3a (not fig. 3bHudsonaster matutinus} . WRIGHT, Mon.
British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palaeontogr. Soc. for 1861),
1862, p. 29. HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p.
294; rev. ed., 1868=1870," p. 337.
Description of 1858. "This species has much the aspect of an
Astropecten; the disk is one-fourth the whole diameter, the rays
rather slender and uniformly tapering, the angles between the bases
of the rays rounded. The plates [of the actlnal side] which appear to
be adambulacral [increase very little in size from the tips of the rays
toward the mouth], are quadrate and a little convex; [the adambula-
cral columns terminated in the mouth area by a pair of pointed
oral plates and not by a single plate as shown in the original illustra-
tion], the marginal [inframarginal] plates oblong, and also convex
[certainly not less than 16 and probably 20 in each column, increasing
rapidly in size toward the axils, whsre there is a single large axillary
plate] ; the disk plates [accessory interbrachials] consist of three at
each angle [one orally and two distally], and a single row [of not more
than seven plates] on each side of the ray, but extending only one-
third or one-half of the length of the ray; they all lie between the
[infra] marginal and adambulacral plates. [Abactinal side unknown.]
The specimen figured was about 2 inches [or 50 mm.-] in diameter
when perfect ; width of disk half an inch, and of rays at the base about
three lines."
Formation and locality. Trenton limestone, Ottawa, Canada.
Holotype No. 140 la is in the Victoria Memorial Museum, Ottawa.
The species has also been identified by Springer in the Lower Trenton
(Kirkfield) at Kirkfield, Ontario.
Remarks. Hall and Billings discussed their asterid genera and
species at different times and finally the former examined Billings' s
material. In this connection Hall showed that figure 3& of Petraster
rigidus was based on the actinal side of Hudsonaster matutinus.
Regarding figure 3a, which is the holotype of this species, he in 1870
wrote as follows:
"The specimen illustrated in figure 3a has a few small intercalated
plates between the marginal and ambulacral [adambulacral] ranges
in two of the axils of the rays, and there are a smaller number of gran-
ules in a similar position but unequally distributed on one side of
142 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
another axil, while two of the axils do not show any such intercalated
plates or granules. In one of the rays, at least, the ambulacral, adam-
bulacral, and marginal plates are distinctly visible, without inter-
calated plates on either side.
"The presence of these unequally distributed plates or granules
is apparently an abnormal structure, probably the result of accident
during the growth of the animal; and this view is sustained by the
fact that the other parts have the ordinary structure of Palseaster,
and in all other respects the specimen agrees with the typical Palse-
aster matutina"
The holotype was examined by the writer and although imperfect
was found to have a normal development, with the plates in their
proper places in at least two of the rays and axils. It certainly is not
an abnormal development of "Palseaster matutina" but represents
a normal individual of a distinct genus and species diverging from
Eudsonaster matutinus toward the forms with large disks and great
axillary areas, as in Lindstromaster and Palasterina. Hall seemingly
was led to make the erroneous remarks above quoted through the
fact that Billings had also included in his description a'nd illustration
(36) an individual of Hudsonaster matutinus, an error due to the
undoubtedly close relationship of this form with Petraster rigidus.
PETRASTER SPECIOSUS (Miller and Dyer).
Plate 23, figs. 5-7; plate 26, fig. 1; plate 27, figs. 1-4.
Palssasterina spedosa MILLER and DYER, Journ. Cincinnati Soc. Nat. Hist., vol. 1,
1878, p. 30, pi. 1, fig. 7. MILLER, N. Amer. Geol. Pal., 1889, p. 266, fig. 381.
Paldeasterina approximata MILLER and DYER, Journ. Cincinnati Soc. Nat. Hist.,
vol. 1, 1878, p. 30, pi. 1, fig. 8.
Original description of Palseasterina spedosa. "Pentagonal; rays
obtuse at their apices; greatest distance from point to point about
2J inches; breadth of body between rays about 1J inches, and dis-
tance from tip of ray to next adjoining tip on either side about 1J
inches.
"The [infra] marginal plates are small and somewhat hemispher-
ical near the termination of the rays, they gradually enlarge and
become square, and then rectangular as they approach the disk, until
at the narrowest part of the disk or body of the fossil they are twice
as long as wide. There are about 50 marginal plates between the
apex of one ray and the next one adjoining, or, in a perfect specimen
of this size, about 250.
"The back or dorsal side is covered with numerous plates (prob-
ably in a complete specimen of this size there would be 1,000 or more),
which are very prominent in the center or somewhat conical, and
seem to have been joined together with deeply serrated edges.
The plates have from three to eight of these indentations,, which
give them a beautiful starlike appearance.
KE VISION" OF PALEOZOIC STELLEROIDEA. 143
"The ambulacral grooves are narrow and deep, as shown by the
sharp ridges on the back of the specimen. The small dorsal plates
which cover the ambulacral pieces are exfoliated in some places,
and show two rows of ambulacral plates coming evenly together
and forming the sharp ridge.''
Original description of Palseasterina approximate, . " Pentagonal;
rays more slender than in P. speciosa; greatest distance from point
to point in the specimen figured 1.15 inches and breadth of the
body one-half inch; another specimen from the collection of Mrs.
M. P. Haines, of Richmond, Ind., measures from point to point 2.5
inches, and has a breadth of body of nine-tenths inch.
"The [infra] marginal plates have about the same form, and are
about as numerous as they are in P. speciosa. The adambulacral
plates are arranged with their length across the rays. The oral
plates are very prominent. [This statement is somewhat in error.
The five prominent points around the mouth are each formed of one
large interbrachial plate and a pair of proximal adambulacrals.] The
space between the [infra] marginal plates and the oral ones [adambu-
lacrals] is filled with numerous small [accessory interbrachial] pieces.
The ambulacral grooves are narrow. Ambulacral ossicles unknown.
"The dorsal side is covered with small conical pieces, which give
it a coarsely granular appearance. The madreporiform tubercle
is prominent, conical, and longitudinally striated.
"This species is distinguished from P. speciosa by its narrower
rays, more contracted body, and smaller dorsal plates."
Emended description. The holotypo of P. speciosa measures:
R = 33 mm., r 18 mm. The holotype of P. approximata measures:
R = 15 mm., r = 6.5 mm.
Rays in young specimens slender distally, but with age they
become obtuse and very wide proximally and here merge into the large
disk. Interbrachial arcs very large.
The abactinal area of disk and rays is bounded by columns of very
prominent inframarginal plates. Beginning at the apex of each ray
they are very small and globular, soon they become quadrate and
gradually shorter but wider, so that near the middle of each interbra-
chial arc they have a crowded appearance. In a full-grown specimen
there are about 50 and hi a half-grown example about 45 of these
plates between the distal ends of adjoining rays. In the distal region
these plates are rounded and nearly smooth but gradually become
granular and strongly convex, each with one prominent point for an
articulating spine, none of which, however, have been observed. On
the actinal side these inframarginals are also very prominent and
pustulose but here are flat. They are most prominent near the center
of the interbrachial arcs.
50601 Bull. 88 15 10
144 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
The supramarginal columns begin at the very apex of the rays
and for more than one- third of their length adjoin the inframarginals
where ambital plates begin to appear. The supramarginal columns
are continuous over the disk in a broad curve, with about 18 to 20
in a column, or from apex to apex of adjoining rays there are from
35 to 40 of these plates. In the axillary areas they are separated
from the inframarginals by three or four rows of ambital plates. At
the apex of the rays the supramarginals are highly convex and
nearly circular in outline but soon pass into more and more dis-
tinctly stellate plates. This is the form of all the abactinal plates
inside of the inframarginals ; they are stellate, highly convex, each
with a central node for an articulating spine and a few granules
that are the bases for smaller spines.
Each radial column appears immediately beneath the two terminal
or rather distal supramarginal plates and then continues as a column
to near the center of the disk. They are more prominent than the
other columns excepting the marginals and have not less than 25
plates in each one. On each side of the radials are five columns of
radial accessory plates which appear to continue as columns over the
disk in broad curves joining those of the next ray. Between these
columns in the axillary areas are additional accessory plates. These
columns of accessory plates appear singly, first on one side and later
on the other, and not in pairs simultaneously, one on each side of the
radial columns.
Ambital areas well developed in the axillary region, where there
are about three columns of these plates. The columns pinch out
rapidly dis tally and none are present in the outer third of the rays.
The plate arrangement of the central part of the disk can not be
made out.
Madreporite of medium size, highly conical, and conspicuous,
with numerous sharp single or bifurcating ridges; on the under side
are seen two outwardly directed spiral cones (the white lines of the
drawing representing the spiral tubes in the madreporite), reminding
one of the brachia in Atrypa (pi. 27, fig. 4).
Adambulacral plates depressed, convex, subquadrangular in out-
line in young specimens, but in fully grown individuals a number of
these in the central region of the column are much drawn out laterally
and are here two or three times as wide as long. In a half-grown
individual there are about 20 of these plates in a column, but in a
mature specimen there appear to be not less than 26. These plates
end inwardly in an apex, against which terminate the ridges of the
ambulacral plates. The adambulacral and inframarginal columns
are closely adjoining in the distal two-thirds of the rays but proxi-
REVISION OF PALEOZOIC STELLEROIDEA. 145
mally they are separated from each other by numerous accessory
interbrachial plates, of which there appear to be not less than 30 in
each area. The apexes of these interbrachial areas form the oral
armature and each consists of two adambulacral plates, back of
which is a rather large interbrachial plate.
Ambulacra! grooves narrow, tapering, and deeply V-shaped.
Ambulacral plates slightly alternating, about as long as wide, and
proximally superposed, with about 26 in each column. Each plate
has an L-shaped ridge with the point proximally directed. The
podial openings are situated in the outer proximal corner of the
plates and between the inner angles of two adambulacrals.
Formation and locality. Richmond and Maysville formations of
the Upper Ordovicic. The holotype of P. speciosa was found by
Mr. W. C. Barnhart, on Twin Creek, near Winchester, in Preble
County, Ohio, and is now in the Dyer collection of the Museum of
Comparative Zoology, Harvard University (No. 14). Four other
less well-preserved specimens are in the Harris collection of the
United States National Museum and were found in the vicinity of
Waynesville, Ohio. The holotype of P. approximata and another
similar-sized specimen but not so well preserved are also from the
vicinity of Waynesville, Ohio. The first is in the Dyer collection
of the Museum of Comparative Zoology (No. 15), and the other in
the United States National Museum. There is a slab with two
young individuals much etched with acid but otherwise excellent,
showing the actinal side, in the Gurley collection of the University
of Chicago (No. 10979). Finally there are two fine specimens, not
fully grown, showing well the actinal and abactinal sides, in the
Haines collection of the same university (Nos. 10839 and 10840),
found near Hamilton, Ohio; these are apparently from the very top
of the Maysville formation.
Remarks. P. speciosa is described by Miller and Dyer from the
abactinal side and P. approximata from the actinal side. These
writers point out that the rays are more slender and the disk more
contracted in the latter, but when one considers that it is less than
half the size of the former and that during growth these forms con-
tinually add accessory plates, this difference hi form is seen to be
one of growth. The ' 'smaller dorsal plates" noted by these writers in
Mrs. Haines's specimen of P. approximata are a character which the
present writer does not regard as of value unless they be constant
at this lower geologic horizon (Maysville at Hamilton), in which event
this specimen can later on be distinguished by another specific name.
Cat. No. 60609, U.S.N.M.
146 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
PETRASTER (?) AMERICANTJS (D'Orbigny).
Plate 26, fig. 2.
Asterias GRAHAM, ANTHONY, and JAMES, Amer. Journ. Sci., eer. 2, vol. 1, 1846,
p. 441, figure but no description.
Ccelaster americanus D'ORBIGNY, Prodr. de Paleont., vol. 1, 1849, p. 22 (name
proposed).
Asterias anikonii DANA, Manual Geol., ed. 1, 1863, p. 221, with figure but no
description.
Palasterina (tyjamesi DANA, Amer. Journ. Sci., ser. 2, vol. 35, 1863, p. 295; Manual
Geol., ed. 2, 1864, p. 221, with figure but no description.
Palseaster jamesi HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 286;
rev. ed., 1868=1870, p. 329. MEEK, Geol. Surv. Ohio, Pal., vol. 1, 1873,
p. 62, pi. 4, fig. 4.
Meek's description. " Through the kindness of Mr. U. P. James,
of Cincinnati, the discoverer of this fossil, I have now before me the
original typical specimen, which I found not in a condition to be
fully and systematically characterized, nor perhaps to afford the
means of determining, beyond doubt, its generic affinities. It
evidently attained a large size, and has a comparatively large disk,
with its five rays rather broad at their inner ends, somewhat rapidly
tapering, and apparently longer than the diameter of the disk. It
evidently lies [does lie] in the matrix in such a manner as to show the
ventral side [the inner dorsal aspect of the ventral plates], but pre-
sents the appearance of having the marginal and disk plates of this
side removed [they are present]. The five pairs of oral pieces [proxi-
mal adambulacral plates] are seen in place, and proceeding outward
from these can be seen a row of adambulacral pieces on each side
of each of the rather wide [narrow] ambulacral furrows. These
latter pieces are [proximally] nearly three times as long as their
diameters in the direction of the length of the rays, and quite promi-
nent at their inner ends, where they are articulated together by little
processes and corresponding sockets or sinuses; while the outer
two-thirds of each is flattened and distinctly more depressed than
their inner ends. Outside of, and alternating with, the outer flat-
tened ends of these adambulacral pieces, a row of [inframarginal]
nearly square, or more or less oblong pieces, may be seen all along the
margins of the rays, and connecting at their axilla in such a manner
as to leave room for several disk [or interbrachial] plates between
them and the oral and inner adambulacral pieces."
In each interbrachial area there are inserted between the infra-
marginals and adambulacrals interbrachial plates as follows: Proxi-
mally one hexagonal plate upon which rest two similar plates mar-
gined by the inframarginals.
"At some places within the [narrow] ambulacral furrows, remains
of ambulacral ossicula can be seen deeply interlocking with the
inner ends of the adambulacral pieces, but their exact form, and the
REVISION OF PALEOZOIC STELLEROIDEA. 147
position and arrangement of the ambulacra! pores, can not be
clearly made out in the specimen studied.
"The two oral pieces [proximal adambulacrals] of each of the
five pairs are separated by deep sutures, and have an irregular form,
being longer than wide, and, like the adambulacral pieces, which
they somewhat exceed in size, flattened and depressed at their outer
ends, and elevated into crestlike prominences farther in, with a
lateral process or thickening on the outer side of each, in which a more
or less distinct pit may be seen." Between the pairs of oral adam-
bulacral plates are inserted small, single, sub quadrangular plates,
the proximal plate of each pair of ambulacral ossicles. Orad to
each pair of oral armature plates are situated single hemispheric
plates.
"The surface, and the arrangement of the dorsal parts, are un-
known; but they were probably armed with short spines, as some
remains of little spines are seen in the matrix at the margin of the
disk, at one place.
"As the extremities of none of the rays can "be clearly seen, the
exact breadth of the whole fossil, between the ends of opposite
rays, can not be exactly given, but it is evident that the length of
one of the rays, measuring from the middle of the disk, must have
been a little more than 2 inches, and consequently that the diam-
eter across from the extremities of opposite rays could not be less
than about 4 inches. The breadth of the disk is 1.20 inches;
and that of the rays at their inner ends, 0.70 inch."
Formation and locality. The single specimen was found in the
Maysville formation (Ordovicic), by Mr. U. P. James, on or near
Vine Street hill, Cincinnati, Ohio, and is now in the James collection
of the Walker Museum of the University of Chicago.
Remarks. This large species is quite distinct from any other star-
fish in having three rows of interbrachial plates, one piece in the proxi-
mal, two in the medial, and five or possibly more in the distal row.
While the arrangement of these interbrachial plates is different than
hi the other species of Petraster, their position is that of this genus.
Therefore this species is for the present referred to Petraster. The
preservation of this specimen is such that its generic and specific
characters will remain hi obscurity until other material is found.
It lies with the actinal side buried in the limestone while all of the
abactinal plates have been worn away. What one sees is therefore
the inner or dorsal aspect of the ventral skeleton.
PETRASTER SMYTHI McCoy.
Petraster smythi McCoy, Geol. Surv. Victoria, Prodr. Pal. Victoria, dec. ], 1874,
p. 41, pi. 10, figs. 1-16.
Original description. "Five broad semielliptical lobes meeting
at slightly rounded reentering angles, leaving the length and the
148 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
width at base of the rays nearly equal and less than the width of the
disk. The upper surface is covered with crowded irregularly polyg-
onal tumid plates. Madreporiform tubercle very large (1J lines in
diameter), irregularly porous, and rugged with branching vermicular
ridges, excentric toward base of the two posterior rays. Ambulacral
groove very narrow, bordered with a row of large transversely oblong
adambulacral plates, wider than long, about 6 in 2 lines at middle
of ray; margin of the rays bordered with a rather smaller row of
similar marginal plates; between the row of adambulacral and mar-
ginal plates an intercalary row of small irregular plates. Width of
disk between the rays, 7 lines; from tip to tip of rays, about 1 inch 2
lines; length of ray, about 5 lines.
" Very rare in the fine sandy Upper Silurian rocks of Moonee Ponds,
Flemington, a little north of Melbourne [Australia]. ' '
Remarks. The holotype exposes the abactinal side, but the plates
of two of the rays are absent, exposing the actinal skeleton, which is
therefore seen from its inner side. So far as one can judge of the
description and illustration, the species is a small but genuine Petras-
ter. Abactinally the rays have conspicuous radial and supramar-
ginal columns, each with about 17 ossicles, that are tumid and are
all closely adjoining. Outside of these in the distal portion of the
rays are the equally conspicuous inframarginals, and they margin
not only these parts but the entire animal; proximally the inframar-
ginals separate more and more from the supramarginals, forming
small ambital areas, occupied by a number (can not be determined)
of rounded plates that now appear not to have been closely adjoining.
Madreporite large, radiately striate, situated in an interradius and
well inside the ambital area.
Ambulacralia large, about 12 in a column, and opposite one another.
Adambulacrals smaller, about 14 in a column. Interbrachial struc-
ture unknown.
Genus LINDSTROMASTER Gregory.
Lindstromaster GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, pp. 344, 346.
Hisingeraster STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
pp. 224, 225 (same genoholotype as for Lindstromaster).
Original description. "Palseasterinidae with flat pentagonal disk
and five short blunt rays.
''The plates of the disk are large, polygonal, tuberculate, and
crowded into a close tessellate pavement, which completely covers
the interradial areas.
REVISION OF PALEOZOIC STELLEROIDEA. 149
"Marginal [inframarginal] plates well developed and conspicuous,
and but slightly smaller than the adambulacrals. Both supra- and
inframarginals appear to be present.
"The abactinal [actinal] furrows are large and subpetaloid. They
are bounded by large tuberculate adambulacral plates. The ambu-
lacral plates are boot-shaped [and slightly alternating]. The pores
for the podia are large, and occur on the suture between the adam-
bulacrals and ambulacrals.
"The adambulacral elements in the oral armature are prominent,
and consist of five pairs of subtriangular plates.
"Type-species: Asterias antiqua, Hisinger 1837. Silurian: Got-
land." No other species of this genus is known.
Remarks. Of this genus only the actinal side is known and the
published figure from a drawing by Mr. George Lilejevall is one of
the best of Paleozoic starfishes. Lindstromaster differs from Petras-
ter in but one generic character, if it be a generic one, in that the
ambulacral furrows are large and subpetaloid in the former and
narrow and slowly tapering in Petraster. However, since the abac-
tinal area is unknown in Lindstromaster there probably will be dis-
covered in this area differences distinguishing it from Petraster. It
is also probable that the marginals in Lindstromaster which Gregory
thinks are "both supra- and inframarginals" lying directly super-
posed are only inframarginals. In Siluric starfishes these plates
have not attained the superposed position so marked in recent genera.
The supramarginals are here never true marginals and always lie
more or less inside the inframarginals, which are the true marginals of
these early genera. For other remarks see Palseostella.
The name Hisingeraster was published after Gregory's name and
is based on the same genotype.
When more material is obtained and the genus is restudied, com-
parisons should also be made with Palasterina primseva, as the two
forms appear to have much in common.
LINDSTROMASTER ANTIQUT7S (Hisinger).
Asterias antiqua HISINGER, Lethsea Suecia, 1837, p. 89, pi. 26, fig. 6.
Lindstromaster antiqua GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, pp. 343, 347,
pi. 36, figs, la, 16 (complete synonymy given here). SCHONDORF, Jahrb.
nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 225.
A single specimen is known from "Bed C," Mount Hoburg, Got-
land, corresponding to the Wenlock of England. It is in the Angelin
collection of the Mineralogical Museum at Copenhagen.
150 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genus PALASTERINA McCoy.
Palasterina McCoY, British Pal. Foss., 1851, p. 59 (not defined). SALTER, Ann.
Mag. Nat. Hist., ser. 2, vol. 20, 1857, pp. 324, 327. BILLINGS (part), Geol.
Surv. Canada, Can. Org. Rem., dec. 3, 1858, p. 76. ZITTEL, Handb. Pal.,
vol. 1, 1879, p. 453. STURTZ (part), Palseontographica, vol. 36, 1890, p. 226;
(part), Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 43, 60.
SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910,
p. 220.
Palxasterina GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 349 (complete
synonymy here).
The term Palasterina was used by McCoy in the following words:
"Before I was aware Prof. Forbes had described them [the three
species named below] it seemed to me that the U [rasterella] ruthveni
and U. Jvirudo, as well as the similar American species, might be easily
separated from the great starfishes forming the recent genus Uraster,
by their small size and much more simple skeletons, and I had named
the genus Urasterella in my manuscript. The U. primdevus I thought
generically distinct from the other two, as the rays were not contracted
at base, etc.; and I had named it Palasterina, from its resemblance
to the recent genus Asterina."
The three species mentioned by McCoy had been described by him
in manuscript, but on learning that descriptions had been previously
published by Forbes, the former, in his "British Palaeozoic Fossils/'
adopted the names of Salter. In writing of these three forms, as
above quoted, he uses the generic name Palasterina (not Palseasterina,
the generally accepted form), and while a type-species is selected, yet
McCoy did not point out a single generic character of present value.
His statement that "the rays were not contracted at base" is a
feature which at that time distinguished Palasterina from Urasterella,
but now many Paleozoic genera are known having rays "not con-
tracted at base."
While McCoy does not strictly propose the generic name, Salter is
the first to accept and define Palasterina, which he does in the following
words :
"Pentagonal, depressed, the arms a little produced, with three or
five principal rows of tubercles above, combined with a plated disk
which fills .up the angles ; ambulacra rather shallow, of subquadrate
or slightly transverse ossicles, bordered by a single row of squarish
large plates, the lowest of which (ad-oral adambulacral plates, Huxley;
angle-ossicula, Forbes) are large and triangular, bearing combs of
spines (Upper Silurian)."
Recently Gregory, in a paper entitled "On Lindstromaster and the
classification of the Palseasterids" (1899), redefines the genus, basing
his definition largely on P. lonneyi. In this connection he writes:
"The species [P. lonneyi] is of interest, as the structure is better
preserved than in the type of the genus [P. primseva], and thus we are
KEVISION OF PALEOZOIC STELLEKOIDEA. 151
largely dependent upon it for our knowledge of the generic charac-
ters " (p. 350).
Emended description. Disk large, with moderately developed inter-
brachial arches. Rays five, short, and tapering more or less rapidly.
U R about twice as great as r."
Abactinal area of rays distally with one medial column of large,
thick, and subquadrangular radial plates, on each side of which are
similar columns of supramarginal and inframarginal pieces. In this
region the columns adjoin and the ossicles are crowded, but
proximally between the radial and supramarginal and between the
latter and the inframarginal columns (ambital areas) are inserted
numerous accessory smaller plates. The latter are most numerous
in the ambital areas and at the base of the rays. The ossicles are
packed into a tessellate pavement. In the center of the disk there
seems to be a prominent central disk piece and at the bases of the
rays there is a somewhat disconnected ring of ten most prominent
plates, the five basal pieces of the radial and five interradial that
give rise to the ten supramarginal columns. Inside of this ring there
are probably numerous, very small pieces with no definite arrange-
ment. A number of the proximal plates of both the radial and
supramarginal columns may be separated from one another by small
accessory pieces.
Madreporite abactinal.
Inframarginal plates bounding the entire outer edge and common to
both sides of the animal. These plates are, however, small (smaller
than the adambulacrals) and bear tufts of small spines.
Adambulacrals subquadrangular, and closely adjoining, each plate
bearing on its outer lateral edge a tuft of three spines. Oral armature
consists of five pairs of triangular adambulacral plates which have
slender spines on the inner edges.
Interbrachial areas extending to near the distal ends of rays and
occupied by numerous "isolated rounded or subangular [accessory
interbrachial] plates bearing [single] small spines."
Ambulacrals slightly alternating and about as numerous as the
adambulacrals. Each plate has an L-shaped ridge, posteriorly
directed. The podial openings are on the sutures between the
ambulacrals and adambulacrals.
Genoholotype. Uraster prims&vus Forbes. The above diagnosis is,
however, based largely upon Palasterina bonneyi Gregory. Dr.
Bather sent the writer a number of wax squeezes of the genoholo-
type, and while the parts preserved are clear and sharp, yet the speci-
mens are not complete enough to work out the generic characters.
Therefore, for the present, P. bonneyi is taken as the genoholotype.
Distribution. Ludlow shales (Upper Siluric) of Great Britain.
152 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Contains :
P. primseva (Forbes). Ludlow of England.
P. bonneyi Gregory. Ludlow of England.
P. (?) ramseyensis Hicks. Tremadoc of Wales.
For other remarks see Palseostella.
Since the foregoing was written, Schondorf 's work cited above has
come to hand. He studied wax squeezes of P. primseva and deter-
mined that the genotype is devoid of marginalia, and therefore refers
Palasterina to the Cryptozonia. He writes: " Special marginalia ap-
pear not to be present, at least the marginal rows are in no way dis-
tinguishable from the other intermediary [accessory interbrachial]
plates. Accordingly Palasterina primseva Forbes sp. does not belong
as stated by Gregory (p. 349) to the asterids with distinct marginal
plates, the Phanerozonia, but with the asterids having inconspicuous
or no marginalia, the Cryptozonia' 7 (p. 222).
The present writer holds that there are inframarginalia present all
along the edge of the interbrachial areas and the rays, and that they
are most typical along the distal portion of the rays. Here the
inframarginalia are ontogenetically youngest and less affected by
subsequent growth displacement and alteration than are the older
ones in the interbrachial areas. In the latter region it is true that
the marginalia are not conspicuous, but so long as one can make
them out to be inframarginalia, size alone can not be the criterion
that will permit of the conclusion that no marginalia are present in
Palasterina. However, the difference in interpretation between
Schondorf and the writer is not a great one, as neither lays great
taxonomic value on whether the marginalia are prominent or u not
prominently developed" (p. 223). So long as the inframarginalia are
more or less prominent, one of the generic characters of Palasterina,
a phanerozonian, is present, and when they are so inconspicuous as
not to be made out among the other ossicles, we have one of the
generic characters of Schuchertia, a cryptozonian.
Schondorf also states that the ambulacralia, and the adambulacralia
as well, are all opposite and do not alternate with one another. The
present writer states that the ambulacrals are " slightly alternating, " H
and throughout this work he has given the arrangement of these
ossicles his special attention. In a great many cases, and especially
in the Ordovicic species, he has had difficulty in ascertaining what
was their actual position in the living animals, for nearly all the
fossils have been squeezed more or less. He has always given their
arrangement as they impressed him in the fossil when no evident dis-
tortion could be seen. The writer would not lay special stress upon
slight alternation among the ambulacralia, for in many Ordovicic
species there is some variation in each ray from opposite to slightly ,
alternate. Of course, when the alternation or opposite arrangement
REVISION' OF PALEOZOIC STELLEROIDEA. 153
is complete and fixed throughout the rays, he would lay great value
on the character. It may be said that in the earliest Asteroidea the
chronogenetic tendency is from slight alternation to complete and
fixed opposite arrangement among the amhulacral ossicles.
PALASTERINA PRIM^EVA (Forbes).
Uraster primxvus FORBES, Mem. Geol. Surv. Great Britain, vol. 2, pt. 2, 1848,
p. 463; Mem. Geol. Surv. United Kingdom, dec. 1, 1849, p. 2, pi. 1, figs.
2a, 26; in McCoy, Brit. Pal. Foss., 1851, p. 60. MURCHISON, Siluria, 1854,
p; 221, fig. 39.
Palasterina primxvus McCoy, Brit. Pal. Foss., 1851, p. 59 (nomen nudum).
SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 327, pi. 9, figs. 2a-2c.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palseontogr.
Soc. for 1861), 1862, p. 26, fig. 16a. QUENSTED, Petrefactenkunde Deutsch-
lands, vol. 4, 1876, p. 74, pi. 92, fig. 35. STURTZ, Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, p. 44; vol. 56, 1900, pp. 214, 224. SCHONDORF,
Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 220.
Palszasterina primseva GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 349. SPEN-
CER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, pp.
37, 38, fig. 30.
Formation and locality. A common species in the Ludlow rocks.
Underbarrow, Kendal, Westmoreland, and Leintwardine, Shropshire,
England.
Remarks. Dr. Bather furnished the author with wax squeezes of
specimens Nos. E4990-E4993 (E61 is quite another species and
abactinally more closely related to Eudsonaster) in the British
Museum (Natural History). They are from the Lower Ludlow at
Kendal, Westmoreland. These show little of value abactinally, but
actinally are well preserved along the ambulacra. Unfortunately,
however, none of the marginals are present and but little of the
interbrachial areas. When better material is at hand a comparison
should be made also with Lindstrb'master antiquus (Hisinger), as the
two look suspiciously alike.
PALASTERINA BONNEYI Gregory.
Palseasterina bonneyi GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, pp. 349, 350,
text figs. 1-3, and pi. 16, figs. 2a, 26. SCHONDORF, Jahrb. nassauisch. Ver.
Naturk., Wiesbaden, vol. 63, 1910, p. 223.
"This species has hitherto been included in P. primseva, the type-
species of the genus, from which it differs by having shorter and
blunter rays, more numerous actinal interbrachial plates, and more
adambulacral plates in each series ; the character of the abactinal ray
plates is also different in the species, the longitudinal series being
widely separated at the proximal end of the rays in P. bonneyi"
(Gregory).
Formation and locality. Ludlow shales. Leintwardine, Shrop-
shire. Types in British Museum (Natural History).
154 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
PALASTERINA (?) RAMSEYENSIS Hicks.
Palasterina ramseyensis HICKS, Quart. Journ. Geol. Soc. London, vol. 29, 1873,
p. 51, pi. 4, figs. 21-23. WOODWARD, Geol. Mag., dec. 2, vol. 1, 1874, p. 96.
This starfish needs to be restudied in the light of modern knowl-
edge. The specimens represented by Hicks's figures 21 and 22 seem
to be referable to Petraster rather than to Palasterina, while that of
figure 23 appears to be nearer Mesopalseaster.
Formation and locality. Said to be from the Tremadoc, but the
development of the starfish seems to be in harmony with Upper
Ordovicic time. The specimens are from Ramsey Island and St.
Davids, Wales, and are in Mr. Lightbody's collection.
Genus URANASTER Gregory.
Uranaster GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 347. SCHONDORF, Jahrb.
nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 225.
The name is "from Uranus, the grandfather of Pluto; the starfish
resembles the living genus Plutonaster."
Generic description. Animal deeply pentagonal, above medium size,
with well-developed interbrachial arches, and five long slender rays.
Inframarginal plates marginal, large, and elongated in the direction
of the rays. These plates are apparently common to both the actinal
and abactinal areas. Abactinally adjoining the inframarginals is
another column of large submarginal plates ( ?supramarginals)
alternating with the former, and sometimes there are smaller acces-
sory pieces between these columns in the angles of the plates. The
rays have no medial or radial columns of prominent plates, but the
entire space between the submarginal columns is occupied by numer-
ous closely fitting, convex, or tubercular plates which are arranged
in alternate transverse rows, each having from three to four pieces
on each side of a ray.
Interbrachial accessory plates numerous, polygonal, and, according
to Baily 's figure of U. Tcinahani, arranged in crescentic rows conforming
to the shape of the disk margin. These plates do not appear to
extend between the marginals and adambulacrals beyond the mid-
length of the rays.
Adambulacral plates tubercular, subquadrangular in outline, and
widest transversely in the medial third of the column.
Ambulacral furrows very narrow, deep, and tapering gradually.
Ambulacra! plates traversed by strong transverse ridges.
Madreporite unknown.
GenoJiolotype. Palasterina Icinahani Baily. Upper Ordovicic. A
second species is U. elizse, Spencer, also from the Ordovicic of Great
Britain.
Remarks. Uranaster differs from Petraster, Palasterina, and
Lindstromaster in having the abactinal area of the rays between the
REVISION OF PALEOZOIC STELLEROIDEA. 155
two pairs of marginal columns filled with small tuberculate plates
arranged in transverse rows. The other genera have the plates in
longitudinal columns.
URANASTER KINAHANI (Baily).
Palasterina kinahani BAILY, in KINAHAN, Man. Geol. Ireland, 1878, pp. xv, 26,
pi. 2, fig. 1; Mem. Geol. Surv. Ireland, Expl. Mem. Sheets Nos. 169, 170,
180, 181, 1879, pp. 58, 59, fig. 6.
Uranaster kinahani GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 348. SCHONDORP,
Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 225.
This species occurs abundantly as natural molds in the Caradoc
(Upper Ordovicic), near Ballymadder Point and Bannow, Wexford,
Ireland. A specimen is in the British Museum (Natural History),
No. E 194.
URANASTER ELIZAS Spencer.
Uranaster elizx SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palaeontogr. Soc. .for
1913), 1914, pp. 18, 20, 23, figs. 18, 23.
From the Ordovicic of Great Britain.
Genus PAL^OSTELLA Stiirtz.
Palxostella STURTZ, Palseohtographica, vol. 36, 1890, p. 230, pi. 31, figs. 42, 43;
Verb, naturb. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 42, 58, pi. 1, figs.5^8,
Palsenectria STURTZ, Verb, naturb. Ver. preuss. Rbeinl., etc., vol. 50, 1893.
p. 12 (genoholotype the second specimen of Palseostella solida).
Description. Animal attaining a large size, with large pentagonal
interbrachial arcs. Rays five, stout. Ambulacral grooves wide and
apparently regularly tapering.
Abactinally the inframarginals are also the marginals for this
area. Near the distal ends of the rays the supramarginals adjoin
and lie either partially over or inside of the inframarginals and
continue nearly to the axillae. There also seems to be a single radial
column, confined, however, to the distal third of the rays. Other-
wise the abactinal skeleton consists of very numerous, small, irregu-
larly shaped, loosely interlocking plates without definite arrangement.
Madreporite abactinal.
Actinally the animal is bounded by subquadrate, numerously
spine-bearing, inframarginal plates which increase rapidly in size
towards the axillae, where they are ponderous and nearly twice as
wide as long. Ambulacral plates alternating, L-shaped, with single
columns of podial openings. Adambulacrals subquadrate, prominent,
increasing regularly in width proximally and bearing spines like
the marginals. Interbrachial areas large, reaching to near the
distal ends of rays, widely separating the inframarginals from the
adambulacrals in the axillary region, and the space occupied by
very numerous, small, spine-bearing, apparently irregularly dis-
posed plates.
156 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genoholotype and only species. P. solida Stiirtz. Two speci-
mens are known from the Lower Devonic roofing slates, Bundenbach,
Germany.
Remarks. The starfishes from the Bundenbach roofing slates are
at best but poor fossils and were it not for the great skill of Mr.
Stiirtz in cleaning them out of the matrix their systematic position
could not well be determined. At best they are compressed to almost
paper thickness and distorted during solidification and shortening or
elongation of the slates. Under these circumstances it appears
best to redescribe the detailed abactinal characters as given by
Stiirtz. He mentions numerous paxillae and finds many characters
common to both Palseostella and the living Nectria.
Palseostella is closely related to Palasterina, also to Lindstromaster ,
and may be a descendant of the latter. The actinal structure in
Lindstromaster and Palseostella is nearly alike, both having large
inframarginals and adambulacrals, but the interbrachial areas in
the former are occupied by large closely adjoining plates, while in the
latter these are small and somewhat loosely interlocking. Abac-
tinally they can not be compared, since that side is unknown in
Lindstromaster. The greatest difference between Palasterina and
Palseostella is that the former has small inframarginals while in the
latter they are very large. Then the supramarginal and radial
columns in Palasterina are better developed and comparatively
larger, while in Palseostella they are shorter and most prominent in
the distal half of the rays. In the latter genus the central region
of the disk has no ring of fifteen basal radial and supramarginal
plates as in Palasterina.
From this evidence it seems probable that the line of descent is
from Petraster to Lindstromaster to Palseostella in marntaining promi-
nent columns of inframarginals, while the line from Petraster to
Palasterina tends toward the reduction or elimination of these plates
as a conspicuous skeleton element.
Palsenectria is based on the second specimen, and the genus is
described as follows: " Should the described starfish [the second
specimen of Palseostella solida Stiirtz] prove not to be identical with
Palseostella, which is possible, even though not probable the other
specimen is in the British Museum I propose for it the new name
Palsenectria devonica." Unless the author takes the time to work
out his new forms, or decides the problem presented for himself, he
should not expect some one else to do this for him. Under these
circumstances the genus and species are regarded as not defined.
Genus PSEUDOPALASTERINA Sturtz.
Pseudopalasterina STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
pp. 219, 224.
Genoholotype and only species. Palasterina follmanni Sturtz.
REVISION OF PALEOZOIC STELLEROIDEA. 157
PSEUDOPALASTERINA FOLLMANNI (Sttirtz).
Palasterina follmanni STURTZ, Paleeontographica, vol. 36, 1890, p. 226, pi. 29,
figs. 29-31a.
Pseudopalasterina follmanni STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc..
vol. 56, 1900, pp. 219, 224.
Actinally this species has ambulacrals and narrow adambulacrals
and outside of the latter the interbrachial and abactinal areas consist
of a membrane with small spine-bearing plates. It has no inframar-
ginal nor radial and supramarginal columns, features characterizing
Palasterina.
Formation and locality. Lower Devonic of Bundenbach, Germany.
NEW GENUS.
NEW SPECIES.
Prof. Weller got on Fountain Creek, at Waterloo, Illinois, in the
St. Louis limestone, a small pentagonal asterid (No. 14396, Univer-
sity of Chicago collection) showing only the actinal side, that differs
from all other known forms of the large-disked genera. The five rays
are completely within the pentagon, with the interbrachial areas large
and straight-sided. These areas are made up of a few large tesselate
plates, of which it seems that seven compose the interbrachial mar-
gins and inside these there seem to be transverse rows, each with five,
three, two, and one ossicles. It could not be made out whether the
inframarginals and supramarginals are directly superposed, but this
may be so.
Ambulacral furrows exceedingly narrow, a mere line. These are
bounded by columns of large, very ornate adambulacrals, each with
seven ossicles. Those of opposite columns alternate decidedly with
one another and on its outer side each bears two short but dis-
tinct spines.
Adambulacral oral armature in five very prominent pairs of pieces,
long and pointed, and about twice as long as any other adambulacrals.
Measurements: R = 8.5 mm., r = 7 mm.
Family LEPIDASTERIME Gregory (emend.).
Lepidasteridse GREGORY (part), Geol. Mag., dec. 4, vol. 6, 1899, p. 352 (includes
Lepidaster&n&Etheridgaster)', Treat. Zool., vol. 3, Echinoderma, 1900, p. 255.
Helutnihasterinse GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 258.
Helianihasteridse STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 204. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 235.
Primitive multi-rayed Phanerozonia apparently with the general
structure of Pabeasteridse, but with as many as 24 rays. Madreporite
known only in Helianthaster, where it is interbrachial and on the
158 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
edge of the disk, more actinal than abactinal. Inframarginals or
supramarginals bounding the rays.
Contains :
Lepidaster Forbes.
Eelianthaster Koemer.
Lepidaster ella, new genus.
Remarks. For a discussion as to why there are multi-rayed that
is, more than the normal five-rayed asterids, see the family Palse-
osolasteridse.
Genus LEPIDASTER Forbes.
Lepidaster FORBES, Mem. Geol. Surv. United Kingdom, British Org. Hem.,
dec. 3, 1850, p. 1, pi. 1, figs. 1-3. WRIGHT, Mon. British Foss. Echinod.,
Oolitic, vol. 2, pt. 1 (Palseontogr. Soc. for 1861), 1862, p. 35. WOODWARD,
Geol. Mag., dec. 2, vol. 1, 1874, p. 9. ZITTEL, Handb. Pal., vol. 1, 1879,
p. 454. STURTZ, Palseontographica, vol. 36, 1890, p. 222, pi. 28, figs.19-20;
Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 52, 72.
Generic description. Disk very large, over 50 mm. in diameter,
and apparently composed of heavy, closely set, polygonal plates.
Rays 13 in number, fairly stout, not slender, and tapering fairly
rapidly in the distal half. R = 52 mm. , r = 25 mm. A little more than
half the length of the rays extends beyond the disk.
Actinally, the rays are bounded on each side of the very narrow
ambulacral furrow by two columns of strong, tumid, closely ad-
joining, transversely oblong, adambulacral and inframarginal plates.
The latter are also the marginals for the abactinal side. The adam-
bulacrals are tuberculate and probably bore small spines; there
are 25 ossicles in each column. Ambulacrals unknown, but the
arrangement of the adambulacrals indicates that they were slightly
alternate.
Madreporite and interbrachial areas unknown.
Abactinally little is known. The rays are described as having
" numerous small polygonal nearly flat ossicula, closely set, and of
various sizes." To judge by similar rays in other genera, it would
appear that Lepidaster may have had radial and supramarginal
columns of ossicles.
Genoholotype and only species. L. grayi Forbes. The holotype
was found in the quarries of Wenlock limestone in the Castle Hill at
Dudley, England.
Observations. Forbes's illustration leads the writer to believe
that the abactinal plates have been deranged and that originally
the larger pieces were arranged in supramarginal columns, while the
smaller ones composed the radial row of ossicula. This type of
structure generally goes with the actinal arrangement as here seen in
Lepidaster , a duplicate of the same structure in Hudsonaster } Poise-
aster, and Mesopalseaster. Stiirtz compares the abactinal skeleton of
REVISION OF PALEOZOIC STELLEEOIDEA. 159
Lepidaster with the recent ScytJiaster, which is made up of plates of
various sizes without regular arrangement. It seems more natural
to compare Lepidaster with Siluric and Ordovicic genera than with
recent forms, and particularly when nearly identical structures are
found in the genera mentioned.
Forbes also describes Lepidaster as having the ambulacral grooves
" towards the extremity, in some instances partially filled up by small
polygonal intervening plates." His figures show a few of these
plates arranged in single columns. This anomalous structure is
undoubtedly best explained by accepting Stiirtz's conclusion, that
they are the separated ambulacral pieces, since otherwise none of
these plates appear to be present. As a fifth actinal, or a single
ambulacral column, the structure is out of all* harmony with star-
fishes.
Genus HELIANTHASTER Roemer.
Eelianthaster ROEMER, Palseontographica, vol. 9, 1863, p. 147, pi. 28. QUENSTEDT,
Petrefactenkunde Deutschlands, vol. 4, 1876, p. 82, pi. 93, fig. 32. ZITTEL,
Handb. Pal., vol. 1, 1879, p. 453. STURTZ, N. Jahrb. fur Min., etc.,
1886, vol. 2, p. 148; Palaeontographica, vol. 32, 1886, p. 81; vol. 36, 1890,
p. 218, pi. 27; Verb, naturh. Ver. preuss. Eheinl., etc., vol. 50, 1893, pp. 12,
30. GREGORY, Treat. Zool., vol. 3, Echinoderma, 1900, p. 258. CLARKE,
Bull. N. Y. State Mus., No. 121, 1908, p. 61, pi. 10. SCHONDORF, Jahrb. nas-
sauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 34.
Generic characters. Disk large; rays 14 to 16 in the genotype.
Interbrachial actinal areas well developed but not over-large for a
many-rayed asterid, either integumentary or granular, distally con-
cavely cut out and margined by several small inframarginals, about
seven on each side of a larger axillary piece.
Madreporite marginal, large, more actinal than abactinal, radi-
ately striate, interbrachial, and situated immediately outside of the
inframarginals. In the fossils it appears to be of the actinal surface.
Rays long and slender, protruding beyond the disk more than
twice the length inside the disk. Adambulacrals opposite one
another, with small lateral spines, prominent and large, wider than
long, bounding the rays, and terminating proximally in pairs of
modified pieces. These make the adambulacral parts of the oral
armature. Ambulacral grooves narrow, ambulacral ossicles not
seen but probably arranged opposite one another.
Abactinally the disk appears to have a scale-like covering. The
rays have median columns of radialia, outside of which are stout
columns of supramarginal ossicles.
GenoJiolotype (type by monotypy). H. rhenanus Roemer (citations
as above). Lower Devonic of Bundenbach, Germany. The above
definition is based on the illustrations of Stiirtz. There is a second
species in Germany, and H. filiciformis occurs in England.
50601 Bull. 8815 11
160 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
HELIANTHASTER FILICIFORMIS Woodward.
Helianthaster filiciformis WOODWARD, Geol. Mag., dec. 2, vol. 1, 1874, pp. 7-10,
fig. on p. 8.
A natural mold in sandstone measuring about 45 mm. in total
diameter. Has 11 rays. The disk "must have measured 10
mm. in breadth. The arms are 17 mm. in length, and near their
junction with the body are 5 mm. broad. " The specimen needs to
be restudied and modernized.
The holotype is from the Upper Devonic at Great Inglebourne,
Harberton, South Devon, England, and is now in the collection of
Mr. Champ ernowne.
LEPBDASTERELL.A, new genus.
Plate 25, fig. 2.
Lepidasterella SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914, p. 23.
Much like Lepidaster, but with 24 rays instead of 13 as in that
genus. Rays extending considerably beyond the disk. The abac-
tinal side only is known, but it shows that the rays have slender radial
columns, the ossicles of which are disposed alternatingly with the
larger and more prominent supramarginals. Outside of these in
some rays may be seen the edges of the inframarginalia, and these
have well developed but slender, rather long spines.
There appear to be no ambital areas, as the inframarginals of
adjoining rays meet and at their junction begins the disk. The
disk appears to have been made up of more or less closely adjoining
polygonal plates whose arrangement is not discernible in the specimen.
The madreporite and its position are unknown.
Actinal area unknown.
Genoholotype and only species. L. babcocki, new species.
Remarks. The essential difference between Lepidaster and Lepid-
asterella is that the former has 13 rays while the latter has 24.
LEPIDASTERELLA BABCOCKI, new species.
Plate 25, fig. 2.
Helianthaster, new species CLARKE, Fifty-ninth Rep. N. Y. State Mus., 1906, p. 37,
and plate.
Helianthaster gyalum CLARKE (part), Bull. N. Y. State Mus., No. 121, 1908, p. 61
(not pis. 12, I3=Pal3eosolaster (?) gyalum}.
Description. Rays, 24. Length of best preserved ray in the
holotype, 28 mm. Width of ray at base, 3 mm. Width of disk,
probably not less than 20 mm.
Rays slender and regularly tapering. Their structure is only
known abactinally, from a natural mold in a fine sandstone. Medi-
ally there is a narrow but high column of radial plates, each one of
KEVISION OF PALEOZOIC STELLEROIDEA. 161
which alternates with those of the supramarginal plates. The
latter are slightly tumid, subquadrate in outline, regular in size,
but decreasing gradually distally. Outside of these and margining
the rays may be seen a little of the inframarginalia, which bear pairs
of short, slender, and finely striated spines. There are from 31 to
33 closely adjoining plates in each one of the abactinal columns;
there appear to be no accessory plates of any kind in the rays.
Disk circular in outline, somewhat tumid and covered by small
many-sided plates that originally appear to have been closely adjoin-
ing.
Remarks. As this is the only American starfish with more than
five rays, it is readily distinguished from all others. The abactinal
arm structure is practically that of Hudsonaster, with the difference
that in L. labcocki the inframarginals do not project beyond the
supramarginals and are therefore not seen as a rule from the dorsal
side in the natural casts.
Locality and formation. The holotype, an excellent specimen, was
found in the lower part of the Upper Devonic, near Ithaca, New
York, by Mrs. G. W. Babcock, after whom the species is named;
it is in her possession. The writer's attention was called to this
interesting specimen by Prof. H. S. Williams of Cornell University.
Some years after this starfish was described in manuscript, Clarke
directed attention as above cited to two others found in the Cashaqua
shales of the Portage in Hunt's quarry, at Interlaken, New York.
Cat. No. 60627 (cast), U.S.N.M.
FAMILY, unknown.
Genus ASTROPECTEN Link.
The genus Astropecten is based upon living species, and it is not
probable that the Devonic forms are identical.
ASTROPECTEN (?) SCHLUTERI Sturtz.
Astropecten schluteri STURTZ, Palaeontographica, vol. 32, 1886, p. 87, pi. 10, figs. 1,
16, 2, 2a; Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 51, 71.
In the roofing slates of the Lower Devonic at Bundenbach, Ger-
many. A single very much distorted and compressed example is
the basis of this species.
Genus ATAXASTER Jaekel.
Ataxa.ster JAEKEL, Zeits. geol. Gesell., vol. 55, 1903, Protokol, p. 14 (106).
Genoholotype and only species. A. pygmseus Jaekel. Ordovicic
(DJ, St. Benigna, Bohemia.
Description. "A small asterid whose ambulacrals in the distal
parts are alternate, but in the proximal are opposite."
Remarks. The above is all that is known about Ataxaster, and
for the present the genus has no value.
162 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
CRYPTOZONIA Sladen.
Specialized Asteroidea without true marginal plates. In other
words, there are no large and conspicuous marginal plates bordering
the animals, but in primitive forms large adambulacrals do margin
the species. In other primitive forms more or less of the infra-
marginals may be present, but if so they remain adjoining the adam-
bulacrals.
Ambulacrals alternating or directly opposite one another.
Oral armature generally with the adambulacral elements the most
conspicuous; more rarely the ambulacral elements are most promi-
nent.
Contains the following families:
Stenasteridae. Schuchertiidse.
Monasteridae. Palasteriscidae.
Urasterellidae. Schcenasteridae.
Calliasterellidae- Palaeosolasteridae.
Compsasteridae.
Remarks. This widely accepted order, which appears as if founded
in nature on a single grand phylum, probably has no such value.
In other words, asterids without either one or both of the marginal
columns bounding the animal developed several times. Therefore,
the absence of these ossicles is no indication of direct phyletic
relationship, but such are simply cases of parallel development. As
long as asterids retain the primitive characters of few columns of
rather strong ossicles and the adambulacral type of oral armature,
they are clearly Phanerozonia, but when the ponderous skeleton
breaks up into a more flexible one of small pieces, then the marginal
and abactinal columns vanish as such in the mass of small plates.
On the other hand, ponderosity may even be retained with the loss
of the inframarginals alone. In all of these Paleozoic forms, the
adambulacral type of oral armature is the rule.
Among the Paleozoic "Cryptozonia" it seems certain that at least
the families Stenasteridae, Urasterellidae, Schuchertiidae, and Palaeo-
solasteridae had independent origins in Phanerozonia stocks. The
Urasterellidae and Calliasterellidae apparently had a single origin in
the Phanerozonia subfamily Mesopalaeasterinae by the adoption of
marked flexibility. On the other hand, the families Stenasteridae
and Monasteridae retained rigidity and primitive characters, and
more probably arose in the Hudsonasteridae through the loss of the
iriframarginals alone. The Schuchertiidae also had their origin
directly in some Phanerozonia stock and one naturally looks for it
in forms developing large interbrachial areas as in the Palasterinidae.
In this family, however, the accessory plates are introduced between
the adambulacrals and inframarginals, while in the former family they
KEVISION" OF PALEOZOIC STELLEROIDEA. 163
are introduced outside of the inframarginals and the entire abactinal
skeleton is broken up into an abundance of tiny pieces. Therefore, one
must go back for their origin at least as far as the Palaasteridse. In
this case it was not primarily flexibility that led to this supposed
ordinal change, but it was due to increase of body cavity and enforced
enlarging of the skeleton in the interbrachial and ambital areas,
making the more or less pentagonal disk. It is probable that the
Palasteriscidas were their descendants, while the Schoenasteridse may
have been the final Paleozoic representatives of the same phyletic line.
The PalasosolasteridaB seemingly arose in the Phanerozonia family
LepidasteridaB by the adoption of a flexible skeleton.
It would, therefore, seem that the Paleozoic families of supposed
Cryptozonia construction are to be grouped as follows:
Superfamily Stenasteracea, new.
Includes families Stenasteridse and Monasteridse.
Superfamily Urasterellacea, new.
Includes families Urasterellidse, Calliasterellidse, and Compsasteridse.
Superfamily Schuchertiacea, new.
Includes families Schuchertiidse, Palasteriscidee, and Schcenasteridae.
Superfamily Palseosolasteracea, new.
Includes family Palaeosolasteridse.
STENASTERIDSE, new family.
Rigid, small, primitive, five-rayed Cryptozonia, with rapidly taper-
ing or petaloid rays. Ambulacrals few in number and opposite in
arrangement. Interbrachial areas consisting of the single pairs of
large basal adambulacral oral armature plates.
Abactinal skeleton unknown, but in some forms seemingly integu-
mentary. No spines as yet known. It is probable that this family
will be referred to the Auluroidea.
Contains:
Stenaster Billings.
Tetraster Nicholson and Etheridge.
Genus STENASTER Billings (emend.).
Plate 32, fig. 1.
Stenaster BILLINGS, Geol. Surv. Canada, Can. Org. Bern., dec. 3, 1858, p. 77.
STURTZ, Palseontographica, vol. 36, 1890, p. 220. GREGORY, Geol. Mag.,
dec. 4, vol. 6, 1899, p. 352. SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1
(Palseontogr. Soc. for 1913), 1914, p. 22.
Urasterella STURTZ (not McCoy), Verb, naturh. Ver. preuss. Rheinl., etc., vol.
50, 1893, pp. 40, 41, 56.
Generic characters. Small rigid asterids, the largest example having
11 = 20 mm., / = 4.5 mm. Disk small, without interbrachial arcs.
Rays five, short, stout, lanceolate or petaloid.
Abactinal area devoid of a preservable skeleton. For this reason
when this side is at hand it is nearly always mistaken for the actinal,
so nearly alike are the two.
164 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Madreporite unknown.
Adambulacral plates with granular surfaces, few in number, never
more than 17 hi a column, subquadr angular and thick, greatest width
about the middle of the rays, rapidly diminishing in size distally and
making the margin of the rays and disk. The great proximal adam-
bulacrals are the orals, occupying the axils in pairs, acutely triangular,
more or less pointed, and terminating orally.
Ambulacral plates large, directly opposite one another and the
adambulacrals as well, h- -shaped, with the widest portion in the per-
radial center of the rays; there are as many as 16 or 17 ambulacrals
in a column. Laterally each plate is more or less excavated, both
distally and proximally, thus leaving very large, subcircular podial
openings.
Spines of any kind are so far unknown.
Genoholotype (the first species and the one selected by subsequent
workers). Stenaster salteri Billings.
Remarks. This genus has not had a good standing, due largely
to the fact that Palseaster Hall was poorly known when Billings
described Stenaster and further because he included two distinct
generic types in his genus. A careful reading of Stenaster, however,
reveals that Billings actually based his genus on S. salteri, the first
form following the generic description, and this selection has been
followed since. Later on, McCoy applied the name Urasterella in
a loose manner to the second species (Palseaster pulcJiellus) , and
although it is a form of this genus it is not the genotype. Hall (1 868)
directed attention to this work of McCoy, but erroneously assumed
that all of Stenaster was synonymous with Urasterella. It is true,
as stated by Hall, that Billings's second species is congeneric with
Urasterella, but the first species and genotype is certainly quite dis-
tinct from McCoy's genus. Sttirtz (1890) also points out the same
thing and gives a short and emended diagnosis of " Urasterella
(Stenaster)." In 1893 the same writer applied McCoy's name
Urasterella to Stenaster salteri and made S. pulchellus the genotype
of Urasterella, a proceeding that violates the rules of nomenclature.
Stenaster as here restricted differs from Urasterella in being more
decidedly rigid and not so flexible as the latter, while the rays are
far shorter and petaloid in form. For these reasons, Stenaster has
comparatively few adambulacrals and ambulacrals, while there are
many more of these ossicles in Urasterella. Finally, the former
reveals no abactinal plates or spines of any kind, while the latter
has a skeleton of small plates that terminate in long inarticulate
spines or rods.
Actinally Stenaster is closely related to Tetraster, and the only
marked difference is that the former has petaloid rays and more
pronounced adambulacral oral jaws.
REVISION OF PALEOZOIC STELLEROIDEA. 165
There would be no need to point out here the difference between
such distinct genera as Stenaster and Palseaster were it not that
Hall regarded the former as a synonym of the latter. In Stenaster
the rays are petaloid and not slender and regularly tapering, as in
Palseaster. The most marked distinction, however, is the fact that
these two genera belong to different orders, for Palseaster has columns
of iirframarginals that are completely absent in Stenaster. Further,
in the last-named genus, the interbrachial areas are made up each
of two pairs of oral armature pieces, while in Palxaster there are,
distaUy to these, additional large single axillary interbrachials.
Stenaster and the related genus Tetraster are very primitive in
construction. Actinally they are seen to have the structure of
Hudsonaster minus the inframarginal columns and the single axillary
interbrachials. It is this difference that makes the former genera
members of the order Cryptozonia, while Hudsonaster is the most
primitive of Phanerozonia. Until the abactinal area is known ha
Tetraster and Stenaster, positive phyletic relationship can not be
stated, but in a number of specimens of the latter genus the evidence
at hand indicates decided specialization through the complete removal
of the heavily plated skeleton of Hudsonaster.
Distribution. Ordovicic of America. The following are the
known species:
8. salteri Billings. Black River and Trenton.
8. (?) oUusus (Forbes). Bala of Ireland and Wales.
S. (?) coronella (Salter) . May Hill oandstone (Siluric) of England.
S. (?) confluens Trautschold. Upper Carboniferous of Russia.
STENASTER SALTERI Billings.
Plate 32, fig 1.
Stenaster salteri BILLINGS, Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858,
p. 78, pi. 10, figs, la, 16. WRIGHT, Mon. British Foss. Echinod., Oolitic,
vol. 2, pt. 1 (Palseontogr. Soc. for 1861), 1862, p. 28, fig. 15 (2) on p. 24.
Original description. "This species has rather short, broad rays,
which are narrower where they are attached to the very contracted
body than they are at about the center of their length. In conse-
quence of this form, the sides of the rays are not parallel, but a little
curved outward. As however only two specimens have been col-
lected, and both appear to be a little flattened by vertical pressure,
it may be that this leaflike shape of the rays is accidental [an
abundance of specimens show this form to be the natural one].
* * * The adambulacral plates are oblong [granular and from 7
to 17 in each column in the smallest and largest examples], and
the sutures between them are nearly at right angles to the ambu-
lacral grooves; those next the body are a little sloping outward.
166 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Their length is about twice their breadth, and they are so disposed
that the greater dimension is transverse or at right angles to the
groove; the extremities which lie next to the grooves are angular,
and some of them appear to have the contiguous pores partly exca-
vated in them. The oral plates [adambulacrals] are acutely triangu-
lar, the sharpest angle being toward the mouth [and form the inter-
brachial areas]. The plates are smooth [the adambulacrals are all
distinctly granular]. The ambulacral pores are very large, and the
ossicles are much contracted in the middle and greatly expanded
along the median line of the bottom of the groove." These plates
are directly opposite one another and the adambulacrals, and in
number equal those of the latter.
The type-specimen figured by Billings measures: R = 14 mm.,
r=3.5 mm., R = 4n Width of a ray at base about 4 mm., at about
mid-length 5 mm. The largest specimen: K- = 20 mm., r 4.5 mm.
In the University of Toronto there is a specimen that in every
way, except one, has the characters of Stenaster salteri. It was
found associated with many other individuals at Kirkfield. It
differs from its associates in having what appears to be a distinct
disk, rather large, with concave sides, filling in the spaces between the
rays. One looks in vain, however, for plates or spines, as the inter-
brachial areas are nothing more than an amorphous mass of cal-
cium carbonate. These areas are very distinct and stand out prom-
inently, but because they do not reveal any plated or spinif erous struc-
ture the writer regards them as false characters, produced during the
permineralization of the specimen. This conclusion is further sup-
ported by the fact that otherwise the characters are those of S.
salteri.
Formation and locality. Three specimens (the type, No. 1398, and
two fragments) are from the Trenton limestone at Belleville, Ontario,
and are now in the Museum of the Canadian Survey. Another
specimen in that museum is on the same slab with Petraster rigidus
and has been labeled Palseaster matutina. In Mr. Ulrich's collection
there are four isolated rays of this species obtained by him from the
Black River limestone at Curdsville, Kentucky. A further specimen
preserving two rays, from Government House Bay, Ottawa, was found
by Mr. Walter R. Billings. In the Museum of Comparative Zoology
there is still another individual which appears to be of this species;
it was found by Mr. J. B. Perry at Pant on, Vermont. Finally, an
abundance of material was obtained in the Black River limestones
at Kirkfield, Ontario; there are more than twenty specimens from
this locality in the Walker collection of the University of Toronto,
gathered by Mr. Townshend.
Cab. No. 60628, U.S.N.M.
REVISION OF PALEOZOIC STELLEROIDEA. 167
STENASTER (?) OBTUSUS (Forbes).
Asterias primseva SALTER and SOWERBY, Quart. Jour. Geol. Soc. London, vol.
1, 1845, p. 8, 20 (table) (rumen' nudum).
Uraster obtusus FORBES, Mem. Geol. Surv. Gt. Britain, vol. 2, pt. 2, 1848, p. 463;
Mem. Geol. Surv. United Kingdom, dec. 1, 1849, p. 2, pi. 1, fig. 3. MURCHI-
SON, Siluria, 1854, p. 182, fig. 17.
Palxaster obtusus SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 326.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Paleeontogr.
Soc. for 1861), 1862, p. 24. SALTER, Mem. Geol. Surv. Gt. Britain, vol. 3,
1866, p. 289, pi. 23, fig. 1.
Stenaster (?) obtusus STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 153; Verh.
naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 41, 56.
Stenaster obtusus SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for
1913), 1914, pp. 22, 23, 31, text fig. 21, pi. 1, figs. 6, 7.
The little that is known of this species appears to be in harmony
with the characters of Stenaster as here defined. It occurs in the
Caradoc beds (Ordovicic) at Drumcannon, Waterford, Ireland, and
in the Bala rocks west of Bala Lake (Moel-y-Garnedd) , North Wales.
STENASTER (?) CORONELLA (Salter).
Palxaster coronella SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 326.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palseontogr.
Soc. for 1861), 1862, p. 25.
Stenaster (?) coronella STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 153.
The description of this species is very short and almost nothing is
known of its characters. It occurs in the May Hill sandstone (Siluric)
at Gunwick Mill, Malvern, England.
STENASTER (?) CONFLUENS Trautschold.
Stenaster conftuens TRAUTSCHOLD, Nouv. Mem. Soc. Imp. Nat. Moscou, vol. 14,
1879, p. 11, pi. 2, fig. 5. SCHONDORF, Palseontographica, vol. 56, 1909, p. 337.
Based on a fragment of the distal part of a ray. Appears to be
a crj^ptozonian. Generic position unknown. From the Upper Car-
boniferous (Moscovian) of Moscow, Russia.
Genus TETRASTER Nicholson and Etheridge (emend.).
Plate 33, fig. 4.
Tetraster NICHOLSON and ETHERIDGE, Mon. Silurian Foss. Girvan Dist., Ayrshire,
fasc. 3, 1880, p. 324, pi. 21, figs. 3-8 (not figs. 1-2= Eudsonaster batheri, and
9-10== Urasterella girvanensis) .
The name has reference to the four columns of actinal plates of
the rays.
Remarks. The authors of Tetraster distinguished their genus from
Palseaster on the ground that it had on the actinal side but four
columns of plates, while that genus as typified by P. matutinus } now
Hudsonaster matutinus, has six. In this definition they are correct,
but they complicated their good intentions not only by referring to
168 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Tetraster another Cryptozonia form (Urasterella girvanensis, new
species), but also by confusing with their genotype specimens that
certainly have the character of Palseaster so far as six columns of
actinal ossicles are concerned. This latter admixture the writer has
taken out of the genotype and has described it in this work as Hudson-
aster ~batheri. The adjusting of this matter has been made possible
through the kindness of Dr. Bather, who made for this work wax
squeezes of the various specimens. After seeing these and the other
material originally referred to Tetraster by Nicholson and Etheridge,
their various drawings became clear. Their figures are fairly good,
but in places do not bring out 'all the characters, but it was the
mixture of three generic forms under Tetraster that made it for a time
impossible to determine on what the genus should stand.
Emended description. Small, five-rayed asterids, E, = about 5 mm.,
r= about 2 mm. Abactinal side unknown. Actinally convex and
in its general expression reminding much of living species with large
marginals, but in Tetraster the comparatively massive margining
plates are adambulacralia.
Adambulacrals compose the entire outer actinal areas of Tetraster
and margin the entire annual. Distally the pieces are small and the
12 ossicles of each column increase rapidly in size with the largest
basal pieces of adjoining columns making the prominent interbrachial
areas. Ambulacrally all are somewhat excavated, stand high above
the furrow and each ossicle terminates in a ridge that joins those of
the ambulacrals. All of these plates are abundantly granular.
Ambulacral furrows not wide but rapidly becoming very deep
proximally, with excavated adambulacral sides. The columns meet
in front of the two largest interbrachial adambulacrals. Podial
openings are large and issue deeply from beneath the adambulacrals
and between the lateral sutures of the ambulacralia. The pieces of
the ambulacral columns are as many as there are adambulacrals and
are placed directly opposite one another. Medially the ambulacralia
are elevated, roof-like, and without a central gutter.
While the ab actinal area is unknown, it appears that the adambu-
lacrals also margin this side of the animal. The nature of the skele-
ton inside of these columns may be of small pieces as hi Urasterella,
or, what seems more probable, consist of large radial and supramar-
ginal ossicles, without accessory pieces.
Spines of any kind are unknown.
Genoholotype. Tetraster wyville-thomsoni Nicholson and Etheridge
as above emended. T. (?) imbricatus (Salter) is also placed under
this genus for the present.
Tetraster as now defined is so distinct in its great adambulacral
development that it will not be confused with any other Paleozoic
genus except Stenaster. Both genera, so far as the actinal area is
REVISION OF PALEOZOIC STELLEEOIDEA. 169
concerned, are closely related. The differences, however, are worthy
of generic rank. In Stenaster the rays are petaloid, with the adambu-
lacrals largest at the mid-length of the rays, while in Tetraster the
rays are triangular, with the greatest ossicles in the axils.
At first the genotype looks as if one had a direct representative of
living Phanerozonia, but a little study quickly shows that the large
actinal margining plates are not marginals or inframarginals, but
adambulacrals, and therefore indicate a cryptozonian form. For a
while it seemed that these large plates must be inframarginals, but
then adambulacrals should be present, and they were looked for
along the ver'tical ambulacral walls, but even here there are no
diminutive pieces of these columns. Among Paleozoic asterids the
writer has not seen a form without adambulacrals, and when there are
but four actinal columns it is always the inframarginals that are
diminutive or absent.
This species may be identical with T. (?) inibricatus (Salter),
which see for further remarks.
Distribution. Restricted to the Upper Ordovicic at Thraive, Gir-
van, Scotland. The specimens occur as very sharp natural molds in
a micaceous sandstone, from which good wax squeezes can be made,
revealing all the skeletal characters. The originals are in the collec-
tion of Mrs. Robert Gray, Edinburgh, Scotland.
TETRASTER (?) IMBRICATUS (Salter).
Palseaster imbricatus SALTER, Mem. Geol Surv. Gt. Britain, vol. 3, 1866, p.
289, pi. 23, fig. 8.
Original description. "P. rigidus, uncialis. Radii breves conici,
nee ad basin valde expansi, subcarinati, tuberculis seu jugis obliquis
remotiusculis asperi.
"Ray conical, its length about five lines, its breadth at base less
than two lines. Only two rays are preserved; these are convex,
slowly tapering, somewhat carinate above (we do not see the lower
side), and ornamented by tubercles gathered into curved oblique
rows, which run forward from the avenues toward the carina above,
and are separated from each other by a space little more than their
own breadth.
" Locality: Caradoc rocks, Llanfyllin, Montgomeryshire [Wales].
The specimen is in the collection of Mr. Prosser."
Remarks. As the writer has not seen the holotype, his conclusions
are based on the lithographic figure, which adds but little knowledge
to that in the description. As there are but two columns of dorsal
ray plates, it hardly seems that the species is a phanerozonian. In
general it looks most like Tetraster, but as the dorsal side of this
form is unknown, no direct comparisons can be made between them.
On the other hand, the imbrication of the plates, their large size,
170 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
and completely opposite arrangement suggest that T. (?) imbricatus
may be an auluroid. Until the original specimen is restudied no
definite assignment can be given it.
MONASTEKID^E, new family.
Progressive but primitive five-rayed Cryptozonia, with massive
plates, especially the adambulacrals, which are common to the actinal
and abactinal areas. Ambulacrals opposite. Interbrachial areas con-
sist of a number of pah's of adambulacrals crowded orally Abacti-
nally the radial and supramarginal columns consist of large, closely
adjoining plates, and are similar to those of Hudsonaster.
Contains :
Monaster Etheridge.
Genus MONASTER Etheridge.
Palseaster (Monaster} ETHERIDGE, jr. (part), Mem. Geol. Surv. New South Wales,
Pal., No. 5, pt. 2, 1892, pp. 70, 71.
Etheridgaster GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 353 (genoholotype,
Palxaster clarlcd).
Emended description. Animal large, very stout, with five thick
petaloid rays. Disk very large, with marked ray angles.
Abactinal area of rays with prominent radial, and supramarginal
columns of large, contiguous, strongly convex, hexagonal plates,
densely covered with small granules radially arranged. The ossicles
of the radial columns alternate with those on each side and are largest
in the mid-length of the rays, decreasing in size both distally and
proximally, but most rapidly distally. This growth gives the ray
columns a decided petaloid form. Outside of the supramarginals the
animal is bounded by single columns of prominent, short, but very
wide plates, which are the abactinal aspect of the actinal adam-
bulacrals. These plates bear numerous spiniferous tubercles. The
ossicles of the axillary disk region are not determinable. In the center
of the disk there appear to be five distinct radials and a central disk
plate, and around these probably were numerous small accessory
pieces.
Madreporite abactinal, large, oval, radially striate, and situated
near the axillary margin.
Actinally the rays consist almost entirely of the adambulacrals,
which are numerous, abundantly tuberculate, short, and very wide.
These columns increase rapidly in width, join and form angular
axillse, and leave between themselves deep petaloid ambulacral
grooves. The interbrachial areas consist of the oral extensions of
adjoining columns of adambulacrals, of which there are a great num-
ber of pairs, terminating in the mouth in five small pairs, the oral
armature.
KEVISION OF PALEOZOIC STELLEEOIDEA. 171
Ambulacrals in two columns, opposite, short but wide, and about
as numerous as the adambulacrals. Podia! openings unknown.
Genoholotype. Palseaster darlcei De Koninck. This is also the only
species of Monaster and occurs in the "Upper and Lower Marine
series" of the " Permo-Carbonif erous " of New South Wales.
Remarks. Mr. Etheridge proposed Monaster as a subgenus of
Palseaster, from which it differs so radically as to give it easily the
rank of a genus. He defines the group as follows : In Palseaster " the
adambulacral plates, bordering the ambulacral avenues, are small
and quadrangular, followed by large transverse marginal plates. In
our Permo-Carboniferous species, on the contrary the adambulacral
plates are transversely elongated, and occupy nearly the whole of
the actinal surface on each side the avenues. The marginal plates
[having reference to the plates along the margins which consist of
adambulacrals, and also form part of the abactinal skeleton], in con-
tradistinction to those of Hall's Silurian species, are here smaller and
subdorsal in position. The question now presents itself, of what value
in a classificatory sense is this character ? Hall lays particular stress
on the position of these plates on the actinal side of Palseaster. He
says it 'has two ranges of plates on each side of the ambulacral
groove; marginal [inframarginal] and adambulacral plates on the
lower side, besides ambulacral or poral plates. The upper or dorsal
side has three or more ranges of plates/ In the case of our specimens,
only one set of plates, excepting those of the ambulacral grooves,
are, as before stated, absolutely actinal; the marginals [same usage
as before] are strictly so, or, at the least sub-dorsal. Under these cir-
cumstances, I purpose distinguishing our Australian species under
the subgeneric name of Monaster (from the one or single row of
adambulacral plates on each side of an ambulacral avenue) ."
The original definition is here given at length to show that it can
only apply to the structure in Palseaster darkei De Koninck, which
is also the first species following the generic discussion. The author
in the same work describes a new species which he also refers to
Monaster, Palseaster (Monaster) giganteus, but its structure is radi-
cally different and does not conform with the subgeneric definition
by Etheridge. Elsewhere in this work it is taken as the genotype of
the new genus Australaster . Gregory in the serial above cited takes
as the genotype of Monaster the last named species, the third of
Etheridge, and transposes the former as the genotype to his new
genus Eiheridgaster. This the writer holds can not be done even
though Etheridge selected no genotype for Monaster. That author's
genotype, as the name implies, refers to a species with but a single
column of plates on each side of the ambulacrum. The species selected
by a subsequent author should have the structure implied in the
172 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
original generic definition and when none actually applies, then the
first species following the generic diagnosis. That Gregory fully
appreciated the structural difference between Palseaster clarkei and
P. (M.) giganteus is shown in his reference of the former to the order
Phanerozonia and the latter to the Cryptozonia. Our disagreement
is not as to structure but in the interpretation of the rules of nomen-
clature. Under these circumstances P. clarkei is retained as the geno-
type of Monaster, since it appears that P. giganteus has both infra-
marginals and adambulacrals on the actinal side, a structure quite
different from that described for and present in Monaster Etheridge.
After the above was written in 1900, the writer explained by letter
to Professor Etheridge what Gregory had done. Etheridge replied
under date of June 6, 1900: "The laws of zoological nomenclature
should have guided Doctor Gregory to have selected the first species
described, as the subgeneric type, in the absence of any statement to
the contrary on my part. As you say, Eiheridgaster is certainly a
synonym of Monaster."
Monaster is most like Hudsonaster. It retains the primitive pon-
derous skeleton of the latter, but curiously has lost the inframarginal
columns of ossicles, although the supramarginals are present.
MONASTER CLARKEI (De Koninck).
Palseaster clarkei DE KONINCK, Mem. Soc. roy. Sciences, Liege, ser. 2, vol. 2, 1877,
p. 166, pi. 7, figs. 6, 6a; (David), Mem. Geol. Surv. New South Wales, Pal.
No. 6, 1898, p. 127, pi. 7, figs. 6a, 66.
Palseaster (Monaster) clarkei ETHERIDGE, jr., Mem. Geol. Surv. New South Wales,
Pal. No. 5, pt. 2, 1892, p. 71, pi. 14, figs. 1, 2; pi. 15, fig. 4.
Etheridgaster clarkei GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 353.
Of this species three natural molds have been found in the "Upper
Marine " and "Lower Marine' 7 series (Carboniferous) of Northum-
berland County, New South Wales. The holotype in the Clarke
collection was destroyed by fire.
The species attained a large size, the rays from the center of the
disk to their extremities are not less than 5 cm., and in another
specimen are 6.5 cm.
URASTERELLID^E, new family.
Txniasteridse GREGORY (part), Geol. Mag., dec. 4, vol. 6, 1899, p. 351 (includes
Tseniaster, fStenaster, Urasterella, Protasteracanthion, Salteraster).
Roemerasterinx GREGORY, Treat. Zool., vol. 3, Echinoderma, 1900, p. 255.
Specialized Cryptozonia, with alternate ambulacra, and with adam-
bulacral type of oral armature. Rays five, rather flexible, long and
gently tapering, proximally united without forming interbrachial
areas. No inframarginals discernible as such at maturity, the actinal
margin being occupied by well-developed adambulacrals. Abactinal
area composed of numerous small plates arranged in columns and
KEVISION OF PALEOZOIC STELLEKOIDEA. 173
quincunx. The radial and supramarginal columns may be dis-
cernible in somewhat larger plates. Adambulacrals many, like coins
set on edge. *
Contains:
Urasterella McCoy
Genus URASTERELLA McCoy (emend.).
Plates 27 to 30.
Urasterella McCoY, British Pal. FOBS., 1851, p. 59 (not defined). HALL, Twentieth
Kept. N. Y. State Cab. Nat. Hist., 1868, p. 289; rev. ed., 1868=1870, p. 332.
McCoY, Geol. Surv. Victoria, dec. 1, 1874, p. 42. ZITTEL, Handb. Pal.,
vol. 1, 1879, p. 453. STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 152;
Palseontographica, vol. 36, 1890, p. 219. GREGORY, Geol. Mag., dec. 4, vol. 6,
1899, p. 352.
Stenaster BILLINGS (part), Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858,
p. 77. STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893,
pp. 40, 56.
Eoemeraster STURTZ, Palaeontographica, vol. 32, 1886, p. 85 (genoholotype, Asterias
asperula Roemer); vol. 36, 1890, p. 220; Verh. naturh. Ver. preuss. Rheinl.,
etc., vol. 50, 1893, pp. 52, 73.
Protasteracanthion STURTZ, Palseontographica, vol. 32, 1886, p. 90 (genoholotype,
P. primus Stiirtz= Asterias asperula Roemer); Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, pp. 54, 75.
Palasteracanthion STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 153 (genoholo-
type, "A, primus "= Asterias asperula Roemer). STURTZ, Verh. naturh. Ver.
preuss. Rheinl., etc., vol. 50, 1893, p. 75.
Salteraster STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp.
43, 60 (genoholotype, Palseaster asperrima oalter).
fEoactis SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913),
1914, p. 30 (genoholotype, E. simplex Spencer=? Urasterdla girvanensis).
Generic characters. Disk very small, without interbrachial arcs.
Rays five, very long and slender at maturity, but shorter and rapidly
tapering when immature.
Abactinal side with very numerous small stellate plates, all of
which, excepting one or three medial columns, are drawn out into
more or less long, blunt, stout, erect, nonarticulating rods. The disk
is occupied by a central plate and a first ring with five plates, which
in some species are seen to be the basal radial plates, followed by a
second ring with 1 5 plates. Five of the latter are the second proximal
radial plates, and 10, basal plates of the supramarginal columns.
Between the central plate and the first ring there may be inserted a
few accessory, much smaller pieces. These disk ossicles may remain
more prominent throughout life or may have the size of the other
abactinal plates, in which case their arrangement is not plainly made
out. In the center of the ray the adial column may be the more
conspicuous throughout life or it may be more or less completely
suppressed when the supramarginals adjoin and these may then
174 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
be the more prominent columns. Or the entire abactinal area of the
rays may be composed of ossicles arranged in columns and in
quincunx when the plates are of nearly equal size. The number of
columns varies with age and in different species, the greatest number
observed being 13 at the base of a ray.
Ambital plates often not readily distinguishable at maturity from
the radial and supramarginal columns. The growing distal tips of
the rays of Urasterella have no ambital plates, but shortly after their
introduction they begin to develop the rodlike extension so charac-
teristic of this genus. Of these columns there may be as many as
four at the base of a ray on each side of the supramarginals.
Inframarginals, as such, usually not distinguishable at maturity
from the ambital plates. Their position is rather abactinal than
actinal. At the distal end of rays, however, these plates lose their
ambital character, passing over somewhat to the actinal side, and
closely adjoin the adambulacrals. In the young of U. ulrichi the
inframarginals are well developed in the axillary regions and here
there is also a single interbrachial marginal plate. More dis tally,
however, the inframarginals have the character of the ambital plates.
In none of the mature specimens have been seen well-developed or
larger axillary inframarginals or interbrachial marginal plates. In
U. asperula the axils have actinally single plates which may be
remnant inframarginals.
Madreporite abactinal, of medium size, slightly convex, depressed
and granules triate.
Adambulacral plates very numerous, coin-shaped, and arranged on
edge with the actinal surface pustulose. Each plate on its actinal
surface bears two or three short, thick spines, and on its ambulacral
side there is another similar spine. Along the outer edge of these
plates toward the abactinal side there is another row of spines, in
pairs, which are long and slender, flat, and longitudinally grooved
on two sides. The adambulacral columns terminate in small tri-
angular plates of the oral armature. In the young of U. ulrichi five
very stout, short, pointed spines (tori) are inserted inside of the plates
of the oral armature.
Ambulacral columns with the ossicles generally slightly alternating,
but they may also be opposed, one plate to each adambulacral piece,
more or less wedge- or club-shaped, and those of each column par-
tially overlapping. Medially the columns loosely adjoin. Each
plate is excavated laterally, along the proximal edge, leaving a more
or less long, slender podial opening between adjoining plates.
Genoholotype. Uraster ruihveni Forbes. The above diagnosis,
however, is based on Stenaster grandis Meek, a form thought to be
in generic harmony with the genotype.
KEVISION OF PALEOZOIC STELLEKOIDEA. 175
Distribution. The species occur in North America and Europe and
range from the Ordovicic to the Upper Carboniferous. The following
are the known species:
U. pulcJiella (Billings). Trenton.
U. grandis (Meek). Richmond.
U. fiuxleyi (Billings). ?Chazy.
U. ulrichi, new species. Lowville.
U. girvanensis, new species. Upper Ordovicic of Scotland.
U. (?) asperrima (Salter). Bala of Wales.
U. (?) constellata (Thorent). Siluric of France.
U. ruthveni (Forbes). Ludlow of England.
U. Jiirudo (Forbes). Ludlow of England.
U. selwyni McCoy. Siluric of Australia.
U. asperula (Roemer). Lower Devonic of Germany.
U. } new species. Ithaca beds. Upper Devonic.
U. montana (Stschurowsky). Upper Carboniferous of Moscow,
Russia.
Remarks.- McCoy in suggesting the name Urasterella did it in the
following words:
11 Before I was aware Prof. E. Forbes had described them it seemed
to me that the U. [raster] RutJiveni and U. Hirudo, as well the similar
American species, might be easily separated from the great star-
fishes forming the recent genus Uraster, by their small size and much
more simple skeletons, and I had named the genus Urasterella in my
manuscript."
This is all that McCoy did to establish the genus Urasterella,
abandoning his manuscript definition on learning that Forbes had
described the species as those of Uraster. Salter 1 in 1857 paid no
attention to this name, although he was the first to recognize and
define Palasterina, which McCoy proposed at the same time and in
the same manner. It seems that Hall (1868, p. 332) was the first
to revive the name Urasterella and regarded Billings's Stenaster as a
synonym by taking his second species, 8. pulchella, as the genotype.
Hall's remarks are as follows: "Mr. Salter, apparently overlooking
my description of Pal&aster niagarensis, has placed both the above-
cited European species [ Uraster rutTiveni and U. Jiirudo] under Palae-
aster, as defined by himself; but as it has been shown that this genus
has two ranges of plates on each side of the ambulacral groove, these
forms can not with propriety be arranged with typical Palseasters.
" There was the same reason for adopting the generic term Uras-
terella as for adopting Palasterina, both of which were proposed by
McCoy at the same time, and the typical forms of each were specified.
Mr. Salter has for some reason recognized the genus Pabea,ster pro-
iAnn. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 326.
50601 Bull. 88 15 12
176 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
posed by me a little earlier, but probably published after that of McCoy;
and has given the two species of Urasterella to that genus. Now
that there seems no reason for continuing them under that generic
term, they should be restored to their proper position as indicated
above. The two species of Forbes must either be placed under
Urasterella as proposed by McCoy, or fall under the later genus
Stenaster of Billings. Were the latter genus of my own proposing
I should nevertheless return to Urasterella, and I have no doubt
that Mr. Billings will accord with this view."
Palseaster pulcJiellus is a true Urasterella, but this does not make
the genus Stenaster a synonym of Urasterella, since S. salteri Billings
has quite another structure and is here accepted as the genotype of
Stenaster.
Meek 1 also recognizes McCoy's name, but because of certain pecul-
iarities in S. salteri, which he recognizes as the genotype, and also
since he had not seen the dorsal side of McCoy's S. grandis, he prefers
to retain Stenaster for the latter.
McCoy returned to his genus in 1874, after going to Australia, and
described a new form, U. selwyni. It is important to quote here
his remarks :
" Genus Urasterella (McCoy )= Stenaster (Billings).
"Gen. Char.: Small starfishes, with five moderate rays, narrowed
at the base, and without disk. Ambulacral grooves narrow, bor-
dered on the under side, with only one row of large (adambulacral)
plates; no marginal plates. Upper surface with numerous rows of
small tubercular plates. Confined to Silurian rocks.
"The late Mr. Salter and Mr. Billings refer the starfishes of this
type to the subsequently published genus Pateaster of Hall; but,
as Prof. Hall objects that his genus Palseaster has ambulacral, adam-
bulacral, and marginal plates, and the types of my genus U. ruthveni
and U. Jiirudo of the English Ludlow rock, like our Australian species
and the American Pals&aster or Stenaster pulchella, have only one row
of plates on each side of the ambulacral groove, I return to the use
of my old generic name."
Zittel 2 says that Stenaster is a synonym for Urasterella and defines
the latter genus. Stiirtz 3 points out the errors in the diagnosis of
Billings and that of Zittel, redefines the genus, and correctly states
that Stenaster Billings contains species of two genera, that is, Sten-
aster salteri, the genotype of Stenaster, and S. pulchella, which he refers
to Urasterella. In this way the genus Urasterella has come to be
recognized.
The figure and description of Uraster ruthveni given by Forbes*
appears to be of a species generically identical with the American
1 Pal. Ohio, vol. 1, 1873, p. 67. s Palseontographica, vol. 36, 1890, p. 219.
2 Handb. Pal., vol. 1, 1879, p. 453. * British Org. Rem., dec. 1, 1849, p. 1, pi. 1, fig. 1 .
REVISION OF PALEOZOIC STELLEROIDEA. 177
Stenaster pulchellus Billings and this is the general consensus of opinion.
The writer unfortunately had no material of the genotype of Uras-
terella for study. However, since U. grandis is closely related to U.
pulchella and shows so much more detail, it is here accepted as the
genotype for the time being.
Professor Hall (1870) wrote: " In Stenaster = Urasterella, however,
we have the absence of adambulacral plates." This is nevertheless
not the case, for the corn-shaped marginal series in S. pulchellus are
the ambulacral plates, since weU-developed inframarginal plates, as
in Palseaster, are absent in mature Urasterella. The inframarginals
are, however, seen to be present in U. pulchella or in the distal ends
of the rays in other species, though proximally they are so modified
as not to be readily distinguishable from the ambital plates. In the
axils of the young of U. ulrichi the inframarginals are well developed,
along with single interbrachial marginal plates, but in mature speci-
mens of other species none have been seen.
Of Roemeraster the writer has seen three excellent specimens skil-
fully prepared by Sttirtz which clearly show that this genus has the
generic structure of Urasterella. Stiirtz 1 describes this genus as
having "strong marginal plates" inside of the "small, cornered adam-
bulacrals." Our specimens do not show such plates, and if present
they can not be adambulacrals. This fact can be demonstrated
in a well-preserved distal end of a ray, where two columns are present
on each side of the ambulacrals. The innermost column of closely
adjoining plates increase in size, retain iheir position beside the ambu-
lacrals, and become the most prominent of the entire animal, while
the outermost column soon change in aspect, the plates become
separated and assume the characters of the ambital pieces. This
is likewise true in the young of U. ulrichi where the axillary areas
also have distinct interbrachial marginal and two larger proximal
inframarginal plates. Passing distally, the latter also soon change
then- form, are separated one from another and are not distinguish-
able from the ambital plates. Stiirtz further describes and figures
five interbrachial ossicles in mature Urasterella asperula which are
also well developed in the young of U. ulrichi. Urasterella is an
easily recognized genus and the triangular ambital plates drawn out
into short blunt erect rods are peculiar to it, a character again re-
peated in Roemeraster. Under these circumstances the latter appears
to be indistinguishable from Urasterella. For other remarks see
U. asperula. On the other hand, Gregory regards Roemeraster as a
good genus and places it erroneously among the ophiuridlike asterids
in the family Tseniasteridse.
i Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, p. 52.
178 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
In regard to Protasteracanthion Stiirtz, see remarks under Uras-
terella asperula, and for Salteraster Sturtz see U. asperrima.
Urasterella differs from Stenaster as redefined in this work in having
long, slender, flexible rays which are never short and petaloid as in
the latter genus. Again, the adambulacral pieces of Urasterella are
very numerous, thin, and coin-shaped, while in Stenaster they are few
in number and comparatively thick; the ambulacral plates are
wholly different.
The long, slender and flexible rays, the apparent absence of in-
framarginal columns, and the absence of distinct interbrachial
plates (these are not the same as the single axillary pieces seen in
U. asperula) at maturity readily distinguish this genus from all other
Paleozoic genera.
For the origin of Urasterella see U. ulrichi.
Eoactis. After this book was completed appeared the work of
Spencer, with the new genus Eoactis. It is probable that the genus
is good and can be distinguished from Urasterella, but the necessary
comparisons can not now be made.
URASTERELLA PULCHELLA (Billings).
Plate 28, figs. 3, 4; plate 30, fig. 5.
Palseaster pulchellus BILLINGS, Geol. Surv. Canada, Rep. Progress for 1853-1856,
1857, p. 292.
Stenaster pulchellus BILLINGS, Geol. Surv. Canada, Can. Org. Hem., dec. 3, 1858,
p. 79, pi. 10, fig. 2. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2,
pt. 1 (Palaeontogr. Soc. for 1861), 1862, p. 28.
Urasterella (Stenaster) pulchellus HALL, Twentieth Rep. N. Y. State Cab. Nat.
Hist., 1868, p. 289; rev. ed., 1868=1870, p. 332.
The largest specimen from Middleville, New York (Hall collection),
measures: 11 = 52 mm., r=ll mm., R = 4.8r. An individual of the
usual size from Trenton Falls, New York (Mus. Comp. Zool., No. 31),
measures: R = 30 mm., r = 4. mm., R = 7.2r. One from Ottawa
(Geol. Surv. Canada collection) measures: R = 25 mm., r = 4 mm.,
R = 6.1r. The smallest specimen seen (Mus. Comp. Zool., No. 36)
measures: R = 7 mm., r = 1.5 mm., R = 4.7r. Width of rays at
base in mature examples from 4.5 mm. to 7 mm.
Original description. " Diameter, 2J inches; rays subcylindrical, 2J
lines in width at the base, with a length of 1 inch; disk 3J lines in
diameter; grooves narrow, bordered throughout by narrow oblong
plates, nine in the length of two lines; the length of these plates in a
direction transverse to the rays is about 1 line; near the disk there
appears to be but one row of marginal plates."
Formation and locality. Restricted to the Trenton limestone.
Most abundant at Trenton Falls, New York. From this locality
REVISION OF PALEOZOIC STELLEROIDEA. 179
there are two specimens in the New York State Museum and six
in the Museum of Comparative Zoology, Harvard University, col-
lected by Dr. C. D. Walcott (Nos. 31-36). The American Museum
of Natural History has one specimen from Canajoharie, New York
(No. 660) ; in the Hall collection, Albany, there is a specimen from
Middle viUe, New York. At Hull, Canada, Mr. W. R. Billings col-
lected two specimens; the type and another specimen were found
by E. Billings at Ottawa, Canada; these are now in the Museum of
the Geological Survey of Canada at Ottawa.
Remarks. This graceful species has the essential structure of
U. grandis and will therefore not require a detailed description.
U. pulchella is from a lower geological horizon, is probably the an-
cestor of U. grandis and has far less numerous abactinal plates,
which on the disk reveal a primitive arrangement reminding one of
Hudsonaster. In the center of the disk there is a single plate around
which is a first ring of 5 basal radial plates followed by another with
15 plates. The plates of the second ring are a little more conspicu-
ous than any other abactinal plates and are disposed as follows:
Five are radial in position and are the second basal plates of the
5 radial columns, while the other 10 are disposed interradially in
pairs immediately on each side of the second basal radials. The
plates interradial in position are the basals of the supramarginal
columns. Between the central ossicle and first ring there appear
to be a few small accessory plates.
The arrangement of the disk pieces above described is plainly
preserved in a very young example (No. 36, MILS. Comp. Zool.).
The radial columns remain the most prominent while the supra-
marginals a little distally from the base of the rays are not readily
distinguishable from the other columns. Here and there an accessory
piece may be inserted between the radials and supramarginals but
the greater width of the rays is attained by increasing the size of
these columns and by the introduction of ambital columns outside
of the supramarginals. In a mature ray on each side of the radials
proximally, there appear to be not more than four columns of plates
but it is possible that a few pieces of a fifth column may be developed.
The form and spinosity of the abactinal plates is as in U. grandis,
except that the nonarticulating spines are less long in U. pulchella.
Inframarginal columns at first view do not appear to be present
but a careful examination near the distal end of a ray reveals two
columns of nearly equal-sized plates on each side of the ambulacral
furrow. Here these plates closely adjoin, the adambulacrals are
elongate subquadrangular and the inframarginals quadrangular in
shape. Proceeding proximally the former become more prominent
and wider but remain throughout closely adjoining. The inframar-
ginals, however, gradually pass over to the abactinal side although
180 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
they remain beside the adambulacrals, become more and more
spinose, longer than wide, and eventually stellate in form and then
are inseparable from the abactinal plates. There are about 42 adam-
bulacrals in each column.
U. pulchella is readily distinguished from U. ulricJii and U. grandis
in being smaller, with more slender and graceful rays, and in its fewer
columns of plates. From U. Jiuxleyi it is distinguished by the less
convex rays and the far smaller number of plates.
Cat. No. 60610, U.S.N.M.
URASTERELLA GRANDIS (Meek).
Plate 27, figs. 6-8; plate 28, figs. 1, 2; plate 30, figs. 1-4.
Stenaster grandis MEEK, Amer. Journ. Sci., ser. 3, vol. 3, 1872, p. 258; Geol. Surv.
Ohio, Pal., vol. 1, 1873, p. 66, pi. 3 bis, figs. 7a-7c. MILLER, N. Amer. Geol.
Pal., 1889, p. 283, fig. 432.
Urasterella grandis MEEK, Geol. Surv. Ohio, Pal., vol. 1, 1873, p. 67.
Palseaster harrisi MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 2, 1879, p. 117,
pi. 10, figs. 2, 2<z.
Original description. "Attaining a very large size, with the body
or disk comparatively small, or only of the breadth of the united
inner ends of the five rays. Rays long, slender, gradually tapering,
and very flexible, widest at their immediate connection with the
body, where they seem to be more or less depressed, but becoming
more nearly terete farther out. Dorsal side of body and arms com-
posed of numerous subtrigonal pieces that rise into pointed tubercles,
or sometimes assume almost the character of short spinules, and
are arranged hi quincunx, so as to form about eight rows near the
middle of the rays; those of the outer two rows being a little larger
than the others. Dorsal pores apparently rather large, and passing
through between the concave sides of contiguous pieces. Ventral
side of body unknown. That of the rays composed of the usual single
row of transverse adambulacral pieces on each side of the well
defined, rather deep, and moderately wide ambulacral furrows.
Adambulacral pieces rather more than twice as long as wide, with
their longer diameters at right angles to the ambulacral furrows,
and rounding over from end to end so as to be most prominent in
the middle; while they do not connect with each other by flat sides,
but have little projecting processes, and corresponding sinuses,
apparently for the purpose of imparting greater flexibility to the
rays."
Emended description. The largest specimen being doubled over
actinally does not permit of exact measurements, but it is, as near
as can be determined, as follows: R = 93 mm., r = 9 mm., R = 10.3r.
Another specimen spread flatly measures: 11 = 49 mm., r=6.5 mm.,
11 = 7.67-. The smallest specimen (holotype of Palseaster Jiarrisi]
REVISION OF PALEOZOIC STELLEEOIDEA. 181
measures: R = 10 mm., r = 2.5 mm., R = 4r. Width of rays at base
in mature examples from 7 mm. to 8.5 mm.
Rays at maturity very long, depressed subcylindrical in outline,
slender and tapering slowly. Disk very small, formed by the united
inner ends of the rays.
Abactinal area reticular, consisting of very numerous, highly
conical, stellate plates, the radial columns and the plates of the disk
most prominent, while the other ray plates (supramarginal and
ambit al) are smaller and irregularly triangular in form. All plates
are more or less drawn out into stout, blunt, nonarticulating spines
which are best preserved along the sides of the rays. On the disk
the ossicles are arranged in a few concentric rows while on the rays
they are in columns and in quincunx. Near the base of a ray in the
largest specimen there are about 13 columns, diminishing to about
4 at the distal ends.
Madreporite of medium size, lobate, radially crenulostriate, situ-
ated between two adjoining rays and about midway between the
margin and center of disk.
Adambulacral plates slightly overlapping the ambulacral columns,
diminishing in size distally, coin-shaped, arranged on edge, and vary-
ing in different specimens from 60 to 110 in each range. In the
youngest known example there are about 20 plates in a column.
Each piece bears three or four short, finger-shaped, articulating
spines, two or three outwardly disposed, and one placed near the
inner base between adjoining plates. Along the outer margin of
the adambulacral plates and abactinally to the thick finger-shaped
spines there is a row of long, slender, flattened and longitudinally
grooved spines, two to each ossicle. Orally the coin-shaped adam-
bulacral plates are somewhat modified, stand less erect and terminate
in pairs of subquadrangular oral armature pieces.
Ambulacral grooves wide, shallow, and furrowed medially by a
narrow angular gutter. Ambulacral plates of adjoining columns
opposite or very slightly alternating, the pieces narrow, equaling
in number the adambulacrals, slightly overlapping each other
proximally and partially interlocking in the medial gutter. The
ambulacral plates continue in undiminished size to the compara-
tively small mouth, are highly convex and club-shaped, with the
attenuated ends laterally disposed. The podial openings are between
the attenuated ends of adjoining plates.
Locality and formation. In the Richmond formation at Richmond,
Indiana (type-locality), and in the vicinity of Waynesville, Ohio,
where Mr. Harris secured six more or less mature and three young
individuals, besides numerous fragments. A ray of this species
was found by Mr. U. P. James near Dayton, Ohio. In the Gurley
collection of the University of Chicago there is a young specimen
(No. 10978) from near Ridgeville, Warren County, Ohio.
182 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Remarks. The type-specimen of Palseaster Jiarrisi Miller is undoubt-
edly a Urasterella, and there are no characters other than size by
which this individual can be separated from U. grandis. Of about
the same age are three specimens in the Harris collection and another
in the University of Chicago, and all were originally referred to
P. Jiarrisi. One of these is nearly again as large as the type, is about
one-half the size of U. grandis, and measures 49 mm. along the greater
radius. The only marked difference between P. Jiarrisi and U.
grandis is the smaller number of adambulacral plates in the former,
a feature common to the young of many fossil starfishes when com-
pared with the adults of the same species. In fact, the variation in
the number of these plates in the two largest examples (64 and 110)
is nearly as great as between the type of Palseaster Jiarrisi (22)
and the smaller of the two largest specimens of U. grandis (64).
U. grandis is closely related to U. pulchella (Billings) of the
Trenton limestone. These species are, however, easily distinguished,
not only by the different geological occurrences, but in that U. grandis
attains a much larger and more robust growth. The latter has also
many more abactinal ossicles with longer nonarticulating spines,
and the adambulacral columns have from 60 to 110 plates, instead
of from 36 to 60 as in U. pulchella. U. grandis appears to be a
direct descendant of the latter and has varied only in attaining a
larger size and a greater number of plates.
U. grandis is also closely related to U. ulrichi, and both are nearly
of one size. The latter, however, has the rays highly convex abac-
tinally, the plates are smaller but as strongly, spinose, and there are
two columns of larger central plates, instead of one as in U. grandis.
In the latter in the mid-length of the ray there are about 9 plates
in a column in 10 mm., while in U. ulrichi there are from 11 to 12 in
the same space.
Cat. Nos. 40885, 40887, U.S.N.M.
URASTERELLA HUXLEYI (Billings).
Plate 29, fig. 2.
Stenaster huxleyi BILLINGS, Geol. Surv. Canada, Pal. Foss., vol. 1, 1865, p. 213,
fig. 197.
E = 58 mm., r = 5.5 mm., 11 = 10. or. Width of ray at base, 6 mm.
Original description. " Deeply stellate, 4 or 5 inches across; body
small, less than half an inch in diameter; rays long; flexuous, sub-
cylindrical, apparently angulated along the medium line on the upper
side, uniformly tapering to an acute point. On the dorsal side the
rays are covered by a multitude of small subangular plates, each
from one-fourth to one-third of a line wide. The central part of the
body is not well preserved in the only specimen collected; but it is
REVISION OF PALEOZOIC STELLEROIDEA. 183
evident from a view of the plates which remain that they are here
larger and more convex than those of the rays.
" Diameter of the body, five lines. Length of each ray, If inches.
Width of the rays at their junction with the body, 3 lines. Depth
of the rays from the dorsal to the ventral side of the body apparently
somewhat less than the width. The total breadth of the specimen,
if the rays were straightened out, would thus be about 4J inches.
"From the manner in which the rays are curved, it is evident that
they possessed a considerable amount of flexibility. The specimen
is somewhat distorted by pressure, but a small portion of one of the
rays near the body seems to retain its natural shape, and it is here
obtusely angulated along the median line. The transverse section
of the ray should be, therefore, subpentagonal. There is still, how-
ever, some doubt on this point. The under side is unknown. " In
places where the specimen is broken the adambulacrals are seen to
be as in other species of Urasterella; there appear to be about 60 or
more of these plates in each column.
Locality and 'formation. The only specimen of this species was
found by Mr. J. Richardson in strata of Chazy age (bed I of the
Newfoundland section) at Point Rich, Newfoundland. The holotype
(No. 554) is in the Museum of the Geological Survey of Canada, at
Ottawa, which has furnished the photograph illustrating the specimen.
Remarks. The holotype was seen at Ottawa and the original
illustration found to be a fairly accurate reproduction of the charac-
ters of the fossil. The abactinal side of the disk has a central disk
plate that is large and tumid. Around it is a circle of six smaller,
highly convex plates, and at the base of each ray medially there is
another single large tumid plate, the basal radial. The abactinal
side of the rays is highly convex and somewhat angulated medially.
The ossicles are small and distinctly arranged in quincunx, of which
there are about six plates in each diagonal row near the base of a ray.
In other words, the abactinal characters of U. huxleyi are very much
like those of U. ulrichi, with the differences that in the latter species
the prominent disk plates are smaller and there are more columns
of smaller ossicles on the stouter and longer rays. Further, in U.
ulrichi there are two columns of larger abactinal plates along the
central area of the rays, while in U. huxleyi all the ray ossicles are
of equal size.
TJ. pulchellus has less convex and more slender rays, and far fewer
abactinal plates.
URASTERELLA ULRICHI, new species.
Plate 29, fig. 1; plate 30, figs. 6, 7.
The smallest specimen measures: R = 14 mm., r = 4 mm., R = 3.5r.
The best preserved specimen measures: R = 45 mm., r = 7.5 mm.,
R = 6/*. The largest specimen measures: R = 78 mm., r=12 mm.,
184 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
R = 6.5r. Width of ray at base in the different individuals 4, 7.5,
and 9 mm., respectively.
Rays short and tapering rapidly when young but at maturity very
long, highly convex abactinally, and tapering very slowly. Disk
comparatively large in the young but small at maturity and formed
by the united ends of rays.
Abactinal area of disk and rays composed of numerous small, tumid,
or highly spinose plates which in the rays are arranged distinctly
in columns and in quincunx. In the angles between all of the
plates are left subcircular small openings. On the rays medially
there are two columns of large convex plates, variously stellate and
not produced into short, blunt rods. These are the supramarginals,
and between them are a few widely separated plates of the radial
columns. In the young the two proximal radial plates are sepa-
rated from each other by from one to three supramarginals, while
the third plate is separated from the second by three or four supra-
marginals. In a mature specimen there appear to be about eight
of these radial plates on each ray and distally they are separated
from one another by five supramarginal ossicles. On each side of the
supramarginal columns are from one (distally) to five (proximally)
columns of smaUer or accessory ambital plates. In a full-grown
specimen there are 11 to 12 in 10 mm., triangular in outline, and
drawn out into short, stout, blunt rods. In the center of the disk
in the youngest specimen (probably also at maturity) there is a
single large depressed plate around which is a first circle with 6
plates followed by a second having 16 pieces. Of the latter, 5 are
basal radials, 10 basal supramarginals, and 1 the madreporite.
All of the disk plates are variously stellate, and are the largest plates
of the abactinal area, the size gradually diminishing distally along
the rays.
Madreporite concentrically striate, somewhat smaller than the basal
supramarginals, between two of which it is placed.
Adambulacral plates seen only in the young specimen, very strong,
carinated, subquadrate, wider than long, and about 19 or 20 in each
column 10 mm. long, not counting the oral armature pieces, which are
subtriangular in outline. Inside of each pair of oral armature plates
is placed a very stout but short subcircular spine which has not been
observed in other species. No other spines are preserved.
Ambulacral furrows very narrow and deep, concealing the ambu-
lacral plates.
In the axils of the youngest specimen, outside of the adambulacrals,
are situated three prominent plates, the proximal one being the
smallest. On each side of these along the ray is a single column of
plates which rapidly diminish in size and separate with increasing
interspaces distally. Their number can not be made out, but they
appear to continue to the tips of the rays, where plates occur that
REVISION OF PALEOZOIC STELLEROIDEA. 185
are not ambital. In the mid-length of the ray, however, these plates
greatly resemble the ambitals. Judging from the manner in which
they appear distally, their form and constant position beside and
outside the adambulacrals leads to the conclusion that they are the
inframarginals. Whether the inframarginals and interbrachial mar-
ginals are present at maturity cannot be determined.
Formation and locality. Three specimens from the top of 'the Black
River limestone immediately beneath the Black River shale, or from
the Vanuxemia bed of the Black River (Mohawkian) group at Minne-
apolis, Minnesota. The specific name is after the distinguished
paleontologist, Dr. E. O. Ulrich, who found all the specimens. They
are in the United States National Museum.
Remarks. This species is distinguished from other Ordovicic
Urasterellas by the few and widely separated plates of the radial
columns, and by the prominence of the supramarginal columns,
which in the other species are not distinguishable from the ambitals.
The convexity of the rays abactinally is also greater than in the
other species excepting U. Jiuxleyi, but that species is further dis-
tinguished in that it has smaller plates, there being from 7 to 10 in
5 mm., while in U. ulrichi there are only 5 to 6 in the same space.
U. ulrichi and U. grandis attained a similar large size. The former,
however, is from a much lower horizon, has very convex rays abac-
tinally, two columns of larger central plates (supramarginals) instead
of one (radial), and the plates are smaller and therefore more
numerous.
The young of this species are very interesting in that they show
distinctly the presence of large inframarginal and interbrachial
marginal plates in the axils. In U. pulchella none of these plates
have been seen in the axils, but the inframarginals are believed to
be present in the very small plates which are indistinguishable from
the ambital pieces situated along the greater length of the rays out-
side of the adambulacrals. The same appears to be true for U. grandis.
The young of U. ulrichi in its general structure reminds one much
of Mesopalseaster sJiqfferi, and it is probable that Mesopalseaster and
Urasterella are derived from the same stock, which also gave rise to
Hudsonaster. The latter retains throughout life the primitive plate
structure, while Mesopalseaster, and particularly Urasterella, develop
in various ways an abundance of accessory plates.
The best preserved specimen of U. ulrichi (pi. 29, fig. 1) has two
nearly fully grown rays and three short stumps. One of these short rays
is so well preserved as to indicate that this condition is not due to poor
preservation, but is apparently a case of accidental loss of parts
during life. The wound has been healed, but no regeneration of lost
parts has taken place, as is so common in similar losses among the
living starfishes.
Cat. No. 60612, U.S.N.M.
186 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
URASTERELLA GIRVANENSIS, new species.
Plate 28, fig. 5.
Tetraster, sp. ind., NICHOLSON and ETHERIDGE, Mon. Silurian Foss. Girvan Dist.,
Ayrshire, fasc. 3, 1880, p. 325, pi. 21, figs. 9, 10.
? Eoactis simplex SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palaeontogr. Soc. for
1913), February, 1914, p. 30, pi. 1, fig. 4.
Urasterella girvanensis SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April,
1914, p. 44.
Description ~by N. and E. "Body stellate, with slowly tapering
arms, which increase little in width throughout their whole length [the
distal ends being unknown]; upper surface unknown, but the body
was convex and probably granular. [The pieces of the disk are seen
through the mouth and consist of many tiny plates arranged in more
or less regular circles about the slightly larger central ossicle.] Ave-
nues of the arms broad, and not bridged over by any of the plates [this
is a slip of the pen, for the ridged ambulacralia lie horizontal and
deep in the grooves and are directly opposite one another; the podial
openings are fairly large and in the usual position laterally]; marginal
ambulacral plates [ = adambulacrals] moderately convex, trans-
versely elongated, or oblong in form, and less in width than the
ambulacral plates; those in the angles of the rays are the largest, and
somewhat more elongated than the others; oral plates not visible."
The interbrachial areas are structurally very interesting because in
each one there is a single narrow but long axillary interbrachial
ossicle that distaily margins the axil, and in front of this is a minute
pair of ad ambulacral oral armature pieces. On either side of the
axillary interbrachial are the most prominent adambulacrals, about
five in number, being larger than the other distal ones, which are
considerably narrower and of fairly uniform size throughout the rays.
Formation and locality. A single specimen from the Upper Ordo-
vicic at Thraive Glen, Girvan, Scotland. A good wax squeeze of the
holotype was furnished by Doctor Bather. The original is in the col-
lection of Mrs. Robert Gray, at Edinburgh, Scotland.
Remarks. This species is clearly a Ur aster ella, a fact which was
also noted by Nicholson and Etheridge, as they state: "In some of
its characters it approaches very closely to T. (?) asperrimus, Salter,
sp." In its interbrachial skeleton U. girvanensis retains youthful
generic characters, seen in the well-developed interbrachial axillary
ossicles. After the above was written appeared Spencer's paper
above cited. Before the writer was aware of this work he had pro-
posed the name U. girvanensis. If we are treating of the same
species Spencer's name takes precedence.
Cat. No. 60611, U.S.N.M.
REVISION OF PALEOZOIC STELLEROIDEA. 187
URASTERELLA (?) ASPERRTMA (Salter).
Palseaster asperrima SALTER, Ann. Mag. Nat. Hist.,ser. 2, vol. 20, 1857, p. 325,
pi. 9, fig. 1. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1
(Palseontogr. Soc. for 1861), 1862, p. 24, fig. 15 (1).
Palseaster asperrimus SALTER, Mem. Geol. Surv. Gt. Britain, vol. 3, 1866, p. 289,
pi. 23, fig. 2.
Palseaster (?) asperrimus STURTZ, Palseontographica, vol. 32, 1886, p. 91.
fTetraster asperrimus NICHOLSON and ETHERIDGE, Mon. Silurian Foss. Girvan
Dist, Ayrshire, fasc. 3, 1880, pp. 320, 321, 324.
Salteraster asperrimus STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50,
1893, pp. 43, 60.
Nicholson and Etheridge wrote of this species as follows: It
"is an unsatisfactory species. The specimens in the Museum of
Practical Geology have the large transverse ossicles very apparent;
but as to whether there is a row inside or outside these, or both,
we are by no means certain. It appears, however, to have possessed
only four rows." Under Tetraster they compare this species with
their "T. sp. ind.," which is a true Urasterella (U. girvanensis) .
It seems best under these circumstances to refer this species to
Urasterella, it being apparently near U. grandis. It is found in the
Caradoc or Baia sandstones (Ordovicic), near Welshpool, North
Wales. Should it prove to be generically different from Urasterella ,
then the name Salteraster Stiirtz can be revived, as he names P. asper-
rimus as the genoholotype.
URASTERELLA (?) CONSTELLATA (Thorent).
Asterias constellata THORENT, Me"m. Soc. ge"ol. de France, vol. 3, 1838, p. 259,
pi. 22, fig. 7.
The original figure of this species does not permit of determining
its generic position. It is described as having but one range of plates
on each side of the ambulacral grooves, and the figure seems to
indicate the presence of a large disk with well-defined interbrachial
arcs. It seems to have more of the characters of Urasterella than
of any other genus.
The specimen was found in the Siluric ( ? Lower) strata in northern
France (Mondrepuis, L'Aisne).
URASTERELLA RUTHVENI (Forbes).
Uraster ruthveni FORBES, Mem. Geol. Surv. Gt. Britain, vol. 2, pt. 2, 1848, p. 436;
Mem. Geol. Surv. United Kingdom, dec. 1, 1849, p. 1, pi. 1, fig. 1; in McCoY,
British Pal. Foss!, 1851, p. 59. MURCHISON, Siluria, 1854, fig. 39-3.
Palxaster ruthveni SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 326.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palaeontogr.
Soc. for 1861), 1862, p. 25.
Urasterella ruthveni FORBES, in McCoY, British- Pal. Foss., 1851, p. 59.
This, the genotype of Urasterella, is unfortunately not well known,
but appears to be congeneric with U. grandis and U. pulchella. It
occurs in the Upper Ludlow beds (Siluric), near Kendal, Westmore-
land, England.
188 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
URASTERELLA HIRUDO (Forbes).
Uraster hirudo FORBES, Mem. Geol. Surv. Gt. Britain, vol. 2, pt. 2, 1848, p. 464;
Mem. Geol. Surv. United Kingdom, dec. 1, 1849, p. 3, pi. 1, fig. 4; in McCoy,
British Pal. Foss., 1851, p. 59. MURCHISON, Siluria, 1854, p. 221, fig. 39-2.
Palxaster hirudo SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 326.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palseontogr.
Soc. for 1861), 1862, p. 25.
Urasterella hirudo FORBES, in McCoy, British Pal. Foss., 1851, p. 59.
This small species has much the appearance of being the young
of U. ruthveni. The form is said to be gregarious and is associated
with that species.
URASTERELLA SELWYNI McCoy.
Urasterella selwyni McCoy, Geol. Surv. Victoria, Prodr. Pal. Victoria, dec. 1,
1874, p. 42, pi. 10, figs. 2, 2a, 3, 3a.
Original description. " Kays five, elongate, gradually tapering from
a little beyond the base, which is slightly contracted, angulated on
the upper side by a prominent ridge along the middle of each ray,
having a row of conical tubercular plates (about eight in two lines),
each side sloping on the dorsal aspect from the middle with about
three rows of conical tubercular plates rather smaller than the middle
row. The five axil plates small, ovate, triangular, very tumid.
Adambulacral plates large, extending to the tubercular margin,
transversely oblong, about twice as wide as long (about nine in two
lines). Ambulacral plates small, in a deep ambulacral groove.
Length of ray from mouth to tip, six lines; greatest width near base,
1J lines. Surface of plates granular.
"This beautiful species is easily known by its strongly angulated
rays on the dorsal side. The traces of oral plates are so very minute
and indistinct that I can not give their character.
"This is the first fossil starfish seen in Australia, and I dedicated
it to my old friend Mr. Selwyn, formerly Director of the Geological
Survey of Victoria, and now Director of the Geological Survey of
Canada, who collected it.
"This species is most nearly related to the Uraster ruthveni of
Forbes from the Upper Silurian grits of Kendal in Westmoreland.
"Common in the fine sandy Silurian beds of range on E. side of
commonage reserve, Kilmore."
Remarks. This is clearly a species of UrastereTla. ActinaUy it
has in each axil one fairly large axillary interbrachial and in this
recalls the same structure in U. girvanensis, new species.
URASTERELLA ASPERULA (Roemer).
Asterias asperula ROEMER, Palaeontographica, vol. 9, 1863, p. 146, pi. 24; pi. 26,
fig. 6; pi. 27.
Roemeraster asperula STURTZ, Palseontographica, vol. 32, 1886, p. 85, pi. 9, figs. 4, 5.
Protasteracanthion primus STURTZ, Palseontographica, vol. 32, 1886, p. 90, pi. 11,
figs. 3, 3a.
Remarks. Of this species there are two specimens in the United
States National Museum purchased of Stiirtz, and two more are
REVISION OF PALEOZOIC STELLEROIDEA. 189
in the Tale Museum Collection. These show both sides of the species.
U. asperula is said to be common in the roofing slate of the Lower
Devonic at Bundenbach, Germany.
There can be no doubt that Asterias asperula Roemer is a true
UrastereTla. All the generic characters are shown, even the longi-
tudinally grooved spines and the rodlike extensions of the ambital
plates. For further remarks on this species see generic discussion.
The abactinal structure of Roemeraster asperulus reminds one much
of UrastereTla pulcJiella of the Trenton, which is the least specialized
species of the genus. The single interbrachial plate retained to
maturity reminds one of the young of U. ulrichi. In this character
U. asperula appears to be primitive. In the proximal third of the
rays in U. asperula the radial columns are the most marked and the
plates continue to increase in size toward the disk. The supramar-
ginals are also well developed proximally, but the two or three
proximal plates are smaller and depressed, so that the ten basals
along with the five radials do not make so prominent a ring of large
ossicles as in U. pulcJiella or in the young of U. ulrichi. Distally
the rays in U. asperula have the plates of nearly equal size, although
the columnar and quincuncial arrangement is retained. The center
of the disk is occupied by a prominent central disk-plate and the
first ring consists of five radial plates as in U. pulcJiella.
Cat. No. 59382, U.S.N.M.
URASTE3ELLA, ne\7 species.
In the paleontological museum of Cornell University there is a
large slender-rayed specimen having the general form and structure
of Urasterella. There is, however, so little of the actinal detail pre-
served that nothing more can be done than to indicate the presence
of this species.
The specimen was found by Mr. J. B. Woodworth at Brookton,
near Ithaca, New York, in the Upper Devonic.
URASTERELLA MONTANA (Stschurowsky).
Palseaster montanus SCHONDORF, Palseontographica, vol. 56, 1909, p. 324, pi. 23,
fig. 1; pi. 24, figs. 20-22 (complete bibliography cited here).
In general this form recalls U. grandis but is somewhat smaller and
stouter. The actinal side alone is known, showing a wide ambu-
lacral furrow bordered by stout columns of narrow adambulacrals.
All of the ossicles of the ambulacrals and adambulacrals are opposite
one another. Oral armature decidedly adambulacral in composition.
Interradial areas described as having "a few plates." A section
through a ray (Schondorf s fig. 22) shows that abactinally on each
side of the radialia there are four columns of ossicles, or nine in all.
Dimensions, about: E, = 35 mm., r = 6-7 nun., R 6r. Width
of ray at mid-length about 6 mm.
190 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Formation and locality .The holotype is from the oldest Upper
Carboniferous (Moscovian) of Mjatschkowa, near Moscow. The
specimen is now in the Bergakademie at Berlin, Germany.
CALLIASTERELLID^E, new family.
Calliasteridse SCHONDORF, Jahrb. nassaulsch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 251.
Cryptozonia with ophiurid expression; a small disk with primitive
plate arrangement and five long, very flexible rays, with radialia
and supramarginalia. No interbrachial areas. Ambulacrals oppo-
site.
Contains :
CalliastereUa, new name.
Genus CALLIASTERELLA, new name.
Plate 31, fig. 3, text fig. 11.
Calliaster mirus TRAUTSCHOLD (not Calliaster Gray 1840, Echinoderma), Nouv.
Me"m. Soc. imp. Nat. Moscou, vol. 14, 1879, p. 108, pi. 13, figs. 3a-3ft, 4.
SCHONDORF, Palaeontographica, vol. 56, 1909, p. 327, pi. 23, figs. 2-5; pi. 24,
figs. 1-18; text fig. on p. 328.
CalliastereUa SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914,
p. 14-
A cryptozonian asterid with the general expression of the ophiurid
Ony chaster.
Description. Disk relatively small, primitive, with rather large,
decidedly spiculate ossicles. Rays five, long and slender, round
in section. R = about 120 mm., r = 30 to 35 mm., R = 4r. Width
of ray at mid-length about 15 mm.
Abactinal disk plates in form and position as in figure 11. There
is a small central five-rayed plate with the rays in the position of the
five arms. Around the central plate is a first ring of five larger basal
radial plates. Then comes a second ring of ten plates, five of which
are the second basal radialia; the other five are interradial in posi-
tion and give rise hi the next ring to the ten basal inframarginal
ossicles. The third ring has five radialia and ten basal inframargi-
nals. All of the ossicles in the three rings are variously six-rayed,
with large rounded openings between the adjoining spicular ends.
Between each pair of basal inframarginals are inserted single elon-
gate interradial disk plates.
Madreporite unknown.
The rays abactinally have a medial column of transverse radialia,
and alternating with these on either side are columns of supra-
marginals. These ossicles are of peculiar construction, enabling
the rays to flex about as in Onychaster.
REVISION OF PALEOZOIC STELLEROIDEA. 191
Ambulacralia small, adambulacralia larger, both very numerous
in each column and in their disposition opposite one another. There
are no inframarginals.
Oral armature of Cryptozonia construction and very much as hi
living Asterias glacialis.
Ocular plates probably not present.
Genoholotype and only species. C. mirus (Trautschold) (cita-
tions as above). The holotype and three other fragments are from
\
R
FIG. ll.-*-THE PLATES OF THE DISK OF CALLIASTERELLA MIRA. SCHEMATIC. AFTER SCH&NDORF. 1, CEN-
TRODORSAL; 2, BASAL RADIALIA; 3, SECOND RADIALIA; 4, THIRD RADIALIA; 5, BASAL INFRAMARGINALS; 6,
PAIRED INFRAMARGINALS; a, POSSIBLE POSITION OF ANUS; mdp, PROBABLE POSITION OF MADREPORITE.
the Upper Carboniferous (Moscovian), at Mjatschkowa, near Moscow.
The holotype is in the University of Petrograd.
Remarks. There is no other Paleozoic asterid worked out in such
detail as is Calliasterella. Not only is the gross skeleton known, but the
detailed construction of all of the essential ossicles and spines as
well. Quarto pages of text and two plates are devoted to the skeleton.
The dorsal skeleton retains primitive construction, and although
the ossicles are much modified, their general orientation is that of
the older Phanerozonia.
COMPSASTERIM;, new family.
Specialized Cryptozonia with decidedly spicular abactinal skele-
ton, recalling that of Asterias. Ambulacra opposite or slightly
50601 Bull. 8815 - 13
192 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
alternating. Rays five, long, thick, and rounded in section. Disk
not large.
This family contains:
Jaekelaster Sturtz.
Compsaster Worthen and Miller.
Genus JAEKELASTER Sturtz.
Jaekelaster SitfRTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900, pp.
235-236, pi. 4, figs. 13-16.
Generic characters. Rays five, thick and long, 55 mm. in length,
16 mm. in greatest breadth, rounded distally. Disk large. Both
rays and disk devoid of marginal plates.
Abactinally without columns of plates, entire surface decidedly
and abundantly covered with "paxillae." Madreporite small,
subcentral, radiately stria te.
Ambulacral furrows wide, widest at about mid-length and nar-
rowing both distally and proximally. Two col mns of podial
openings. Ambulacralia narrow, L-shaped, slightly overlapping
and slightly alternating proximally. Adambulacrals small, di-
rectly opposite the ambulacrals. Outside of the rays the integu-
ment bears spines and "paxillae" abundantly.
Genoliolotype and only species. J. petaliformis Sturtz (citations
as above). Lower Devonic of Bundenbach, Germany.
Remarks. This genus is most closely related to Compsaster ,
in fact the general appearance of the two is very much alike.
Genus COMPSASTER Worthen and Miller.
Plate 31, figs. 1, 2.
Compsaster WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883, p. 327.
Original description. "Body stellate; central area or disk com-
paratively small; rays large, long, more or less fusiform; grooves
deep and bordered by numerous thin, subcircular, adambulacral
plates. The ventral side of the typical species shows about six
rows of plates upon each side of the ambulacral furrows, which
have a regular, transverse as well as longitudinal arrangement.
"This genus is distinguished from all others in the family [their
Palseasteridse] by the number and form of the adambulacral plates,
by the great number of disk plates upon each side of the ambula-
cral furrows, and by the general form of the body and rays."
GenoTwlotype. C. formosus Worthen and Miller. It is also the
only named species of the genus.
The adambulacral plates in Compsaster appear to be much as
in UrastereUa and both genera agree in having no distinct inframar-
REVISION OF PALEOZOIC STELLEROIDEA. 193
ginal plates. The ambit al plates are very numerous, showing a dis-
tinct progression to far more numerous columns of plates than in
Urasterella. They also appear like those in this genus. It is prob-
able that the abactinal area when it is known will be seen to have
spicular ossicles like those of the ambital areas.
COMPSASTER FORMOSUS Worthen and Miller.
Plate 31, figs. 1, 2.
Compsaster formosus WORTHEN and MILLER, Geol. Surv. 111., vol. 7, 1883, p. 327,
pi. 31, figs. 2a, 26.
Original description. "Body deeply stellate; central disk com-
paratively small; rays rigid, large, fusiform, more than twice as long
as the diameter of the central disk, and terminating abruptly in an
obtuse point. The typical specimen furnishes the following meas-
urements: Diameter of the body, 0.72 inch; length of ray from cen-
tral part of disk, 1.78 inches; diameter of the ray at its junction with
the body, 0.41 inch.
"Ambulacral grooves deep; ossicles rather small; adambulacral
plates very numerous, and consisting of series of thin plates upon
each side of the ambulacral furrows, which are rounded upon the
exterior, prolonged below, and each bear a short, sharp spine on the
upper part of the inner side, directed toward the apex of the ray.
There are about eighty adambulacrak upon each side of the furrows,
and they dimmish very slowly in size toward the apex, after passing
the fusiform enlargement of the ray.
"The disk-plates abut upon the adambulacrals in a straight line;
they are subquadrangular, or polygonal, and each bears a strong
central spine. Six longitudinal rows appear on the ventral side on
each side of the ambulacral furrows; they are somewhat uniform in
size, arranged in transverse, as well as longitudinal order, and are
about two-thirds as numerous in a longitudinal row as the plates in
an ambulacral series.
"Oral plates, madreporiform tubercle, and dorsal side unknown,
except so far as we may judge of the spinous character of the plates
from those visible on the ventral side.
"This large and beautiful species is so distinct from any hitherto
described that no comparison is necessary for the purpose of identifi-
cation.
"Position and locality. Okaw bluffs, between Chester and Kaskas-
kia, Randolph county, Illinois; from the second division of the Chester
limestone.
"Illinois vState coUection of 1880, by A. H. Worthen, No. 2476."
The writer has not been able to consult the type-specimen.
194 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
COMPSASTER, new species.
Professor Weller collected an excellent asterid at Fountain Creek,
Waterloo, Illinois (No. 14395, University of Chicago collection),
showing the entire actinal side of a form that is nearest to O.formosus.
It is, however, a smaller and more slender species, but until the
abactinal surfaces of both species are known, one can not say that
they are certainly of the same genus.
Measurements : R = 20.5 mm., r = 4.5 mm.
SCHUCHERTIID^E, new family.
Specializing Cryptozonia with well-developed pentagonal inter-
brachial arcs. Inframarginals well developed in the proximal regions
where they remain adjoining the adambulacrals. Interbrachial
and abactinal areas composed of small, more or less closely adjoining
plates without regular arrangement.
Madreporite unknown.
Contains :
Schuchertia Gregory.
Remarks. This family is the most ancient and primitive of all
large-disked or pentagonal starfishes devoid of marginals. In
Schuchertia the first appearing inframarginals are retained through-
out life and remain adjoining the adambulacrals in the axillary
regions. These columns distally become less distinct and in the
outer third of the rays are not distinguishable as inframarginals
from the other smaller ossicles of the interbrachial areas. This
therefore indicates that during the growth of Schuchertia the infra-
marginals are in the process of elimination and in later forms seem
to be completely disguised or lost. Then the plates of the inter-
brachial and abactinal areas, while small, are usually strong, rarely
spicular or stellate but never decidedly so, while in later genera the
ossicles either break up into smaller and smaller plates, even into
granules, or are spicular or thorny pieces. Even the adambula-
crals which are well developed in Schuchertia may become greatly
modified and almost eliminated.
In seeking for an ancestor for this family, it is plain that it came
from a stock like Hudsonaster. This genus actinally has columns
of ambulacral, adambulacral, and inframarginal plates; abactinally,
inframarginal, supramarginal, and radial columns of plates. In
addition there are five axillary marginals, and on the disk a ring of
five basal radial and five interradial plates giving rise to the radial
and supramarginal columns. The first step from Hudsonaster toward
Schuchertia would be a form of the former genus introducing small
ambital plates. This type is as yet unknown, but is suggested in
UrastereTla. The progressive introduction of ambital plates and no
REVISION OF PALEOZOIC STELLEROIDEA. 195
development of small interbrachial pieces will tend to keep the
inframarginals beside the adambulacrals. This is the case in Schuch-
ertia where the axillary marginals are also retained, but this genus
has progressed considerably beyond this stage, since the inframar-
ginals fail to develop in the later growth, while the supramarginals
and radials, as columns, are no longer distinguishable as such in
any part of the individuals. The tendency of departure from Hudson-
aster therefore seems to be toward the breaking up and elimination
of its essential structure into a mass of variously arranged small
pieces, except the ambulacrals, which remain as columns of stout
plates.
Genus SCHUCHERTIA Gregory.
Plates 32 and 33.
Palasterina BILLINGS (part, not McCoy or Salter), Geol. Surv. Canada, Can. Org.
Rem., dec. 3, 1858, p. 76.
Schuchertia GREGORY, Geol. Mag., dec. 4, vol. 6, August, 1899, p. 351.
Trentonaster STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900, pp.
224, 225 (based on same genoholotype as Schuchertia).
Original description. "Palseasterinidse with tessellate exoskeleton;
the rays are short and the general shape is pentagonal. R: r as 2:1.
Interradial plates numerous, polygonal. Adambulacral plates large
and hexagonal. Ambulacral furrow tapering uniformly toward the
distal end.
" Allied to Palseasterina by the general form of the body and the
absence of the marginal plates."
Emended description. Rays five, short and slender. General
form pentagonal. Disk large, with well-developed interbrachial
arcs.
Abactinal area composed of very numerous, small, tumid, non-
stellate plates which are arranged in quincunx or columns. There
are no radial, supramarginal, or ambital plates recognizable as such,
nor is the margin bounded by columns of inframarginals.
Madreporite indistinguishable amongst the abactinal plates.
Actinally the ambulacral furrows are very narrow, bounded by
columns of adambulacral plates. Adjoining these in the proximal
regions are distinct " axillary marginals" and inframarginals, the
latter distally becoming smaller, and these continue either closely
adjoining or are separated from one another by increasing interspaces.
In the latter case they are not readily distinguished from the inter-
brachial accessory plates.
Interbrachial areas occupied by numerous small accessory plates
and like those of the abactinal area with which they are continuous.
Ambulacral plates apparently slightly alternating.
Genoliolotype (of both Schuchertia and Trentonaster). Palasterina
stellata Billings. The generic description is, however, largely based
on S. laxata, new species.
196 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Distribution. Ordovicic and Siluric of North America. Con-
tains the following species:
S. stellata (Billings). Trenton, Ottawa, Canada.
8. laxata, new species. Richmond of Ohio.
S. ordinaria, new species. Basal Siluric.* Girardeau lime-
stone, Alexander County, Illinois.
Remarks. Schuchertia need only be compared with Petraster and
Palasterina. The former differs in that the incomplete columns of
inframarginals adjoin the adambulacrals and are not separated from
them by interbrachial accessory plates as in the other two genera.
Petraster is readily distinguished by the prominent columns of infra-
marginals bounding the entire animal and abactinally by the distinct
columns of radial, supramarginal, and ambital plates. In Schuchertia
the plates of the abactinal and interbrachial areas are alike and
not distinguishable into the kinds of columns just mentioned for
Petraster.
The fact that in ScJiuchertia the " axillary marginals 7 ' and infra-
marginals remain beside the adambulacrals seems to indicate its
origin in forms like Hudsonaster. The generic structure of the latter
is retained in Schuchertia, to which is added along the margin (mainly
axillary) a series of interbrachial accessory plates. The generic
structure of Hudsonaster is also retained in Petraster, but here, unlike
those in Schuchertia, the interbrachial accessory plates are developed
between the inframarginals and adambulacrals, forcing these columns
apart. The phylum starting in Schuchertia is not a prolific one, but
the other, Palasterinidse, whose inframarginals are true marginals,
is not only more prolific in genera but also has a longer geological
range. Beginning in Petraster of the Ordovicic, it is continued in
the Siluric in Lindstromaster and Palasterina.
SCHUCHERTIA STELLATA (Billings).
Plate 32, fig. 2; plate 33, fig.l.
Palasterina stellata BILLINGS, Geol. Surv. Canada, Rep. Progress for 1853-1856,
1857, p. 290; Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858, p. 76, pi. 9,
figs, la, 16. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1,
(Pakeontogr. Soc. for 1861), 1862, p. 27, fig. 16 on p. 26. QUENSTEDT, Petre-
factenkunde Deutschlands, vol. 4, 1876, p. 74, pi. 92, fig. 34.
Trentonaster stellata STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 56,
1900, pp. 217, 224, 225.
Description of 1858. "Pentagonal; disk about one-half of the whole
diameter; ambulacral grooves narrow and deep, bordered on each
side by a row of small, nearly square adambulacral plates; a second
row consisting of disk plates extends nearly to the end of each ray,
the remainder of the disk covered with smaller plates. All of these
plates are solid and closely fitted together; the disk plates in the
KEVISION OF PALEOZOIC STELLEKOIDEA. 197
angles in contact with the oral plates are much larger than any of
the others.
"In the only specimen in the collection the length of the rays
measured along the ambulacral grooves is three lines; number of
adambulacral plates on each side of the grooves sixteen; the rays
diminish somewhat rapidly in size, and terminate in a rounded point;
diameter of the disk four lines. The plates are all a little worn, so
that the character of their surfaces can not be observed; they were
probably nearly smooth."
Emended description. Measurement of the immature type-speci-
men: R = 6 mm., r = 3 mm. Larger specimen: R = 9 mm., r=4 mm.
Abactinal area composed of a series of plates which are more or
less closely adjoining and on the rays are arranged in distinct but
twisted columns. This twisting is due to the insertion of new columns
of plates, always on the right side, crowding the older ones to the left.
The plates on the rays are subquadrangular to elongate subquad-
rangular, increasing in size proximally, and on the disk are largest
and generally subcircular in outline, or faintly stellate. The plates
appear to be smooth. At the apex of the ray are two somewhat
larger plates followed by three columns and on each side of these
there are two other columns of ossicles. Near the beginning of the
interbrachial arcs the rays have from se^en to eight columns of plates.
Madreporite not distinguished among the abactinal plates.
Actinally the most conspicuous columns are the adambulacrals
bounding the very narrow ambulacral grooves. These ossicles are
subquadrangular in outline near the base of the column, but become
wider than long distally; there are about 15 in a column, terminating
proximally in two larger, prominent, wedge-shaped plates of the
oral armature. Interradially upon each pair of oral pieces is placed
a large, single, pentagonal plate (holds the position of axillary
interbrachials), against which rest two diverging inframarginal
columns, each with 13 or 14 ossicles, and these columns continue
adjoining the adambulacrals. Before attaining the distal ends of the
rays they gradually become smaller and pass over to the abactinal
side. Other actinal disk plates are also present, but apparently are
arranged in quincunx, and are smooth like those of the abactinal side.
Ambulacrals unknown.
Formation and locality. Ordovicic. The holotype (No. 1399, Geol.
Surv. Canada collection) showing the actinal area was found by Mr.
E. Billings, at Ottawa. Another specimen, also in the Museum of
the Geological Survey of Canada, and showing the abactinal area, was
found by Mr. J. S. Stewart on Governor General Bay, near New
Edinburgh, Canada.
Remarks. This little Ordovicic asterid is distinguishable from
other species with large interbrachial arcs in that it is devoid of all
marginal and radial columns of plates. From S. laxata it is differ-
198 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
entiated by its smaller size, less numerous plates, and by the rela-
tively greater size of the disk ossicles.
SCHUCHERTIA LAXATA, new species.
Plate 32, fig. 3* plate 33, figs. 2, 3.
The best specimen measures: R = 18 mm., r= 6 mm., R = 3r. The
University of Chicago individual: R = 23 mm., r= about 8 mm.
R = 2.8r. Other specimens indicate a growth twice as large as the
former one.
Rays short and slender. Disk and interbrachial arcs large but not
nearly so large as in Petraster speciosus. Six specimens are known,
four of which are poorly preserved and but a jumble of plates. The
specific name is given to indicate the generally separated condition
of the plates.
Abactinal areas of rays and disk composed of very numerous,
small but irregularly sized, subquadrate or diamond-shaped, slightly
pustulose plates. The arrangement is mainly in quincunx but a
columnar arrangement is also noticeable.
Madreporite not distinguishable amongst the abactinal plates.
Ambulacral furrows very narrow, hardly revealing the carinated
ambulacral plates. The podial openings are between the plates
along their lateral corners.
Adambulacral plates very convex, a little wider than long, with
the inner edge more or less pointed, against which terminates the
carina of the ambulacral pieces. There are about 15 of the former
plates in 15 mm. along the base of the columns. Actinally the adam-
bulacral columns are the most conspicuous, since the inframarginals
beyond the mid-length of the former columns appear not to be
separable from the adjoining interbrachial pieces. The inner side of
the adambulacrals bears tufts of short blunt spines.
Inframarginals recognizable as such only within the axils where
they adjoin the ad ambulacral columns. Just within each axil there
is a large single plate (axillary marginal) against which rest the infra-
marginal columns. These plates decrease rapidly in size distally and
are soon separated from one another so that the tenth piece from the
axil is indistinguishable in shape and size from the interbrachials.
Interbrachial accessory plates very numerous, smallest in the
inner axillary areas and thence increasing rapidly to a size maintained
throughout the interbrachial areas. In form they are either diamond-
shaped or subquadrate, centrally tumid, and each plate seems to have
borne one central and two or three lateral spines.
Formation and locality. Five specimens are in the Harris collec-
tion, all from the Richmond (Waynesville) , in the vicinity of Waynes-
ville, Ohio. A fine specimen showing the abactinal area (University
of Chicago collection, No. 10982) was found 7 miles north of Hamilton,
REVISION OF PALEOZOIC STELLEROIDEA. 199
Ohio. Another fragment was found by Mr. E. H. Vaupel at Oxford
in the same State.
Remarks. The only Upper Ordovicic species with which Schu-
cTiertia laxata can 136 confounded is Petraster speciosus. However,
there is one fundamental difference between them, so that even
fragments can be distinguished. In P. speciosus the inframarginals
are very conspicuous and bound the outer margin of the entire animal,
while in S. laxata there are no columns of marginal plates, since
the inframarginals closely adjoin the adambulacrals. In other
words, in the former species the interbrachial accessory plates are
between the adambulacrals and inframarginals, while in the latter
form these pieces, which are also far more numerous, are all outside
of the inframarginals. The generally disjointed condition of the plates
in S. laxata will probably always distinguish it from P. speciosus.
Cat. Nos. 60613, 60623, U.S.N.M.
SCHUCHERTIA ORDINARIA, new species.
A regular species of Schuchertia.
The holotjTpe shows the abactinal side only. R = 1 1 mm., r = about
6.3 mm.
This species is most closely related to S. laxata, but is smaller and
with a comparatively larger disk. Therefore the rays do not protrude
beyond the disk so far as in the latter form. Abactinal skeleton con-
sists of minute tumid ossicles that are more cut along their edges
than in S. laxata, and therefore more spicular.
The specimen can not be freed of the adhering clay sufficiently to
make an instructive photograph, and is therefore not illustrated.
Formation and locality. The holotype is from the basal Siluric
Girardeau limestone, Alexander County, Illinois. It is in the Gurley
collection of the University of Chicago (No. 10992).
Family PALASTEBISCID^E Gregory.
Palssbrisingidx STURTZ, Palaeontographica, vol. 36, 1890, p. 246.
Palsechinasteridse STURTZ, Palaeontographica, vol. 36, 1890, p. 246, also p. 247.
Palasteriscidse GREGORY, Treat. Zool., vol. 3, Echinoderma, 1900, p. 257.
Specialized Cryptozonia with very large interbrachial arcs. Abacti-
nal and interbrachial areas either with a smooth membrane or granu-
lar and thorny. No radial or supramarginal columns of ossicles.
Inframarginals not present in any position. Madreporite probably
always actinal. Ambulacrals alternating or opposite; adambulacrals
more or less modified, but remain adjoining the ambulacrals.
Contains :
Palasteriscus Sturtz.
Echinasterella Stiirtz.
Loriolaster Sturtz.
Cheiropteraster Sturtz.
For remarks on the course of development, see Schuchertiidse.
200 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genus PALASTERISCUS Sturtz.
' Palasteriscus STURTZ, Palaeontographica, vol. 32, 1886, p. 95, pi. 14, fig. 1; vol. 36,
1890, p. 223, pi. 28, figs. 23-236; pi. 29, fig. 24; Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, pp. 44, 61.
This genus differs from Loriolaster in being more decidedly stellate,
with longer and more slender rays and less extensive interbrachial
arcs. The abactinal and interbrachial areas are made up of innumer-
able, small, variously shaped, loosely interlocking, stellate plates.
These areas in Loriolaster appear to be smooth.
Madreporite very large, actinal.
The genoholotype, P. devonicus (same citations as above), is the
only species and is very rare in the Lower Devonic roofing slates of
Bundenbach, Germany.
Cat. No. 59389, U.S.N.M.
Genus ECHINASTERELLA Sturtz.
Echinasterella STURTZ, PalseontograpMca, vol. 36, 1890, p. 225, pi. 28, figs. 26, 26a,
27; Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 53, 73.
This form seems to be closely related to Palasteriscus , and without
material for examination it is difficult to point out the differences
supposed to be generic. Sturtz describes the ambulacrals as oppo-
site, but his figures 26 and 27 show that they are slightly alternating.
The genoholotype, of which three imperfect examples are known,
is E. sladeni (same citations as above). It is from the Lower Devonic
of Bundenbach, Germany. Another form which probably belongs
here is E. (?) darwini Clarke, from the Devonic of Brazil.
ECHINASTERELLA (?) DARWINI Clarke.
Echinasterella ? darwini CLARKE, Monog., I, Serv. Geol. Min. Brasil, 1913, p. 315,
pi. 27, figs. 9-12; Bull. N. Y. State Mus., No. 164, pi. 34, figs. 1-3.
Original description. "Of this fine starfish the collections con-
tain two specimens, one, that from which our illustrations are taken,
a sharp cast of an oral surface with the arms extended, in a shale
sufficiently compact to permit a squeeze to be made of the entire ex-
posure. The other is a considerably distorted individual also chiefly
with oral exposure. I have placed the species only provisionally with
Sturtz's genus EchinastereUa, which was described from the Bunden-
bach slates, feeling that in all probability both this species and its
associated Aspidosomaf pontis will be found on close analysis by
expert students of the Asteroidea to pertain to other genera. The
structural features presented by E.f darwini are as follows: The
oral surfaces of the arms are very broad, the ambulacral grooves
narrow, and the plates arranged opposite each other. The ambulacral
plates are turned obliquely inward and are broadened at their inner
ends. As preserved these show only their upper edges except where
REVISION OF PALEOZOIC STELLEROIDEA. 201
turned somewhat more obliquely along the food groove. At the
edges of these very broad ambulacral areas the adambulacral plates
project in a single row on each margin, as rounded nubs or thickened
elongated ridges, from each of which project not less than two and
probably three spines. In no place is the aboral surface of the starfish
presented, but the madrepore plate is distinctly shown in one of the
axillae, indicating its ambulacral position. This plate is round, con-
vex, obscurely radiopunctate, and somewhat ridged. The oral appara-
tus is indicated by thickened plates at the axillae, but their structure
in detail can not be made out. The example serving as the type of this
species has a radial length from center of mouth to tip of longest arm
of 54 mm."
Formation and locality. From the Devonic of Ponta Grossa,
Brazil.
Genus LORIOLASTER Sturtz.
Loriolaster STURTZ, Palaeontographica, vol. 32, 1886, p. 94, pi. 12, figs. 3, 3a, 4;
pi. 13, figs. 1, la, 2, 2a; vol. 36, 1890, p. 208, pi. 26, figs. 36, 4a; Verh. naturh.
Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 47, 63.
Generic description. Animal large, five-rayed, widely pentagonal,
with very large interbrachial arcs entirely inclosing the rays. No
marginal plates and seemingly with a parchment-like integument on
the abac tin al side and in the large interbrachial areas. The plates
seen on the abactinal side resemble the aspect of the actinal plates.
Madreporite unknown.
Ambulacral plates alternating, numerous, considerably wider than
long, with the podial openings indeterminate.
Adambulacrals about as numerous as the ambulacrals, sickle-
shaped, greatly elongated laterally and bearing club-shaped carinse
(the regular arrangement of the latter seems to preclude their being
articulating spines) which have on their outer lateral ends needle-like
spines.
Oral armature pieces large, described as ambulacral.
The only known species is L. mirdbilis Sturtz (same citations as
above), from the Lower Devonic of Bundenbach, Germany. It is
not a common species.
Cat. No. 59380, U.S.N.M.
Remarks. All of the Bundenbach material has been subjected to
great pressure so that nearly all of the finer details have been obliter-
ated and obscured by the adhering slate. However, L. mirabilis
can not be confounded with any of the associated species because of
the constant absence of the marginal plates and the smooth abactinal
and interbrachial membrane. The latter may have been made up of
very small calcareous pieces now obliterated by the great pressure to
which these fossils have been subjected. Also see Cheiropteraster.
202 BULLETIN" 88, UNITED STATES NATIONAL MUSEUM.
Genus CHEIROPTERASTER Sturtz.
Cheiropteraster STURTZ, Palaeontographica, vol. 36, 1890, p. 228, pi. 29, fig. 33;
pi. 30, fig. 32; Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 49, 68.
This genus has its nearest relationship in Loriolaster, from which
it differs in having a far larger oral opening and markedly different
ambulacral columns. These are small plates, in shape like the
vertebrae in teleost fishes, which alternate in adjoining columns and
do not closely adjoin medially. Podial openings unknown.
Madreporite small, actinally situated near the mouth.
Abactinal and interbrachial integument granular and thorny.
Oral armature pieces small, described as ambulacral.
GenoTiolotype and only species. C. giganteus Sturtz (same citations
as above). This large species is very rare in the Lower Devonic
roofing slates of Bundenbach, Germany.
Cat. No. 59381, U.S.N.M.
SCHOENASTERID^:, new family.
Specialized Cryptozonia with well-developed interbrachial arcs.
Abactinal area reticular, composed of numerous small plates. Inter-
brachial areas with distinctly imbricating plates. Oral armature
adambulacral. Adambulacrals well developed in the form of a rope
(hence the name, from schoinos, a rope). Ambulacral plates and
madreporite unknown.
Contains :
Schcenaster Meek and Worthen.
Genus SCHCENASTER Meek and Worthen.
Plates 32, 33, 35.
Schcenaster MEEK and WORTHEN, Proc. Acad. Nat. Sci. Philadelphia, vol. 12,
1860, p. 449; Geol. Surv. Illinois, vol. 2, 1866, p. 277. ZITTEL, Handb. Pal.,
vol. 1, 1879, p. 453.
Original description. " Animal consisting of a flattened pentagonal
disk, with the angles more or less produced in the form of rays or
arms, and the margins between the rays concave in outline, and
fringed with short, flattened, spine-like appendages, which are also
continued part of the way out along the lateral margins of the rays.
Upper side of rays composed of a number of alternating solid plates,
with the dorsal pores passing between them, while the angles between
the rays are filled with similar plates, forming the upper side of the
disk. Under side of the disk composed of numerous small plates,
very distinctly imbricating inward and laterally toward the ambu-
lacra. Ambulacral furrows (in the typical species) wide, deep, with
on each side a single row of comparatively stout, squarish, or oblong
adambulacral pieces, having an obliquely outward, imbricating
arrangement, so as to present somewhat the appearance of a twisted
REVISION OF PALEOZOIC STELLEROIDEA. 203
cord, as seen from below ; farther out these become the marginal pieces
of the free rays. There is ; however, no regular row of marginal
pieces to the disk between the rays. [In other words, there are no
infra- or supramarginals.] Oral pieces ten, anchylosed ( ?) in pairs, so
as to look like five pieces merely emarginate at the outer and inner
ends. (Other characters unknown.)
"Of course it is not possible to give a systematic diagnosis of a
genus like this, of which only imperfect fossil species are known.
All that can be done, in cases of this kind, is to give such of the more
prominent characters as happen to be visible in the particular speci-
mens accessible; while we can not always be sure, until better speci-
mens are obtained, and other species known, whether some of the
characters given may not be merely specific, or in other cases of
more than generic importance. At the same time, we have to regret
our inability to give any information in regard to some of the more
delicate parts that would be the first to claim the attention of the
zoologist in describing existing starfishes.
"In first publishing a description of the beautiful species forming
the type of this genus, we referred it provisionally to McCoy's genus
Palasterina, under the subgeneric name Schcenaster. Later com-
parisons, however, have satisfied us that it can not properly be retained
in that genus; and as it seems to present equally important differ-
ences from all the other established genera known to us, we now
propose to separate it as a distinct genus, under the name Schoenaster.
It is probably most nearly allied to the Silurian genus, Palasterina,
but differs in the peculiar oblique, outward, imbricating character
of its row of plates on each side of the ambulacral furrows, and the
distinct inward imbricating character of the minute, scale-like plates
covering the under side of the disk. There are likewise differences
in the arrangements of the plates and pores of the dorsal side of the
free arms, as seen in our figure 7&, pi. 19.
"As the disk is not seen in the specimen from which figure 7d,
of the plate just alluded to, was drawn, it presents so different an
appearance from fig. 7a of the same plate, showing the upper side
of the fossil, that, looking at these figures alone, doubts might arise
whether or not they belong to the same type. An examination,
however, of the upper side of the same specimen from which 7d was
drawn, as well as of other fragments, shows them to be the same. A
fragment of the same specimen from which fig. Id was drawn also
shows portions of the under side of the disk.
"Another species, apparently of this genus, from the Burlington
limestone, with much narrower ambulacral furrows 1 , shows numerous
i "We should explain here, that the ambulacral furrow of the enlarged ray, represented by fig. 7c of
plate 19, is proportionally too broad, in consequence of the accidental flattening of the specimen, and in
part to the adambulacral pieces being represented proportionally too small. These furrows are more
nearly natural in fig. 7d, but even in that specimen they are proportionally wider than in undistorted
examples."
204 BULLETIN" 88, UNITED STATES NATIONAL MUSEUM.
little, short, flattened, spine-like appendages protruding from these
furrows, with an outward imbricating arrangement or inclination
towards the extremities of the rays. There may have been other
little spines over the outer surface, in addition to those fringing the
margins of the disk, though the specimens retain no traces of them."
Genoholotype. S . fimbriatus Meek and Worthen. Lower Carbonif-
erous (St. Louis) of Illinois.
The other three species referred to ScTmnaster probably do not be-
long in this genus. The specimens have not been studied.
SCHCENASTER FIMBRIATUS Meek and Worthen.
Plate 35, figs. 1-4.
Palseasterina (Schoenaster} fimbriata MEEK and WORTHEN, Proc. Acad. Nat. Sci.,
Philadelphia, vol. 12, 1860, p. 449.
Schcenaster fimbriatus MEEK and WORTHEN, Geol. Surv. Illinois, vol. 2, 1866,
p. 278, pi. 19, figs. 7a-7d.
Original description. "Body regularly pentagonal star-shaped,
with the rays produced into rather acutely pointed arms, which are
convex above, and about equal in length to the diameter of the disk.
Plates of the upper side of the arms and disk, convex, or even tumid ;
near the disk those of the rays hexagonal, heptagonal, or irregular
in form, alternating, and consisting of about five or six longitudinal
rows, with a few much smaller intermediate pieces. Farther out
the rays, they gradually pass into two mesial ranges of oblong, alter-
nating pieces, with their longer diameter parallel to that of the rays;
while on each side of these, minute irregular pieces fill the space
between them and the marginal pieces. Toward the extremities
of the rays, these little intermediate pieces diminish in size and at
last become obsolete, leaving only the two middle and outer, or adam-
bulacral rows. Ambulacral furrows, in apparently undistorted
specimens, deep and nearly or quite twice as wide as the row of
pieces on either side; adambulacral pieces rather thick and strong,
and liable to present considerable differences in their obliquity and
breadth of surface exposed, in consequence of the compression or
distortion of the specimen. Plates of the under side of the disk, very
much smaller than the adambulacral, closely crowded together,
and owing to their imbricating arrangement, presenting much the
appearance of the scales of a fish; immediately on each side of the
rays, they imbricate toward the latter, but near the middle of the
space between any two ambulacra, the imbrication is inward toward
the mouth, so that in tracing the rows parallel to their longer diam-
eter, across between the rays, they are found to describe a nearly
semicircular curve, with a slight angularity near the middle.
"Near the extremities of the rays, the dorsal pores are seen to
pass between the ends of the two mesial ranges of oblong pieces,
REVISION OF PALEOZOIC STELLEROIDEA. 205
but farther in toward the disk they are more irregularly dis-
tributed. Our enlarged figure 7&, pi. 19, represents these pores
and the dorsal plates, as seen in one of the rays, with the convex
outer portion of the plates ground away, in which condition the pores
probably appear larger than natural. In specimens with the tumid
portion of these plates unremoved, the pores are not readily seen,
and the whole dorsal side then seems to be made up of solid, close-
fitting pieces. Greatest diameter across between the extremities of
the opposite rays, about 2.37 inches; diameter of disk, 1 inch; breadth
of ambulacral furrows, about 0.10 inch; length of little flattened
marginal spines, near 0.08 inch.
" Locality and position. St. Clair County, Illinois; in the St. Louis
division of the Subcarboniferous series."
The specimen has not been studied.
SCHCENASTER (?) WACHSMUTHI Meek and Worthen.
Plate 33, fig. 6.
Schcenaster wachsmuthi MEEK and WORTHEN, Proc. Acad. Nat. Sci., Philadelphia,
vol. 18, 1866, p. 259; Geol. Surv. Illinois, vol. 3, 1868, p. 499, pi. 17, fig. 4.
Original description. "Body flattened or much depressed, with
a regular, distinctly pentagonal outline, the angles being produced
into five rather attenuated rays, which are a little convex above,
and apparently as much as two-thirds as long as the diameter of
the disk, if not more. Disk concave in outline on the margin be-
tween the rays, and imparting a slightly alate character to the
latter by extending a little along their margins; like the dorsal side
of the rays, composed above of numerous bmall, slightly convex
plates. Dorsal pores moderately distinct between the plates.
Plates of the under side of the disk about as large as those of the
dorsal side, but flattened, scalelike, crowded, and having the inward
imbricating character of the genus strongly marked. Ambulacra
(as seen in a compressed specimen) very narrow, their marginal
plates moderately large, oval-oblong, comparatively thin, and very
strongly imbricating outward. Between these, two rows of short,
flattened, spinelike scales are seen arising from the ambulacral
furrow, and all inclining outward or toward the extremities of the
rays. (Other characters unknown.)
"Diameter of disk, 1.32 inches; rays apparently extending as much
as 0.90 inch or more beyond the margins of the disk.
"This species will be readily distinguished from our 8. fimbriatus,
of the St. Louis limestone, the only other species of the genus known
to us, by its smaller and less convex plates on the dorsal side, as well
as by its much thinner, less oblique, and more strongly imbricating
row of plates along each side of the ambulacra, and particularly
by its much narrower ambulacral furrows. We have not seen any
206 BULLETIN
traces of the row of short, flattened, marginal spines seen around the
disk of S. fimbriatus, in the form under consideration; nor have
the similar little appendages arising in a double row from the am-
bulacra of the latter been seen in S. fimbriatus. These, however,
may be rather generic than specific characters, and consequently
be found common to both species.
"The specific name is given in honor of Mr. Charles Wachsmuth,
of Burlington, Iowa, the discoverer of the only specimen we have
seen."
Formation and locality. Burlington limestone, Burlington, Iowa.
The specimen has not been seen; it is in the Museum of Compara-
tive Zoology, Harvard University (No. 7).
SCHCENASTER (?), new species.
In the Gurley collection of the University of Chicago there is a
small specimen (No. 10993) from the Chester formation of Pope
County, Illinois. According to the older classification it would be
referred to Schwnaster, but until this genus is thoroughly revised
nothing of generic value can be done with these Carboniferous
asterids. The specimen under discussion seems to be most closely
related to S. (?) wachsmuthi.
Measurements: R = about 10 mm., r = about 3.5 mm.
SCHCENASTER (?) LEGRANDENSIS Miller and Gurley.
Plate 32, figs. 4-6.
Schcenaster legrandensis MILLER and GURLEY, Descr. new gen. and ep. Echinod.,
1890, p. 56, pi. 9, figs. 7-9. MILLER, Sixteenth Rep. Geol. Surv. Indiana,
1891, p. 371, pi. 9, figs. 7-9; N. Amer. Geol. Pal., App. 1, 1892, p. 682, fig. 1246.
Original description. "Body thin, regularly pentagonal, sides
concave, with long, nanow, gradually tapering convex arms. Plates
on the dorsal side of the disk in our specimen apparently anchylosed,
and spines, if any belonged to the margin, broken away. Ventral
side depressed and flat between the arm furrows, where it consists
of very small plates, and if they imbricate inward the overlap must
be very slight. Ambulacral furrows wide, deep; two rows of sub-
quadrangular, ambulacral plates form the bottom of each furrow,
on each side of which there is a row of oblong adambulacral plates,
having an obliquely inward imbricating arrangement, presenting
somewhat the appearance of a twisted cord; these are continued to
the ends of the arms with the same obliquely inward imbricating
arrangement. Five pairs of oral plates.
"This species is small in comparison with the two heretofore
described, has sides less convex, and narrower arms."
Formation and locality. Kinderhook limestone at Le Grand,
Iowa. The specimen has not been seen; it may be in the Gurley
collection of the University of Chicago. It is certainly not a form
of Schcenaster.
REVISION OF PALEOZOIC STELLEEOIDEA. 207
SCHCENASTER (?) MONTANUS Raymond.
Plate 33, fig. 5.
SdiMnaster f montanus RAYMOND, Ottawa Naturalist, vol. 26, 1912, p. 80, fig.
3 on p. 81.
Original description. "Animal small, about an inch in diameter.
Rays short, slender, extending about one-half their length beyond
the disk. Disk large, pentagonal, the margin slightly concave
between the rays. The five proximal plates of the adambulacral
series function as orals, while on the arms beyond the disk the adam-
bulacrals become marginals. The adambulacral plates are rather
small, oval, placed with the long axis diagonal to the axis of the arm.
On the most perfect arm there are 16 of these plates on each side of
the groove, not counting the proximal and distal plates.
"The ambulacral ossicles are small, arranged alternately. On
this specimen they are mostly displaced. The plates on the disk
between the rays are few and small. The marginals are small,
rounded, and do not appear to bear spines, but this appearance may
be due to poor preservation.
" Locality. This species is described from a single specimen col-
lected by the writer in the Madison limestone at Spring Canyon in
the Ruby Mountains, near Alder, Montana. The type is in the
Carnegie Museum, Pittsburgh, Pennsylvania."
PAL^EOSOLASTERID^E, new family.
Specialized multiradiate Cryptozonia with as many as 29 rays,
large disk, and actinal madreporite. Ambulacralia opposite or slightly
alternating.
Contains :
Palseosolaster Stiirtz.
Echinasterias Sttirtz.
Echinodiscaster Delage and Herouard.
Echinostella Stiirtz.
Medusaster Sttirtz.
How rays are introduced in multi-rayed asterids. For paleontologic
purposes it is not necessary to go deeply into this matter, 'but it is
desirable to know when, where, and how the supernumerary rays are
introduced in living forms so that we may be guided in our generic
studies.
Ritter and Crocker 1 have recently summed up what is known on
this subject, and to their paper the reader is referred for further
study.
In nearly all cases the multi-rayed forms have the rays of equal
development, and for this reason it is the general belief that
i Ritter and Crocker, Multiplication of rays and bilateral symmetry in the 20-rayed starfish, Pycnopodia
kelianthoides (Stimpson), Proc. Washington Acad. Sci., vo . 2, 1900, pp. 247-274.
50601 Bull. 8815 14
208 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
adults have the full complement of rays characteristic of a species.
In some forms there are individuals with smaller immature arms,
but it is not certain that these are newly introduced rays, or that
"this disparity in size is due to the regeneration of the halves of
automatically bisected animals 77 (p. 248). It is true that certain
species start with the full complement of rays and introduce none
hi later growth. Others certainly deviate from this rule, and a
species with as many as 37 rays has one small specimen with only
24 plus 2 young sprouting rays. It is therefore plain that "in
Heliaster rays are added until far in adult life, 77 and it is also probable
that "they are added in pairs 77 (p. 249). In Labidiaster as many as
six new rays may bud simultaneously at intervals around the entire
circumference of the disk.
An adult Pycnopodia has from 20 to 24 rays, and "this variation
does not depend upon the size of the specimen. * * * The
number is generally even, * * * but a few specimens with 21
have been found 77 (p. 250). The smallest specimen had six equal
arms and two much smaller budding rays, one larger than the other.
Between these two extremes all intervening stages occur. At least
the original five and probably six rays are of larval origin, all the
others being postlarval. When the sixth ray is introduced is un-
known; the rest as a rule appear in bilateral order that is, in pairs.
They are all developed "in a budding area adjacent to I and II 77
on the side toward the sixth ray until late in adult life. When the
arms are of unequal number it is due to the suppression of a ray.
In Pycnopodia, therefore, ray multiplication goes on through a con-
siderable part of adult life, but not the whole of it, and is variable
with the individuals. All of the rays, whether of larval or post-
larval origin, have the same structural characters.
Finally, it seems that the sixth ray in Pycnopodia has its origin
in, or at least "is in some intimate way related with the larval organ
of the embryo 77 (p. 268). Therefore multiradiation among asterids
has its origin in late larval life.
In Heliaster , Clarke states, 1 "the formation of new rays is funda-
mentally different from that in Pycnopodia." They do not develop
hi distinct generations, but appear entirely independently of each
other. "A considerable number may develop at approximately the
same time, often as many as six or seven and sometimes eight or
nine in H. polybrachius, but they show no definite relation to each
other. 77
The evidence seems "to show that they tend to arise in all four
quadrants of the circumference of the starfish about equally, but
successively rather than simultaneously. 77
i H. L. Clarke, Bull. Mus. Comp. Zo61., vol. 51, 1907, pp. 25-76.
KEVISION OF PALEOZOIC STELLEROIDEA. 209
In Heliaster the original number is five, and the new rays are not
as a rule disposed bilaterally (pp. 63-66).
Verrill (1914) states that Heliaster and allied living forms strikingly
resemble Helianfhaster of the Devonic, " and may indicate a con-
tinuous descent from these ancient forms'* (p. 13). He also says, "I
am inclined to believe that the increase in number of rays has been
due more to the advantage gained in holding their food securely, and
in opening bivalves, than for holding to the rocks, though both go
together' 7 (p. 16).
Genus PALvEOSOLASTER Sturtz.
Plate 34.
Palseosolaster STURTZ, Verb. nat. Ver. preuss. Rheinl., etc., vol. 56, 1900, pp.
226-229, pi. 2, figs. 1-3.
Helianthaster CLARKE (part), Bull. N.Y. State Mus., No. 121, 1908, p. 64, pi. 11.
Generic characters. Disk very large, with ' 27 to 29 rays in the
genotype, which project one-third their length beyond the disk.
Animal large, up to 190 mm. in greatest diameter. Disk and rays
without marginalia. Length of rays 95 mm., width of rays 7 mm.
Abactinal area decidedly and abundantly spinous and devoid of all
columns of plates.
Actinally the great disk is also decidedly spinous. Rays slender,
with wide ambulacra; ambulacral plates very narrow, seemingly
alternating, but more probably opposite, h- -shaped, with large podial
openings in two rows in each ambulacrum. The rays are bounded
by narrow columns of small adambulacral ossicles that bear spines
on their outer sides. Mouth circular and very large. The nature of
the stout oral armature can not be made out.
Madreporite actinal, placed near the mouth, interbrachial, very
large, striate, more or less oval, about 34 mm. in greatest width.
The general aspect is said to be much like the living Solaster affinis
Brandt.
GenoJiolotype. P. gregoryi Sturtz (citations as above). Lower
Devonic of Bundenbach, Germany. Another species occurs at this
locality and a third is found in the Upper Devonic of New York.
Remarks. For present purposes we may state that Palseosolaster
has from 25 to 29 rays and that the ambulacralia are arranged opposite
one another; the other Sturtz genera, Echinasterias, Ec7iinodiscaster }
and Echinostella, are so much like Paldeosolaster that they may be
disregarded.
Clarke recently erroneously referred another very well preserved
species of this genus to Helianthaster (H. roemeri Clarke), but it
plainly has no direct relationship with H. rhenanus. This is seen
in the greater disk, larger number of rays, decidedly actinal position
210 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
of the madreporite and the absence of interbrachial inframarginals.
It is clearly a Palseosolaster.
PAL^JOSOLASTER (?) GYALUM (Clarke).
Plate 34, fig. 1.
Helianthaster gyalum CLARKE, Bull. N. Y. State Mus., No. 121, 1908, p. 63, pis.
12, 13.
Original description. The New York specimens, i i Helianthaster
gyalum nov. are smaller than H. rhenanus. The arms are more
numerous and appear to be quite uniformly 24-25. Compared to
H. rhenanus they are relatively short, but very long compared with
the size of the disk, which is much suppressed, and on none of the
specimens, all showing the oral surface, is any distinct evidence of
it visible, so deeply do the arms cut into it and so closely do they
lie together. Notwithstanding this apparent retreat of the disk the
madreporiform plate is very large. This organ is preserved in but one
example, but here it overlaps two adjoining interbrachial angles and
the mouth parts pertaining thereto. Instead of being a flat or
concave elongate plate, as in H. rhenanus, it is highly convex and
circular; its surface markings less distinct and coarse in that species.
"The great oral aperture is margined by a series of pronounced
' jaws' or sharp projecting elevated angles the sides of which take
origin from the margins of adjoining arms. These oral projections
are slightly expanded at their tips into blunt points comparable to
but smaller than the 'Hocker' of H. rhenanus, but like them carry
small spines projecting inward. The solidity and strength of these
mouth parts is indicated by their prominence and elevation as shown
in figure. * * * The reentrant angle at the base of each arm is
narrow, long, and acute, much more extreme in these respects than
in H. rhenanus and very much more elevated."
Actinally the rays have two columns of plates of which the ambu-
lacrals are by far the most prominent, are opposite one another or
slightly alternate, and occupy the comparatively wide ambulacral
furrows. The adambulacrals are narrow columns and apparently as
many in number as the ambulacrals. Laterally each adambu-
lacral bears several spines.
Locality and formation. A slab with three individuals from the
Portage (Upper Devonic) at EaiTs quarry, Ithaca, New York, is now
in Cornell University.
Remarks. The writer has not seen these specimens, but the illus-
trations seem to indicate that the " pronounced jaws" are made up
of small ossicles of which there are at least five in each column.
Further, that these ossicles are more probably the continuation of
the narrow adambulacrals than of the ambulacral columns into the
great oral area, and that proximally to these lie the minute pairs of
oral armature pieces.
REVISION OF PALEOZOIC STELLEROIDEA. 211
The madreporite in P. (?) gyalum, as in the forms of Palseosolaster,
is very large, actinal in position, and placed even closer to the mouth
than in any other species. Not only this, but it lies completely cov-
ering one proximal ambulacral furrow and across parts of two adja-
cent rays. Its general position and size are in harmony with Palseo-
solaster and not at all with Helianihaster.
That P. (?) gyalum can not be referred to Helianihaster is therefore
seen in the different position of the madreporite, the greater number
of rays, the wider ambulacral furrow, and the improbability of its
having three columns of abactinal ray plates in place of an integument
bristling with spines. All of these differences are in harmony with
Palseosolaster. Further, if P. (?) gyalum had interbrachial inframar-
ginals as does Helianihaster, they should show somewhere on these
five specimens, all of which preserve the actinal side. While these
differences may not appear to be great, they make of Helianihaster a
phanerozonian and of Palseosolaster a cryptozonian.
PAL^EOSOLASTER ROEMERI (Clarke).
Heliarthaster roemeri CLARKE, Bull. N. Y. State Mus., No. 121, 1908, p. 64, pi. 11.
From the Lower Devonic of Bundenbach, Germany. The holo-
type is in the New York State Museum.
Genus ECHINASTERIAS Sturtz.
Echinasterias STURTZ, Verh. nat. Ver. preuss. Rheinl., etc., vol. 56, 1900, p. 230,
pi. 2, fig. 4; pi. 3, fig. 5.
The genotype has 25 rays, but otherwise appears to have all the
generic characters of Palseosolaster.
Genoholotype and only species. E. spinosus Sturtz (citations as
above) . Lower Devonic of Bundenbach, Germany.
Genus ECHINODISCASTER Delage and Herouard.
Echinodiscus STURTZ (not Echinodiscus Worthen and Miller 1883, Echinoidea),
Verh. nat. Ver. preuss. Rheinl., etc., vol. 56, 1900, pp. 231-233, pi. 3, figs.
6-10.
EcMnodiscaster DELAGE and HEROUARD, Trait6 de Zoologie Concrete, 1904.
Echinodisdtes SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, 1914, p. 15.
One of the specimens has 29 rays and a greatest diameter of 160
mm. The writer fails to see how this genus is to be distinguished
from Palseosolaster. It may be more abundantly spinose, and the
madreporite is somewhat smaller and more coarsely striate, but these
characters can hardly be of generic value.
Genoholotype and only species. Echinodiscus multidactylus Sturtz
(citations as above). Lower Devonic of Bundenbach, Germany.
Cat. No. 59386, U.S.N.M.
212 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genus ECHINOSTELLA Sturtz.
Echinostella STURTZ, Verb. nat. Ver. preuss. Rheinl., etc., vol. 56, 1900, pp.
233-235, pi. 3, fig. 11; pi. 4, fig. 12.
The only specimen has 28 or 29 rays and a greatest diameter of
150 mm. Abactinally the rays and the central area of the disk
stand out rounded above the remainder of this side of the animal,
which is not the case in Palseosolaster; otherwise the two genera aie
very much alike.
GenoJiolotype and only species. E. traquairi Sturtz (citations as
above). Lower Devonic of Bundenbach, Germany.
Genus MED US ASTER Sturtz.
Medusaster STURTZ, Palseontographica, vol. 36, 1890, p. 229; pi. 31, figs. 34,35;
Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 54, 73.
A form with from 12 to 15 rays. Thought to have relationship
with living Solaster.
GenoJiolotype and only species. M. rhenanus Sturtz (citations as
above). Lower Devonic, Bundenbach, Germany.
Cat. No. 59385, U.S.N.M.
PALEOZOIC OPHIURIDS.
A great deal has been written about these animals, but in general
the true structure of some of them was not known until very recently.
Authors have described these delicate and generally poorly pre-
served fossils in such broad terms that a complete restudy of all of
the material is required before it can be said that most of the genera
or even the species are established. The way has been successfully
blazed by Schondorf in Europe, and his methods of study and results
need to be applied to the American fossils. It was the writer's hope
to do this, but it is now plain that his present duties will prevent his
doing so.
In general it may be said that but few Paleozoic so-called ophiurids
are true Ophiuroidea. They are ophiurid-like animals, with open
ambulacral furrows, but without the essential characters of the
Asteroidea. Nor are they transitional in structure between the
Asteroidea and Ophiuroidea. Therefore Schondorf has separated
them under the term Auluroidea. It seems to the writer that they
originated in the Asteroidea early in the Ordovicic in some crypto-
zonian stock near Urasterella, with large and square ambulacralia.
(Stenaster and Tetraster are probably true aulurids.) Out of the Aulu-
roidea, probably in late Devonic time, arose the true Ophiuroidea.
The essential differences between the Auluroidea and Ophiuroidea
will become plainer to the student after a study of the text and
figures presented in this work.
REVISION OF PALEOZOIC STELLEROIDEA.
213
Subclass AULUROIDEA.
Auluroidea SCHONDORF, Palaeontographica, vol. 67, 1910, p. 60; Jahrb. nassauisch.
Ver. Naturk., Wiesbaden, vol. 63, 1910, pp. 234, 247, 251. SPENCER, Mon.
Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, pp. 24,48. '
Protophiuroidea and Euophiuroidea SOLLAS and SOLLAS, Philos. Trans. Roy. Soc.
London, ser. B, vol. 202, 1912, pp. 214,222.
Auluroidea are Paleozoic brittle-stars, with a more or less well-
developed central disk, whose border between the rays is more often
concave than convex ; in other words,
star-shaped. The rays appear al-
ways to be five in number, simple,
long and slender, and more or less
sharply marked off from the disk as
appendages. This means that the
body cavity does not extend from
the disk into the rays as in Aster-
oidea. On the actinal side the rays
have open ambulacral furrows with
the ambulacralia never anchylosed
into vertebrae ; they are arranged in
two columns, the pieces of which
are either alternate or opposite one
another.
Original definition. "The ambu-
lacral water-vascular system lies in
a circular canal that is situated wholly within the ambulacralia medi-
ally between the adjoining columns of ossicles. From it diverge short
side branches that either partially penetrate the substance of the
individual ambulacrals, or lie in the suture
between adjoining ambulacrals; in either
case they extend into the ventrally open,
broad ambulacral furrow. Ambulacrals free,
not coossified, those of adjoining columns
either alternate or opposite, but always di-
rectly opposite the adambulacrals. Ambu-
lacrals ventrally composed of a medially
situated, internally concavely hollo wed-out
body, which, as the plates of the two columns
are normally united, forms the medial ambu-
lacral canal; and of a lateral process extend-
ing to the adambulacrals, with depressions
on both sides of it in the ventral surfaces of
the ambulacrals and adambulacrals [see fig.
14]. [These large lateral depressions are not podial openings through
or between the plates as in asterids, but appear to be cavities for the
ampullae or for these and podial insertions. Dorsally the ambulacrals
FlG. 12. A SECTION THROUGH THE ABM OF AN
ASTEKID, AFTEK SCHONDORF. A , AMBULACRA-
LIA; Ad, AD AMBULACRALIA; Amp, AMPULLA;
D, DORSAL SKELETON; F, PODIA; ipl, ACCES-
SORY OSSICLES; Mo, ATTACHMENT FOR UPPER
LONGITUDINAL MUSCLES; Mo 1 , UPPER TRANS-
VERSE MUSCLE ; Mu, ATTACHMENT FOR LOWER
LONGITUDINAL MUSCLE; Mv>, LOWER TRANS-
VERSE MUSCLE; Mv, VERTICAL MUSCLE ; Bo,
UPPER MARGINALIA; Ru, LOWER MARGINALIA;
Wr, RADIAL WATER- VESSEL.
FlG. 13. A SECTION THROUGH THE
ARM OF A TRUE OPHIURID, AFTER
SCHONDORF. Bs, VENTRAL SHIELD;
C^CUTIS; F, PODIA; L, BODY CAV-
ITY; N, NERVE-RING; oZm, uZm,
UPPER AND LOWER INTERMEDIARY
VERTEBRAL MUSCLES; Rs, DORSAL
SHIELD; Ss, LATERAL SHIELD; W,
VERTEBRA; Wr, RADIAL WATER-
VESSEL.
214
BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Ad.
FIG. 14. A SECTION THBOUGH THE ARM OF AN AULUBID,
WITH ALTERNATING AMBULACEALIA, AFTEE SCHON-
DOEF. A, AMBULACEALIA OF THE DOESAL AND VEN-
TRAL SKELETON; A 1 , VENTEAL EXTENSION OF THE
AMBULACEALIA; Ad, AD AMBULACEALIA = LATERAL
SHIELDS OF OPHIUEIDS; F, PODIA, ON LEFT THE CANAL
PASSES THBOUGH THE PLATE, ON EIGHT THE PLATE IS
DISSECTED DOWN TO THE CANAL; Wr, INTEESKELETAL
EADIAL WATEE- VESSEL.
are covered with spines and tubercles.] Disk with concave, or con-
vex ( ?) margins, with or without marginal ossicles. When the latter
are present, they never extend along the rays, but are wholly re-
stricted to the disk. The rays on both sides are margined by the
adambulacrals. Ventrally the rays have broad open ambulacral fur-
rows, bounded laterally by the
adambulacrals. A typical mad-
reporite [probably always] lies
in one of the ventral interradii.
"The group [subclass] is re-
stricted to the older Paleozoic."
Remarks. This subclass of
Stelleroidea appears to be an in-
dependent development whose
structure partakes of that of
the Asteroidea and the Ophiu-
roidea, though more like the
latter. The Auluroidea agree
with the asterids in having open
ambulacral furrows, and a true madreporite, which is, however, ven-
trally situated, but otherwise the class is more like the ophiurids, since
the body cavity does not extend into the rays as in true starfishes.
However, the water-vascular canal in the Auluroidea does not lie
outside of the ambulacrals as in the Asteroidea, but within these
ossicles along the mid-line of the ambulacrum in a canal that is cut
out of the sides of adjoining ambulacral columns. On the other
hand, the Ophiu-
roidea have the am- AA A A i* A4 A A Ad
bulacralia coossified
and modified into
vertebral ossicles;
there are no open
ambulacral furrows,
and the entire rays
are covered by four
columns of shields or by an integument more or less studded with
calcareous plates and granules.
It appears that the Auluroidea all have dorsally four columns of
plates, of which the two medial ones are the most prominent. These
are undoubtedly the dorsal aspect of the thickened ambulacrals.
Outside of the disk they are usually convex and more or less orna-
mented with granules, but within the disk are less prominent and take
on other expressions, commonly with concave surfaces. The plates
outside of these margin the rays and are the adambulacrals; while
in some forms they are prominent and tuberculate, usually they are
made up of narrow vertical pieces that bear spines along the ventral
or only the distal ventral edge.
FIGS. 15 AND 16. VENTEAL AND DOESAL AEM STBUCTUEE OF AULUBOIDEA,
WITH THE AMBULACEA ALTEENATING. SCHEMATIC, AFTEE SCHSNDOEF.
A, AMBULACEALIA; A 1 , VENTEAL EXTENSIONS OF AMBULACEALIA; A 3,
AD AMBULACEALIA; FS, PODIAL CAVITIES.
KEVISION OF PALEOZOIC STELLEKOIDEA. 215
The Auluroidea can be divided into two orders on the basis of
whether the ambulacrals on each side of the median line are arranged
alternately (the Lysophiurae), or are opposite to one another (the
Streptophiurae). The following classification is a modification of
the one offered by Gregory in 1897, and kis work was in turn a
modification of that of Bell in 1892.
After the present work was written, its author first became aware
of the works of Sollas and Sollas, and Spencer. The reader should
consult those papers for important studies on the Auluroidea.
Order LYSOPHIUR^E: Gregory.
Encrinasterise BRONN (part), Klassen u. Ordnungen d. Tierreichs, vol. 1, 1859,
p. 287. SCHONDORF, Palaeontographica, vol. 57, 1910, p. 61.
Ophio-Encrinasterise STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 56,
1900, p. 198. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 63, 1910, pp. 234, 237.
Lysophiurse GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1030 (contains
Protasteridse and Palaeophiuridae) ; Treat. Zool., vol. 3, Echinoderma, 1900,
p. 274. SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London, ser. B, vol.
202, 1912, p. 223.
Ophiurasterix SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 251.
Auluroidea in which the ambulacral ossicles of adjoining columns
are alternate to one another. The individual plates are separate and
not coossified into vertebrae.
Remarks. The Lysophiurae are Paleozoic or primitive Auluroidea,
that is, ophiurid-like animals, with wide and open ambulacral furrows
in which the ambulacralia are free and alternately arranged. The
ambulacralia are common to the actinal and abactinal sides.
This order contains the following families:
Protasteridse. Have boot-shaped ambulacralia ventrally.
Palseophiuridae. Have subquadrate ambulacralia ventrally.
Encrinasteridae. Have boot-shaped ambulacralia ventrally
and large disks that are bordered by marginalia.
Family PROTASTERIDSE Miller.
Protasteridse MILLER, N. Amer. Geol. Pal., 1889, p. 216. GREGORY, Proc. Zool.
Soc. London for 1896, 1897, p. 1031; Treat. Zool., vol. 3, Echinoderma, 1900,
p. 274. PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 366.
Paldeophiomyxidse STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 202.
"Lysophiurae with boot-shaped ambulacral ossicles, each composed
of a 'body ' in the median line of the arm, and a lateral 'wing' at right
angles to it" (Gregory). There is either a well-marked scale-covered
or granular disk, and five flexible arms. The adambulacral plates are
rather stout, with many lateral spines.
216 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Schondorf states that Gregory's family Palseopliiuridae has no
value and is to be merged into the one under discussion. The reasons
for this are given beyond under the former family heading.
The family contains:
Tseniaster Billings. Alepidaster Meek.
EopTiiura Jaekel. Gregoriura Chapman.
Bohemura Jaekel. Bundenbachia Stiirtz.
Palseura Jaekel. PalseopJiiomyxa Stiirtz.
Protaster Forbes.
Genus T^ENIASTER Billings.
Plate 36, figs. 1-3.
Tseniaster BILLINGS, Geol. Surv. Canada, Can. Org. Rem., dec. 3, 1858, p. 80.
CHAPMAN, Canadian Journ., n. ser., vol. 6, 1861, p. 517. WRIGHT, Mon.
British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Paleeontogr. Soc. for 1861), 1862,
pp. 24, 34. HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 300;
rev. ed., 1868=1870, p. 338. ZITTEL, Handb. Pal., vol. 1, 1879, p. 445.
STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 150; Palseontographica, vol.
32, 1886, pp. 78, 83. MILLER, N. Amer. Geol. Pal., 1889, p. 285. GREGORY,
Geol. Mag., dec. 3, vol. 6, 1889, p. 26. STURTZ, Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, p. 20. J. F. JAMES, Journ. Cincinnati Soc. Nat.
Hist., vol. 18, 1896, p. 138. GREGORY, Proc. Zool. Soc. London for 1896, 1897,
p. 1035; Treat. Zool., vol. 3, Echinoderma, 1900, p. 250. SPENCER, Mon.
Brit. Pal. Asterozoa, pt. 1 (Paleeontogr. Soc. for 1913), 1914, pp. 8, 19.
Taeniura GREGORY, (not Tseniura Miiller and Heule 1837, sting-ray), Proc. Zool.
Soc. London for 1896, 1897, p. 1035.
Original description of Tseniaster. " Generic Characters. Body
deeply stellate; no disk [there is a disk] or marginal plates; rays
long, slender, flexible, and covered with small spines; two rows of
large ambulacral pores; adambulacral plates elongated and sloping
outward so that they partly overlap each other; ambulacral ossicles
contracted in the middle, dilated at each end. Generic name from
tainia, a riband."
Genolectotype (the first species of Billings is here selected as the
genolectotype) . Palxocoma spinosa Billings.
Remarks. Hall restudied the type-material of T&niaster and
records his observations as follows :
"In reviewing the characters of Protaster and Eugaster, I became
satisfied that there was an intimate relation between these and
Tseniaster of Billings, and in order to satisfy myself on this point,
I have * * * received from Sir William E. Logan, permission to
examine specimens of T&niaster spinosus and T. cylindricus.
"An examination of the specimen illustrated in fig. 3 (ut sup.)
reveals what I conceive to be a disk not at all unlike the disk of Prot-
aster, but less extended than in the Lower Helderberg species. The
structure of the ray is precisely of the same character as the ray of
REVISION OF PALEOZOIC STELLEKOIDEA. 217
that species which I have named Protaster forbesi, the proportions of
plates and relations of parts showing specific differences/'
"Mr. Billings remarks, under the description of T. spinosus, that
' the ambulacral ossicles appear in some places to alternate with each
other, but this is owing to a distortion; those on one side of the furrow
are opposite to those upon the other. 7
"Now the specimen of this species which I have examined, and
which I suppose to be the one figured upon Plate X, figures 3d, Z>,
has the ambulacral plates alternating * * *.
"In the specimen of T. cylindricus examined the ambulacral
plates are less distinctly alternate, but the relation of the adam-
bulacral plates and the pores are the same as in the other form.
Not having seen the specimen showing the dorsal view, figure 4o- of
plate X, I can only remark that the structure of the rays is very
similar to that of Protaster."
Without restudying the type-material of Tseniaster, or paying
attention to Hall's statements that T. spinosus has what he con-
ceived "to be a disk not at all unlike the disk of Protaster," Gregory
(1897) regarded Tseniaster as a Cryptozonia starfish and made of
Billings's second species, T. cyliiidricus, a new genus, Tseniura. The
latter he correctly refers to the primitive ophiurids. Under these
circumstances it is desirable to repeat here what Gregory wrote
about Tseniaster.
Tseniaster "was described as diskless, which in respect to the type
species T. spinosus is correct. In that species the oral skeleton
consists of five pah's of large adambulacrals, as in ordinary Asterids.
The affinities of the true Tseniaster appear to me to be with such forms
as Palssaster ruthveni [= Urasterella ruthveni]. It is asteroid in the
oral armature, in its alternately arranged ambulacral ossicles, and in
the absence of a disk. I therefore consider Tseniaster a genus of
Asteroidea. The second species placed by Billings in this genus
has, however, a well-marked disk, and has the oral armature com-
posed of five pairs of Ophiuroid syngnaths. It must, therefore be
included among the Ophiuroidea."
To adjust this matter, the writer restudied the genotype of Tseniaster
at Ottawa (two specimens, No. 1404, the originals of figures 3a-3d
of Billings's Plate X). There can be no doubt that we have here a
genuine lysophiurid. There is a small disk present in the individual
figured as 3a and 3Z>, but it is not one with a round outline, as in
typical ophiurids, but is concave along the margin between the rays.
Actinally this disk is abundantly covered with prominent but slender
spines.
The ambulacrals are distinctly alternate throughout the rays and ap-
pear as in the sketch presented on page 218; this condition is not due
to distortion or other causes. The side plates are narrow, and each
218 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
bears three spines (see fig. 17B); toward the mouth the two columns
diverge far more than one would assume from the original figure, and
in consequence there are long and slender syngnaths and not the
asterid-like mouthpieces figured in the illustration. (See fig. 17C.)
3
FIG. 17. DIAGRAMS OF T^NIASTEB SPINOSUS (BILLINGS). A, To SHOW THE SPINOSE DISK WITH CONCAVE
SIDES. B, ACTINAL PLATE ARRANGEMENT: C, AMBULACRALS; ad, ADAMBULACRALS OR SIDE PLATES,
WITH THEIR SPINES. C, MOUTH PLATES: a, AMBULACRALS; S, SYNGNATH.
The specimen of figures 3c and 3d is too poorly preserved to add
anything further. Associated with these types there is another and
much larger specimen with a well-marked disk, of which Billings
said nothing in his original description. This form is clearly of
another species (probably T. cylindricus) , and is mentioned here so
that other workers will not confound it with the original material
of T. spinosus.
In 1900 Gregory referred Tseniaster to the asterid family Tsenias
teridse, and placed here as well the unrelated genera Stenaster, S alter-
aster, and Urasterella. From what has been stated above it is plain
that T&niaster can not be closely related to the cryptozonian genus
Urasterella , since the latter has an abundance of narrow ambulacralia,
asterid in type, while these ossicles in the former are far fewer in
number and of the form seen in other and related Lysophiurse. It
is very probable that Tseniaster -arose in the same stock that gave
rise to Urasterella and Stenaster, but the former genus is clearly one
of the aulurids.
In 1896 Gregory took out of T&niaster the second species of
Billings (T. cylindricus) and based on it the new genus Tseniura.
As will be seen, this name can have no standing, as the generic char-
acters are those of Tseniaster, and, further, the name is preoccupied
since 1837.
Thinking T. cylindricus most closely related to the primitive
ophiurids PalseopTiiura and St&rteura, Gregory defined T&niura as
follows :
"Diagnosis: Palseophiuridse with a small pentagonal disk, not
bounded by marginal ossicles. The ambulacral furrow is broad.
The oral skeleton is conspicuous and the syngnaths each composed
of two separate pieces. The two jaws of each oral angle are closely
attached; the mouth frames are separated, and each of them is a
short, thick, slightly bent bar."
REVISION OF PALEOZOIC STELLEEOIDEA. 219
Genotype and only species. Palxocoma cylindrica Billings.
11 Remarks: This genus differs from PalseopTiiura by the absence
of marginal ossicles from the disk, and from Sturtzura by the smaller
size of the disk and the form of the syngnaths. In Sturtzura the
jaws end bluntly against a jaw plate, whereas in Tseniura they appear
to taper to a point and have no jaw plate."
The two cotypes of T. cylindricus (1405a) were lost for many years,
but have recently been rediscovered in the collections at Ottawa.
In addition, there is another specimen in that Museum under this name,
and it is probable that it was so identified by Billings (No. 1405).
This was the only specimen studied, and its generic characters agree
very well with the originals of the cotypes (figs. 4b and 4c of pi. X),
and appear to be in harmony with the genotype of Tseniaster. The
only important difference is that here the ambulacralia are directly
opposite one another, and not slightly alternating, as in T. spinosus
(see fig. 18), a difference, it seems to the writer, not in itself alone
worthy of being the base of a new genus distinct from Tseniaster.
Tseniaster has the following species:
T. spinosu^s (Billings). Lower Trenton.
T. cylindricus (Billings). Trenton.
T. schofiarize Ruedemann. Upp^r Trenton.
T. elegans Miller. Richmondian (Waynesville).
T. meafoadensis Foerste. Richmondian.
TXENIASTER SPINOSUS (Billings).
Plate 36, fig. 1; text fig. 17.
Palseocoma spinosa BILLINGS, Geol. Surv. Canada, Rep. Progress for 1853-1856,
1857, p. 292.
Tseniaster spinosus BILLINGS, Geol. Surv. Canada, Can. Org. Hem., dec. 3, 1858,
p. 81, pi. 10, figs. 3a-3d. MILLER, N. Amer. Geol. Pal., 1889, p. 285, fig. 439.
Revised description ly BiUings. "The specimens collected are
about seven lines in diameter; the rays linear-lanceolate, one line
in width at the base, and covered at the sides with numerous small
spines.
"In the view of the enlarged specimen (pi. X, 3s), the ambulacral
ossicles appear in some places to alternate with each, other, but this
is owing to a distortion. Those on one side of the furrow are oppo-
site those upon the other. The adambulacral plates are elongated,
and so placed that the outer extremity of the one lies upon the inner
extremity to the next. The rays are flexible."
For remarks on this species, see the discussion under Tseniaster.
Formation and locality. Two specimens from the Lower Trenton,
Falls of Montmorency, east of Quebec, Canada. The original of
figures 3a to 3d is in the Victoria Memorial Museum, Ottawa
(No. 1404).
220 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
T^NIASTER CYLINDRICUS (Billings).
Plate 36, figs. 2, 3; text fig. 18.
Palseocoma cylindrica BILLINGS, Geol. Surv. Canada, Rep. Progress for 1853-
1856, 1857, p. 292.
Txniaster cylindricus BILLINGS, Geol. Surv. Canada, Can. Org. Rem., dec. 3,
1858, p. 81, pi. 10, figs. 4a-4c. WRIGHT. Mon. British Foss. Echinod., Oolitic,
vol. 2, pt. 1 (Palseontogr. Soc. for 1861), 1862, p. 34. PARKS, Trans. Cana-
dian Inst., vol. 8, 1908, p. 371.
Txniura cylindrica GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1035.
Lapworthura cylindrica PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 371.
Original description. "One inch and a half in diameter; rays
five [very flexible], covered with spines, subcylindrical, regularly
rounded on the upper side, flattened
on the lower, about one line in width
at base, and regularly tapering to an
acute point."
Formation and locality. In the Tren-
ton limestone at Ottawa, Canada. The
. i8.-DiAGRAMOp7cTmAL BAY-PLATES cotypes (14050), along with a third
OF TJSKIASTKR CYLINDKICUS BILLINGS, specimen (1405), are in the Victoria
a, AMBTJLACKAL; ad, SIDE PLATES. _* . , v ,
Memorial Museum at Ottawa.
T^ENIASTER SCHOHARL Ruedemann.
Tseniaster schoharise RUEDEMANN, Bull. N. Y. State Mus., No. 162, 1912, p. 88,
pl. 3, fig. 1.
Original description. The holotype "is small, the arms about 15
mm. long, and it is without a disk. The arms are slender and flexible,
about 1.3 mm. wide at the base, and almost as high in lateral view,
originally probably cylindrical. The ventral view shows a straight or
slightly zigzagged ambulacral canal, and on both sides of this squarish
depressions surrounded by the ambulacral and adambulacral ossicles,
the covering lower arm plates not being retained if they existed.
The inner (ambulacral or vertebral) ossicles appear as narrow and
outwardly curved ridges, the outer or adambulacral ossicles as ridges
bent in the opposite direction with a projection in the middle of the
outer arch. The ambulacral ossicles are not directly opposite nor
regularly alternating, but those of the right side on the ventral view
advanced about one-third the length of the ossicles beyond those of
the other side.
"The lateral view of one other arm exhibits the ossicles as ver-
tical bars, that are thickened at both ends resembling vertebrae
and terminating at the dorsal side with a flat surface and forming
there an apparently continuous layer. They appear almost twice
as wide as the intervening spaces. The dorsal surface bears bundles of
obliquely forward directed spines, one bundle corresponding to each
ossicle. These spines appear in the right arm to proceed from
REVISION OF PALEOZOIC STELLEROIDEA. 221
small spine-bearing plates attached to the adambulacrals, as in the
closely related Bundenbachia. On one arm two bundles are seen
to proceed from each segment, one oblique, the other vertical to
the arm. The oral skeleton consists of stout, bifid pieces, much
resembling the oral pairs of adambulacrals of the genotype."
Formation and locality. Upper Schenectady formation, the
eastern shale phase of the Upper Trenton limestones. From near
Schoharie Junction, New York. The holotype is in the New York
State collection at Albany.
T^ENIASTER ELEGANS Miller.
Tseniaster elegans MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 5, 1882, p. 41,
pi. 1, figs. 6-6c. J. F. JAMES, Journ. Cincinnati Soc. Nat. Hist., vol. 18,
1896, p. 139. PARKS, Trans. Canadian Inst., vol. 8, 1908, pp. 365, 371.
Protaster elegans PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 368.
Original description. "This species is founded upon more than 30
specimens occurring on a single slab, but showing only the ventral
side, with the exception of the ends of some of the rays.
"The body is deeply stellate, and the rays long, slender, and
flexible, and margined on either side with a row of spines.
"From the length of some of the rays, a complete specimen,
properly spread, would probably have a diameter of an inch and a
quarter, or more. The rays taper to a sharp point, are rounded on
the dorsal side, and margined on the ventral side by a row of spines,
one spine arising from each plate. Only two rows of plates seem to
form the dorsal side of a ray. The ventral side of each ray is marked
by a furrow in the middle, separating two series of plates or ossicles.
These plates are a little longer in the direction of the ray than wide,
and alternately break joints at the middle, where they are slightly
contracted."
Formation and locality. This extraordinary slab is in the Harris
collection of the United States National Museum. It was found near
Waynesville, Ohio, in the Richmond formation of the Ordovicic
period.
Cat. No. 40878, U.S.N.M.
TffiNIASTER MEAFORDENSIS Foerste.
Txniaster meafordensis FOERSTE, Bull. Sci. Lab. Denison Univ., vol. 17, 1914, p.
326, pi. 4, figs. 5-7.
The reference to this species is inserted as this book is going through
the press. The form appears to be common in the lowest Richmon-
dian on Workman Creek, three miles southeast of Meaford, Ontario,
Canada. The types are probably in the Victoria Memorial Museum,
Ottawa, Canada.
222
BULLETIN 88, UNITED STATES NATIONAL MUSEUM,
Genus EOPHIURA Jaekel.
Text figs. 19, 20.
EopUura JAEKEL, Zeits. geol. Gesell., vol. 55, 1903, Protokol, p. 14 (107), figs. 1, 4.
GenoJwlotype. EopMura, species not named. To give this genus
proper standing according to the rules of nomenclature, the form is
here named E. bohemica, new species. It is the only species of the
genus, and occurs in the Ordovicic (Dj) at Osek, Bohemia.
The genus is not defined in the regular manner, but from the general
discussion and the illustrations the following has been made out:
Kays outside of the disk,
apparently petaloid i n
form, with wide-open am-
bulacral furrows, having
narrow alternating ambu-
lacralia that are \ shaped,
with the podial openings
excavated on each side of
the lateral extensions.
The adambulacrals are
very large, drawn out lat-
erally, rectangular, lie hori-
zontally, and in no way
resemble the side plates of
typical ophiurids, or, for
that matter, even the Lys-
ophiurse. The lateral
edges of these ossicles are
slightly elevated, tubercu-
lated, and each bears about
four long and very slender
spines. Inside of the disk
the adambulacrals are nar-
row and stand vertically
as in Paleozoic ophiurids ;
the columns first diverge
The two proximal pieces of
mouth extensions, with the
FIG. 19. END OF A KAY OF EOPHIUKA BOHEMICA, MUCH EN-
LARGED, AFTER JAEKEL. a, AMBULACRALIA; ad, ADAMBULA-
CRALIA; Afg, PIT OF PODIA; ads, LATERAL EXTENSION OF
AD AMBULACRALIA; Rr, RADIAL FURROW; St, SPINE BASES.
and then converge in the mouth area.
each column make the adambulacral
terminal ossicles of adjacent columns side by side but not coalesced,
forming the syngnaths. The two most proximal ambulacrals rapidly
decrease in size, enlarging the mouth area, with none of these ossicles
on each side of the two final adambulacrals.
The disk seems to be small, with concave interbrachial sides.
Actinally these areas are covered with scale ossicles. Madreporite
ventral, in the outer and lateral part of an interradius. Immediately
outside of the syngnaths occur interradially two small ossicles,
reminding one of vestigial inframarginalia.
REVISION OF PALEOZOIC STELLEROIDEA.
Genus BOHEMURA Jaekel.
Text fig* 21.
223
Bohemura JAEKEL, Zeits. geol. Gesell., vol. 55, 1903, Protokol, p. 18 (111), fig. 6.
This genus, like the foregoing, is not defined in the regulation way.
Jaekel's fine drawing shows that it is closely related to Eophiura,
differing mainly in that the rays do not end bluntly, but have slowly
tapering whip-like terminations. These ends are of course ontoge-
netically young and reveal the progressive ray development. Here
FIG. 20. ORAL SKELETON OB- EOPHIUEA BOHEMICA, AFTER JAEKEL. a, AMBULACRALIA; ad, ADAMBU-
LACRALIA; Ma, MADREPORITE AMONG THE SCALE PLATES.
the rays are made up of overlapping and alternating adambulacrals
that distally have no ambulacral grooves. Gradually an ambulacral
groove is developed and it becomes rapidly wider as one examines
the rays more and more proximally, and the adambulacrals turn
over and flatten out, finally making the great open ambulacral
furrows. The illustration here reproduced, figure 21 on page 224,
will show the generic characters as drawn by Jaekel.
Oenoholotype and only species. B. jahni Jaekel. Ordovicic (DJ
at Zahorzan, Bohemia.
Genus PALvEURA Jaekel.
Text fig. 22.
Palseura JAEKEL, Zeits. geol. Gesell., vol. 55, 1903,Protokol,p.l6(109)-17(110),fig. 5.
Oenoholotype. Palseura, species not named. No species is again
mentioned, nor is it stated that the generic name is new. To give
the name standing, the form figured is here named P. neglecta, new
genus, new species. It occurs in the Ordovicic (D t ) of Bohemia;
the exact locality is not given, but may be Osek.
50601 Bull. 88 15 15
224 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Jaekel here again fails to diagnose his genus, but some of the
characters can be clearly made out from his excellent drawing.
The disk is small and tends to have convex interbrachial areas. So
far as the oral disk area is concerned, the general structure is that
of Eophiura, with this marked difference the opposite and asterid-
like ambulacralia of the rays do not vanish proximally as in that
genus but crowd over and finally
rest wholly upon the adambulacrals,
and end against the distal edges of
the syngnaths . These are made up
of the coossified terminal adambu-
lacral ossicles.
Genus PROTASTER Forbes.
Text fig. 23.
Protester FORBES, Mem. Geol. Surv.
United Kingdom, dec. 1, 1849, pi.
4, pp. 1-2; in McCor, British Pal.
Foss., 1851, p. 60. MURCHISON,
Siluria, 1854, p. 221, fig. 39-4.
SALTER, Rep. Twenty-sixth Meet.,
Brit. Assoc. Adv. Sci., Notices and
abstracts, 1857, p. 76; Ann. Mag.
Nat. Hist., ser. 2, vol. 20, 1857, pp.
325, 330 HALL, Nat. Hist. N. Y.,
Pal., vol. 3, 1859, p. 134. SALTER,
Ann. Mag. Nat. Hist., ser. 3, vol. 8,
1861, p. 484. WRIGHT, Mon. Brit-
ish Foss. Echinod., Oolitic, vol. 2,
pt. 1 (Palseontogr. Soc. for 1861),
1862, pp. 23, 31. HALL, Twentieth
Rep. N. Y. State Cab. Nat. Hist.,
1868, p. 293; rev. ed., 1868=1870,
p. 335. QUENSTEDT, Petrefacten-
kunde Deutschlands, vol. 4, 1876,
p. 133, pi. 95, fig. 11. STURTZ, N.
Jahrb. fiir Min., etc., 1886, vol. 2,
pp. 148, 150. GREGORY, Geol.
Mag., dec. 3, vol. 6, 1889, p. 26.
ST^RTZ, Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, p. 18.
J. F. JAMES, Journ. Cincinnati Soc.
Nat. Hist., vol. 18, 1896, p. 137. GREGORY, Proc. Zool. Soc. London for 1896,
1897, p. 1031, fig. 1. WHIDBORNE, Palseontogr. Soc., 1898, p. 206. PARKS,
Trans. Canadian Inst., vol. 8, 1908, pp. 363, 366. SCHONDORF, Jahrb. nas-
sauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 238. SOLLAS and SOLLAS,
PMlos. Trans. Roy. Soc. London, ser. B, vol. 202, 1912, p. 223. SPENCER,
Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, p. 35.
Encrinaster (part) HAECKEL, Gen. Morphologie, vol. 2, 1866, p. Ixvii (no definition).
Also Bee Sturtzura.
FiG.21. VENTRAL VIEW OF A RAY OF BOHEMURA
JAHNI, ENLARGED ABOUT X 2, AFTER JAEKEL.
a, AMBULACRALIA; ad, AD AMBULACRALIA WITH
THEIR LATERAL EXTENSIONS; ddm, SYNGNATHS.
REVISION OF PALEOZOIC STELLEBOIDEA.
225
Original description. " (Order Ophiuridse. Family Euryales.)
Body circular, covered with squamiform plates; genital openings
in the angles of junction of the arms beneath; arms (simple) formed
of alternating ossicula."
Genoholotype. " Species Unica. Protaster sedgwickii. Forbes."
Original description of P. sedgwickii. "The disk is circular, and
shaped like that of an OpTiiura. The arms are five in number, very
narrow, equidistant, and similar.
"The upper and under surfaces of the disk were covered by small,
similar, more or less regular, polygonal or crescentic plates, imbri-
cated in scale-fashion, and having punctated surfaces. Those of
the under side of the
body are smaller and
more regular than
those of the upper.
The mouth is cen-
tral, and rather small
in proportion to the
disk. The buccal ap-
paratus is composed
of ten parts or proc-
esses, arranged in
pairs; half of each
springs from the
origin of each arm in
a diverging manner,
and meets the corre-
sponding half to form
a lanceolate tooth-
1 i lr <* rkrrn'antirkn FlG> 22. ORAL SKELETON OF PAI^EUBA NEGLECTA, MUCH ENLARGED,
u j t? i o II, AFTER JAEKEL> at AMB ULACBALIA; ad, ADAMBULACEALIA.
deeply indenting the
cavity of the mouth. Of how many separate ossicles each of the
buccal processes was composed the specimen affords but very indis-
tinct indication. They seem to affect a slightly falcate form at their
extremities. The arms were composed of alternating, somewhat
quadrate ossicula, the sides of which were deeply indented supe-
riorly, in order to form spiniferous crests. The spines were appar-
ently short, and not equal in length to the length of an ossicle, obtuse,
and few in a row. The under surfaces of the brachial ossicula were
not indented laterally. The central portions of the upper surfaces
of the brachial ossicles were hollowed out slightly, and the sutures
between them deeply impressed. About 12 of the brachial ossicles
were imbedded in the disk, and the parts of the dorsal surface of
the latter, corresponding to the arms and central skeleton, do not
present traces of scales, a feature seen in existing Euryales.
226 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
"The dimensions of Prof. Sedgwick's specimens are as follows:
"Breadth of disk, 8/12ths inch.
"Average breadth between arms, 5/12ths inch. Breadth of
an arm, at its junction with the body, 1/1 2th inch.
"Locality and, geological horizon. Silurian, from Ludlow rocks
at Docker Park, near Kendal, Westmoreland (specimen figured),
associated with Encrinites, and at Benson Knot, also near Kendal,
hi hard sandstones, full of characteristic Ludlow fossils."
Gregory restudied this genus and in 1897 defined it as follows:
"Protasteridse with a well-marked disk; long, tapering, very flexible
arms. Some of the adambulacral ossicles are Y-shaped. Scales of
the disk fairly large. Type species. Protaster sedgwickii, Forbes."
Gregory remarks that "Forbes's figures of the arm-structures are
not satisfactory,"
figure 23 as drawn
by the former.
This illustration
a b shows each set of
FIG. 23. AMBULACHALIA AND ADAMBULACRALIA OF PROTASTEB SEDGWICKII,
AFTER GREGORY, a, NEAR THE DISTAL END; 6, IN THE MIDDLE; c, AT ambulacralia dif-
THE PROXIMAL END. THESE DIFFERENCES ARE DUE TO PRESERVATION.
tion thought to be due to preservation and adhering rock rather than
to structure (see fig. 28, on p. 242).
Madreporite probably abactinal in position.
In regard to the term Encrinaster, see Encrinaster (Aspidosoma) ,
page 242.
Protaster has the following species :
P. sedgwickii Forbes. Ludlow of England.
P. biforis Gregory. Wenlock of Wales.
P. groomi Sollas and Sollas. Ordovicic of England.
' P. (?) whiteavesianus Parks. Black River.
P. (t) salteri (Salter). Ordovicic of Wales.
. P. (?) stellifer Ringueberg. Rochester.
PROTASTER BIFORIS Gregory.
Text figs. 24, 25.
Protaster biforis GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1032, figs.
2, 3 on p. 1033. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 63, 1910, p. 238.
Original description. "Disk fairly large; interbrachial outlines
concave. The syngnaths are simple, prominent, and stout. The
ambulacral ossicles consist of a thick body and a stout curved wing.
The distal margin of the ossicles is notched by a depression for a
ventral muscle-field, which also cuts into the proximal margin of the
adjoining ossicle. Owing to these muscular depressions the arm has
apparently two series of pores.
REVISION OF PALEOZOIC STELLEROIDEA. 227
"The adambulacral ossicles are massive and taper slightly to their
distal ends; they are closely attached and form a regular series of
marginal plates.
"Arms very flexible.
"Dimensions:
Length of longest arm 18 mm.+a;
Diameter of arm at the base 2.5 mm.
Diameter of arm near the distal end 1 mm.
Diameter of mouth 1.5 mm.
Length of syngnath 1 mm.
Width of ambulacral furrow at edge of disk 1.25 mm.
"Distribution. Wenlock Shale, Castell Dinas, Bran, near Llan-
gollen," Wales. Museum Practical Geology, London.
if
24 25
FIGS. 24 AND 25. PROTASTER BIFOEIS. AFTEE GEEGOEY. 24, AMBULACRALIA AND ADAMBULACEAUA.
25, O, OUTLINE OF THE DISK AND AEMS; 6, A PAIE OF SYNGNATHS.
PROTASTER GROOMI Sollas and Sollas.
Protaster groomi SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London, ser. B
vol. 202, 1912, pp. 223, 224.
Occurs in the Middle Ordovicic of Shropshire, England.
PROTASTER (?) WHITEAVESIANUS Parks.
Protaster whiteavesianus PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 368,
figs. 1-6.
This is a large form with a, disk 15 mm. across and with rays 60 mm.
long. It is described in too much detail to quote here. The ambula-
cralia are distinctly alternate and boot-shaped.
Formation and locality. In the Lower Trenton (Kirkfield), at
Kirkfield, Ontario, Canada. The cotypes (five) are in the University
of Toronto (No. 638 T.).
PROTASTER (?) SALTERI (Salter).
Ophiura salteri SALTER and SOWERBY, Quart. Journ. Geol. Soc. London, vol. 1.
1845, pp. 9, 20 (table) (nomen nudum).
Protaster salteri WRIGHT, Mon. British foss. Echinod., Oolitic, vol. 2, pt. 1 (Pala>on-
togr. Soc. for 1861), 1862, p. 33 (nomen nudum). SALTER, Mem. Geol. Surv.
Great Britain, vol. 3, 1866, p. 289, pi. 23, fig. 3 (is inclined to refer it to
Tseniaster}. PARKS, Trans. Canadian Inst., vol. 8, 1908, pp. 364, 367.
Formation and locality. The holotype is from the Ordovicic, near
Cerrig-y-Druidion on the Holyhead road, Wales.
228 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
PROTASTER (?) STELLIFER Ringueberg.
Protaster stellifer RINGUEBERG, Bull. Buffalo Soc. Nat. Sci., vol. 5, 1886, p. 7, pi. 1,
fig. 2.
Generic relations not established.
Original description. "Disk of medium size, flat, circular, slightly
flexible, distinct from the rays on the dorsal side, and has on that
side a quinque-dentate, stellate, central elevation, which again has
a central, stellate depression of about one-half the lateral extension
of the stellate elevation upon which it is impressed. The points of
the star-like figure are opposite the several rays and extend about
two-thirds across the disk.
"Surface finely granulose. Rays slender, almost imperceptibly
tapering in the upper half of their known length; rounding on the
dorsal side, with two rows of regular, quadrilateral, alternating
[ambulacral] plates which can with difficulty be made out, and which
have a fine granulose [integumentary] surface resembling the surface
of the disk; opposite each transverse suture there is a corresponding
linear depression across the surface of the opposite plate on the other
side of the median suture, which at first sight gives the impression
that the ray is composed of opposite plates of only one-half the real
length.
"Ventral side with ten short, oral plates which are slightly sepa-
rated below, and meet at the discal surface, where they are rounded.
"Ambulacral series long, regularly quadrilateral; alternately
arranged.
"Adambulacrals indistinct in the only specimens found showing
the ventral surface.
"Marginal series slightly imbricating; spiniferous; spines rather
short."
Formation and locality. Three specimens were collected in the
Rochester shale of the Siluric at Lockport, New York. They are in
the author's collection. Two other fine specimens were found by C.
J. Sarle at Lockport and are now at Yale University. Another good
specimen is in the Walker collection of the University of Toronto
(No. 1007) ; it was found at Grimsby, Ontario.
Genus ALEPIDASTER Meek.
Plate 36, fig. 4; text fig. 26.
AlepidasterM-EEK, Amer. Journ. Sci., ser. 3, vol. 4, 1872, p. 275; Geol. Surv. Ohio,
Pal., vol. 1, 1873, p. 68, at end of specific description.
Protasterina ULRICH, Journ. Cincinnati Soc. Nat. Hist., vol. 1, 1878, p. 95. J. F.
JAMES, Journ. Cincinnati Soc. Nat. Hist., vol. 18, 1896, p. 139 (genoholotype,
P. fimbriata Ulrich).
Original description. "The only specimen I have seen that is
certainly known to belong to this species [P. ? granuliferus] is very
REVISION OF PALEOZOIC STELLEROIDEA.
imperfect, being merely an incomplete disk, and the inner ends of the
rays. It does not conform to the characters of Protaster given in
Prof. Forbes' s diagnosis, in all respects, since its disk, especially on
the upper side, is covered by an integument composed of a vast
number of very minute grains of calcareous matter, instead of dis-
tinct imbricating scales. It is therefore not improbable that perfect
specimens would show other characters that would warrant the
establishment of a new genus or subgenus for such forms, in which
case the name Alepidaster might be applied to the group."
Genoholotype. Protaster (?) granuliferus Meek.
Original description of
Protasterina. "Rays five,
slender, flexible, and ex-
tending much beyond a cir-
cular and minutely granu-
lar disk, which is provided
with short, slender, and
outwardly directed spines;
inner ray pieces [ambu-
lacra] regularly alternating,
of an hour-glass shape, and
interlocking along the me-
dian line, which is there-
fore not straight but zig-
zag ; outer ray pieces elon-
gated [adambulacrals or
side plates], directed ob-
liquely outwards, so as to
partly overlap each other;
two rows of large pores between the inner [ambulacral] and outer
[side plates] ray pieces; in the type-species these pores appear to have
been occupied by loosely-fitting subpyramidal plates, some of which
have a deep depression in the top, as though they were perforated;
their 1 true nature, however, is very uncertain. Oral pieces ten, each
pair being formed by two of the outer ray pieces. Type, P. fimbriata."
When the original material of Alepidaster and Protasterina is re-
studied, it may be shown that the latter is not a synonym of the
former, but at present the writer does not see any generic differences.
Alepidaster is easily distinguished from Protaster, not only by its
greater geologic age, but also in that its disk is minutely granular
and spiniferous while that of the latter is scale-bearing. Further,
when the exact shape of the ambulacrals is determined, it is probable
that other differences 'will be found there. Particular attention
should be paid to Ulrich's statement that in Protasterina the podial
openings " appear to have been occupied by loosely fitting, sub-
FIG. 26. DIAGRAMS OF ALEPIDASTER GRANULIFERTJS (MEEK).
DRAWN BY E. O. ULRICH FROM A SPECIMEN IN THE UNITED
STATES NATIONAL MUSEUM. A, ACTINAL VIEW OF RAY
PLATES. B, SAME FROM ABACTINAL SIDE. C, SECTION
THROUGH RAY. D, SlDE-PLATE FROM THE INSIDE AND IN
SECTION.
230 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
pyramidal plates/' If these extra plates existed, Protasterina will
be a good genus, differing not only from Protaster, but also from
Alepidaster.
Apparently no one has actually seen the abactinal side of the rays.
The writer also fails to see it in the few specimens studied. It would,
therefore, seem that there are no dorsal plates and that the dorsal
covering is integumentary or finely granular.
This genus contains the folio whig species :
A. (?), new species. Trenton.
A. granuliferus (Meek). Bichmondian.
A.flexuosus (Miller and Dyer). Eden and Maysville.
A. miamiensis (Miller). Bichmondian (Waynesville) .
ALEPIDASTER (?), new species.
In the Walcott collection at Harvard University there are three
specimens (Nos. 28, 29, 30) of a "Protaster" that probably will prove
to be new. They are from the Trenton limestone at Trenton Falls,
New York.
ALEPIDASTER GRANULIFERUS (Meek).
Text fig. 26.
Protaster ? granuliferus (Alepidaster at end of description) MEEK, Amer. Journ.
Sci., ser. 3, vol. 4, 1872, p. 274; Geol. Surv. Ohio, Pal., vol. 1, 1873, p. 68,
pi. 3 bis, figs. 8a, 86. J. F. JAMES, Journ. Cincinnati Soc. Nat. Hist., vol. 18,
1896, p. 138. PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 368.
Original description. "Disk small, apparently circular; rays rather
slender, and of unknown length. Dorsal surface of disk and rays
covered by an integument composed of innumerable minute grains
of calcareous matter. Ventral side of disk not well exposed in the
specimen but apparently provided, in the interradial spaces, with
minute spines directed outward. Oral pieces not well exposed in
the specimen. Arm pieces [ambulacrals] regularly alternating, but
apparently rectangular at their inner ends, and not interlocking along
the minute mesial impressed line, longer transversely than in the
direction of the length of the rays; each largely excavated at its
anterior outer end so as to form a large pore, or porelike depression,
and divided transversely by a furrow into two parts, the anterior of
which is very short, and the posterior longer, and marked by a minute
[muscle] pit at its inner end; about eight or nine of these pieces in
each range of each ray included within the margin of the disk. Outer
arm [or ray] pieces (adambulacral of some) smaller than those of the
the inner ranges, and placed edge upward, with an oblique outward
direction so as to imbricate outward or toward the extremities of
the rays, each bearing one or more minute articulating spines.
" Breadth of disk, about 0.43 inch; breadth of arms at their inner
ends, 0.10 inch."
REVISION OF PALEOZOIC STELLEROIDEA. 231
Formation and locality. In the lower part of the Richmondian at
Moore's Hill, Indiana. It was found by C. B. Dyer, and is now in
the Museum of Comparative Zoology, Harvard University (No. 21).
ALEPIDASTER FLEXUOSUS (Miller and Dyer).
Plate 36, fig. 4.
Protaster flexuosus MILLER and DYER, Journ. Cincinnati Soc. Nat. Hist., vol. 1,
1878, p. 31, pi. 2, figs. 1, la. MILLER, N. Amer. Geol. Pal., 1889, p. 276,
fig. 409. PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 368.
Protasterina ("Protaster" lapsus) fimbriata ULRICH, Journ. Cincinnati Soc. Nat.
Hist., vol. 1, 1878, p. 95, pi. 4, figs. 9-9c.
Protasterina flexuosa and P. fimbriata J. F. JAMES, Journ. Cincinnati Soc. Nat.
Hist., vol. 18, 1896, pp. 139, 140.
Original description of P. flexuosus. " The disk is composed of very
thin, small plates, the order of arrangement of which is not deter-
mined. The specimens examined differ in size, and show the disk
varying in diameter from one-quarter to one-half an inch. The rays
were very flexuous when living, and are found winding and turning
and thrown in different directions in different specimens.
"Four series of plates are seen upon the dorsal side of each ray
near the disk. The two inner series [the ambulacrals seen from the
dorsal side] form an angular ridge, each plate is concave at the uniting
surface, the arrangement is alternate, and the appearance, therefore,
of the top of the ray, is something like the alternate arrangement of
two series of hourglasses. The outer series, or marginal [side] plates,
are spinous, the spines directed toward the point of the ray. A
weathered ray shows three series of pores one row between the inner
series of plates, and one between the marginal plates and inner series,
upon each side of the ray [probably the plates are worn through by
weathering; there should be but two rows of podial openings, the
others being either muscle pits or weathering holes]. The plates are
a little the longest in the direction of the length of the ray, and in
this direction there are about four to the line. The rays cross the
disk on the dorsal side and unite near the center of it, but the speci-
men figured, Plate II, figure 1, has this part of the rays removed.
Plate II, figure la, illustrates the ventral side of a specimen, which
is too indistinct to show the arrangement of the plates. Small
pieces, showing the ventral side of the rays, from other specimens,
have two rows of spines springing from the marginal plates on each
side of the rays."
Original description of P. fimbriata. 1 "Disk of medium size, circu-
lar. Dorsal side of disk, and rays to margin of disk, covered with a
granular integument. Ventral surface of disk provided with a large
number of outwardly directed, short and slender spines. Oral
i Ulrich still maintains that this species is a good one and distinct from P. flexuosus.
232 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
pieces 10, subrhomboidal in outline, arranged in pairs, each pair
being formed of two of the marginal series of ray plates ; on the lower
inner edge there are five spines, and extending from each pair,
over the mouth, is a bundle of rather long ones.
"Rays apparently very flexible, contracted toward the mouth;
six series of plates are exposed on the ventral side of each ray. The
two middle series [ambulacrals] alternate regularly, and interlock
along the mesial line; they are about twice as long in the direction
of the ray, than measuring transversely; each piece is contracted
in the middle on the inner side to admit the wide ends of the two
pieces immediately opposite, and on the outer side to form a pore-
like impression, between them and the outer series of plates; four
[five or possibly six] of these plates in each range of each ray are
included within the disk, and the series terminate abruptly at a
distance of one and a half lines from the inner end of the oral plates;
from the margin of the disk to the extremity of the ray there are
about 16 pieces in each range. The pores mentioned above, are
occupied by a series of loosely fitting, obtusely conical or pyramidal
plates, some of which distinctly show a deep depression in the top,
and may have been perforated. Outer or marginal pieces [side
plates] flat, placed on edge, and directed obliquely outward so as
to overlap each other; the two free edges, that is, the one toward
the point of the rays and the one seen on the ventral surface, are
lined with from 10 to 12 short club-shaped spines, varying somewhat
in length. The marginal [side plates] and middle [ambulacral] series
of plates articulate by means of corresponding prolongations from
the sides of the plates. These prolongations arise from near that
end of the plate which is directed toward the mouth.
" Dorsal side of rays composed of two rows of alternating and
interlocking plates, which near the disk are very deeply sculptured
and about as wide as long, becoming gradually less excavated, and
longer in proportion to the width, toward the point of the rays; on
each side they articulate with the upper edge of the oblique mar-
ginal series of plates [these are the ambulacral plates seen from the
dorsal side, either because there were no dorsal plates or because
there was an integument].
" Breadth of disk, 0.60 inch; breadth of arm at margin of disk,
0.16 inch; length of same from oral plates, 0.88 inch. This species
is related to Protaster flexuosus"
Formation and locality. "This species has been found at differ-
ent elevations from near low-water mark in the Ohio River [that
is, in basal Eden] to the top of the hills at Cincinnati," in the upper
Maysville beds. The type-specimens of P. flexuosus were found
by Mr. C. B. Dyer and are now in the collection of Harvard Uni-
versity (Nos. 17, 18). Those of P. fimbriata were obtained by
REVISIOH OF PALEOZOIC STELLEEOIDEA. 233
Ulrich in the lower Eden at about 100 feet above low-water mark
in the Ohio River at Covlngton, Kentucky. This material is now
in the Ulrich collection of the United States National Museum.
Other poor specimens from a still lower level in the Eden and asso-
ciated with Triarihrus are from the first ward, eastern Cincinnati.
Two specimens are at Yale University.
Cat. No. 60615, U.S.N.M.
ALEPIDASTER MIAMIENSIS (Miller).
Protaster miamiensis MILLER, Journ. Cincinnati Soc. Nat. Hist., vol. 5, 1882,
p. 116, pi. 5, figs. 6-66. J. F. JAMES, Journ. Cincinnati Soc. Nat. Hist.,
vol. 18, 1896, p. 138. PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 368.
Original description. "This species is large, with a proportion-
ally small disk. A specimen having a disk four-tenths of an inch
in diameter, has rays an inch in length. Five specimens have
been examined, all showing the ventral side. The dorsal side is
unknown. Only that part of the disk between the rays is visible
in any of the specimens, and the plates are so anchylosed together
that no special definition of them can be given. The rays are long
and coarser and stronger than usual in this genus, though they were
quite as pliable and flexuous when living as others. Two series of
subquadrangular plates, or ambulacral ossicles, alternating with
each other, constitute the bottom of each ambulacral furrow; these
are bordered by spinous adambulacral plates, which terminate at
the angles of the mouth in only five oral plates."
Formation and locality. In the Richmond formation, about
Waynesville, Ohio. The specimens are in the Harris collection
of the United States National Museum.
Cat. No. 40886, U.S.N.M.
Genus GREGORIURA Chapman.
Gregoriura CHAPMAN, Proc. Royal Soc. Victoria, n. ser., vol. 19, 1907, p. 24, pi. 6,
fig. 1; pi. 8, figs. 1, 3. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wies-
baden, vol. 63, 1910, p. 238.
Original definition. "A Protasterid in which the usual boot-
shaped ambulacrals are laterally developed, and modified into a
subtriangular form. Ossicles on each side of the ambulacral canal
subalternate, excepting at the junction with the mouth frames,
where they are parallel. Adambulacral ossicles narrow, slender,
extending laterally in a line with the proximal border of the ambu-
lacral ossicle. Spine-bearing plates, slender, at right angles to the
adambulacrals, carrying (in the genotype) two conspicuous spines.
Oral skeleton having jaw plates three-fourths the length of the
mouth frames; teeth thick and prominent. No traces of a disk
preserved in the specimen on which the genus is founded. Arms
very slender and very flexible."
234 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genoholotype and only species. G. spryi Chapman (same references
as above).
Found in the Siluric (Melbournian) shale at South Yarra, Vic-
toria, Australia.
Genus BUNDENBACHIA Sturtz.
Text fig. 27.
Bundenbachia STURTZ, Palaeontographica, vol. 32, 1886, p. 83; vol. 36, 1890,
p. 216. GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1033, fig. 4
on p. 1034. PARKS, Trans. Canadian Inst., vol. 8, 1908, p. 367. SCHON-
DORF, Palseontographica, vol. 57, 1910, p. 57.
Rays five, short, basally wide, terminally pointed or blunt. Disk
large, soft and delicate, with concave interradial margins.
Abactinally the disk is covered with symmetrical scales, and
medially there is a pentamerous rosette (the oral armature squeezed
through), from the points of which the rays originate. Rays covered
with a granular integument.
Gregory states : "Ambulacral ossicles with
a dumbbell-shaped body and thin tapering
wing. The body of the ossicle is apparently
divided into two pieces by a transverse de-
pression. The adambulacral plates are small
an( * narrow > an d support a triangular spine-
bearing plate. [There is only one plate here,
FIG. 27.- VENTRAL ABM STRTTC- L J >
TURK OF BUJNDENBACHIA. EN- the adambulacral. I here are no lateral
LARGED. AFTER GREGORY. pieces as in Ophiuroidea.]
"The syngnaths are curved, narrow bars."
Genoholotype and only species. B. beneckei Sturtz.
Remarks. " Bundenbachia differs from Protaster by the irregular
nature and soft plating of the disk, by the presence of spine-bearing
plates attached to the adambulacral ossicles, and by the different
form of the ambulacral ossicles." (Gregory.)
BUNDENBACHIA BENECKEI Sturtz.
Bundenbachia beneckei STURTZ, Palseontographica, vol. 32, 1886, p. 83, pi. 8, figs. 7,
7a; pi. 9, figs. 1, la. GREGORY, Proc. Zool. Soc. London for 1896, 1897, p.
1034, figs. 4.
Formation and locality. Occurs in the Lower Devonic roofing
slates of Bundenbach, Germany.
Genus PAL^EOPHIOMYXA Sturtz.
Palseophiomyxa STURTZ, Verb, naturh. Ver. preuss. Kheinl., etc., vol. 56, 1900,
p. 202.
Palxophwmyxidx STURTZ, Verh. naturh. Ver. preuss. Kheinl., etc., vol. 56, 1900,
p. 202. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 235.
Genoholotype and only species. P. grandis (Sturtz).
REVISION OF PALEOZOIC STELLEROIDEA. 235
PAL^OPHIOMYXA GRANDIS (Sttirtz).
Bundenbachia grandis STURTZ, Palaeontographica, vol. 32, 1886, p. 84, pi. 9, figs.
2, 2a, 3, 3a.
Palseophiomyxa grandis STURTZ, Verh. naturh. Ver. preuss. Rhei.nl., etc., vol. 56,
1900, pp. 196, 202.
Formation and locality. Lower Devonic roofing slates of Bun-
denbach, Germany.
Cat. No. 59388, U.S.N.M.
Family PAL^OPHIURID^E Gregory.
Palseophiuridx GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1034; Treat.
Zool., vol. 3, Echinoderma, 1900, p. 274. SCHONDORF, Jahrb. nassauisch.
Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 237.
"Lysophiuras in which the [alternate] ambulacral ossicles consist
of a bar-shaped or subquadrate 'body' without wings. " (Gregory,
1900.)
Remarks. This family differs from the Protasteridse in "having
the ambulacral ossicles longer than wide, and never divided trans-
versely by muscular depressions. The ambulacral ossicles are either
bar-shaped or thickened to a subquadrate form. They are never
boot-shaped." (Gregory, 1897.)
Schondorf states that Gregory established the families Protas-
terida6 and Palaeophiuridae on the form of the ambulacralia. He
says: "The first family has boot-shaped, and the second family
bar-shaped or subquadrate ambulacrals. The form of the latter is
nothing more than a poor development of the boot-shaped type of
ambulacrals. Accordingly the second family has no value and is
to be withdrawn." In addition the forms of Sturtzura have boot-
shaped ambulacrals, and these "are so little alternate in arrangement
that their alternation appears to me more than questionable."
(Schondorf, 1910.)
Contains the following genera:
Pal&ophiura Stiirtz.
Sturtzura Gregory.
Eugasterella, new name.
Ptilonaster Hall.
Genus PAL^EOPHIURA Stlirtz.
Palxophiura STURTZ, Palseontographica, vol. 36, 1890, p. 233. GREGORY, Proc.
Zool. Soc. London for 1896, 1897, p. 1034.
" Palaeophiuridse with the disk surrounded by rod-shaped marginal
ossicles. The ambulacral ossicles are rods lying parallel to the arm. "
(Gregory.)
Genoholotype and only species. P. simplex Sturtz (citations as
above). Lower Devonic, Bundenbach, Germany.
236 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Genus STURTZURA Gregory.
Sturtzura GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1034. SCHON-
DORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, pp. 215, 237.
fRhodostoma SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London, ser. B, vol.
202, 1912, p. 225 (genoholotype, Protaster leptosoma Salter.)
" Palseophiuridse having thick, subquadrate, ambulacral ossicles
and narrow adambulacral plates. The disk is fragile, and its plates
are small and thin: it has no marginal plates. The mouth-frames
are narrow and separate.
"Distribution. Silurian, England and Australia."
Genoholotype. Protaster brisingoides Gregory.
"Remarks. This genus differs from PalseopTiiura, as the ambu-
lacral ossicles are thick and subquadrate, instead of being in the
form of narrow bars; also by the absence of the strong marginal
plates round the disk." (Gregory.)
Chapman in 1907 refers the genotype back to Protaster, adding:
"In consequence of this determination Sturtzura leptosoma may now
be considered as the type of the genus." This is in violation of the
rules of nomenclature and the generic name will stand or fall on the
basis of Protaster brisingoides.
Schondorf states: "The ambulacra of S. brisingoides are not
at all bar-shaped or subquadrate, if the figures of Chapman ( 1907,
pi. 8, fig. 2) are correct, but are also essentially boot-shaped and
alternate in arrangement. S. leptosoma also has alternating ambu-
lacra. Their identity with LapwortTiura miltoni ranges them with
the Auluroidea, where S. brisingoides undoubtedly also belongs.
That this form and with it also the genus Sturtzura is to be retained
as valid is very questionable, but the final answer may as well wait
until the originals are restudied. In any event it is plain that
both forms can not belong to the Lysophiurse."
Contains the following species:
S. brisingoides (Gregory). Siluric of Australia.
S. leptosoma (Salter). Ludlow of England.
S. leptosomoides Chapman. Siluric of Australia.
STtJRTZURA BRISINGOIDES (Gregory).
Tseniaster australis McCoY, see CHAPMAN 1907, p. 23 (nomen nudum).
Protaster brisingoides GREGORY, Geol. Mag., dec. 3, vol. 6, 1889, pp. 24-27, fig. 1.
CHAPMAN, Proc. Roy. Soc. Victoria, n. ser., vol. 19, 1907, p. 22, pi. 6,
fig. 2; pi. g; fig. 2.
Sturtzura brisingoides GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1034.
Formation and locality. From Siluric sandstones at Moonee Ponds
Creek, Flemington, near Melbourne, Australia. The type-material
is in the British Museum (Natural History), London. Other speci-
mens from the same locality and from the parish of Yering, Upper
Yarra district. Victoria, are in the National Museum at Melbourne.
REVISION" OF PALEOZOIC STELLEEOIDEA. 237
STtfRTZURA LEPTOSOMA (Salter).
Protaster leptosoma SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 331,
pi. 9, fig. 5. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1
(Palaeontogr. Soc. for 1861), 1862, p. 33. QUENSTEDT, Petrefactenktmde
Deutschlands, vol. 4, 1876, p. 133, pi. 95, fig. 12. STURTZ, Palseonto-
graphica, vol. 32, 1886, p. 79; N. Jahrb. fiirMin., etc., 1886, vol. 2, p. 148.
Sturtzura leptosoma GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1035.
CHAPMAN, Proc. Roy. Soc. Victoria, n. ser., vol. 19, 1907, p. 26, pi. 8, fig. 5 .
SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910,
p. 215.
Rhodostoma leptosoma SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London,
ser. B, vol. 202, 1912, p. 223, text fig. 40, pi. 9, figs. 3, 4.
Formation and locality. Common in the Siluric, Ludlow forma-
tion, at Leintwardine, Shropshire, England.
Remarks. Schondorf says S. leptosoma is but a miniature edition
of Lapworthura miltoni, both forms being completely identical. The
disk margin is not convex but concave between the radii.
STURTZURA LEPTOSOMOIDES Chapman.
Sturtzura leptosomoides CHAPMAN, Proc. Roy. Soc. Victoria, n. ser., vol. 19,
1907, p. 26, pi. 7, figs. 1, 2; pi. 8, fig. 4.
Formation and locality. In the Siluric (Melbournian), Moonee
Ponds Creek, Flemington, near Melbourne, Australia.
Genus ETJGASTERELLA, new name.
Eugaster HALL (not Eugaster Seville 1839, Orthoptera), Twentieth Rep. N. Y.
State Cab. Nat. Hist., 1868, p. 290; rev. ed., 1868=1870, p. 332. ZITTEL,
Handb. Pal., vol. 1, 1879, p. 444. STURTZ, N. Jahrb. fur. Min., etc., 1886,
vol. 2, p. 151. MILLER, N. Amer. Geol. Pal., 1889, p. 244. STURTZ, Verh.
naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, p. 20. GREGORY, Proc.
Zool. Soc. London for 1896, 1897, p. 1035.
Eugasterella SCHUCHERT, Fossilium Catalogus, Animalia, pt. 3, April, 1914,
p. 19.
Original description. "Body stellate, consisting of a central alated
disk, and five long, slender, somewhat flexuous rays. Disk composed
of small, polygonal tuberculose or subspinose plates on the ventral
side. Rays consisting of a double series of alternating subquadrate
ambulacral ossicles, and a series of curved adambulacral plates
bordering the grooves, and forming the margin of the ray; the outer
ends overlap the edge of the next plate in advance. Oral plates ten,
arranged in pairs, forming the terminal plates of the adambulacral
ranges. Pores large, arranged in two rows in each ray; penetrating
the ray at the junction of the ambulacral and adambulacral plates
hi such a manner that four different plates border each perforation.
Adambulacral and disk-plates bearing spines. Dorsal surface
unknown."
Genoholotype. E. logani (Hall).
238 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Gregory comments on this genus as follows :
"Palseophiuridae in which the ambulacral ossicles are subhep-
tagonal in form, the central suture along the arm being zigzag, while
the outer angles of the ossicles are cut away for the reception of the
podia. The adambulacral ossicles have a flat base, and thence bend
forward crescentically. The mouth frames are massive, and those
of each pair meet along the middle line of the oral angle.
"This genus I only know from Prof. James Hall's figures, and, in
spite of their clearness, I feel much doubt as to the wisdom of diag-
nosing it from these alone. Its affinities are clearly with the Palse-
ophiuridae, but it approaches the Protasteridse in one respect, for,
owing to the deep depressions in the outer angles of the ambulacral
ossicles, the outer side forms a short rudimentary wing. The genus
differs from all the Protasteridae by the absence of a muscular groove
across the ambulacral ossicles.
" Among the Palseophiuridse it differs from Palseophiura by the
absence of marginal ossicles, and from Sturtzura and T&niura by
the massive nature of the mouth frames."
Contains :
E. logani (Hall). Hamilton.
E. (?) concinna (Ringueberg) . Rochester.
EUGASTERELLA LOGANI (Hall).
Eugaster logani HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 290;
rev. ed., 1868=1870, p. 333, pi. 9, figs. 7, 8.
Original description. "Body stellate, with a small disk and long
attenuate fiexuose rays. Disk composed of numerous small poly-
gonal plates with radiated surfaces; the diameter, measured from the
sinus to its extension on the opposite ray, is about half as great as the
length of the ray measured from the center of the disk. Rays, as seen
from the ventral side, narrow, attenuate, with the ambulacral plates
curving, and near the base of the rays a little wider than long, and
toward the extremities longer than wide, with a strong elevated
transverse ridge. There are about eight pairs of plates inclosed
within the limits of the disk. Pores penetrating the interstices
near the outer extremities of the plates, while near the inner end there
is a depression or pit resembling a partially excavated pore. The
adambulacral plates as seen from below are extremely narrow and
very convex on their outer surfaces, forming the margins of the ray.
Oral plates in pairs, narrow and elongate.
"This specimen measures, from the center of the disk to the extrem-
ities of the rays, about one inch and a half; the widest part of the ray,
which is near the margin of the disk, is about an eighth of an inch
in diameter. The rays toward their extremities have the ambula-
cral field covered by the curving of the marginal plates over the
REVISION OF PALEOZOIC STELLEROIDEA. 239
groove, and which, closely uniting by their margins, give a tereti-
form termination of nearly one-third the entire length, each one
having the aspect of a slender crinoidean proboscis.
" Geological formation and locality. In the Hamilton group, near
Fenner, in Madison County, New York. The specimen was collected
during the geological survey, and is now in the State Museum."
EUGASTERELLA (?) CONCINNA (Ringueberg).
Eugaster condnnus RINGUEBERG, Bull. Buffalo Soc. Nat. Sci., vol. 5, 1886, p. 8,
pi. 1, fig. 3.
This species does not belong to EugastereUa, but until the holo-
type is restudied it can remain in this genus. It is from the Kochester
shale at Lockport, New York, and is in Dr. Bingueberg's collection.
Original description. "Disk flat, thin, alated, composed of very
fine granulose plates. Rays broad at their base; flattened, rapidly
tapering for two-thirds of their length; terminal third attenuate,
rounded; plates, if any, undefinable.
" Dorsal side with four series of plates; two median and two lateral;
the transverse sutures dividing the outer series are continuous with
every other transverse suture dividing the central series. Medial
series with twice the number of plates of the outer, and are raised
above them.
" Sutures depressed. Surface of the plates rounded and finely
granular.
"The lateral series of plates decrease regularly in size after reach-
ing the disk until they end in a pronounced elevation formed by the
last pair of the medial series at a point about halfway from the
border toward the center of the disk measuring at the narrowest
part where both series end. Between these elevations the disk
has a shallow, stellate depression with the points opposite the several
rays. The two outer rows of plates apparently disappear toward
the attenuate tip before the median does so. The imperfect cast
of the upper part of one of the rays is all that is known of the ventral
side; this shows traces of an alternating series of ambulacral plates.
"Marginal plates spiniferous.
"Length of ray, one-half inch."
Genus PTILONASTER Hall.
Ptilonaster HALL, Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, pp. 291-292;
rev. ed., 1868=1870, p. 334. LUTKEN, Dansk. Vid. Selsk. Skrift., ser. 5,
vol. 8, pt. 3, 1869, p. 82. STURTZ, Palseontographica, vol. 32, 1886, p. 83;
N. Jahrb. fur Min., etc., 1886, vol. 2, p. 152. GREGORY, Proc. Zool. Soc.
London for 1896, 1897, p. 1036. STURTZ, Verb, naturb. Ver. preuss. Rbeinl.,
etc., vol. 56, 1900, p. 188.
Palseocoma MILLER (part), N. Amer. Geol. Pal., 1889, p. 266.
Original description. "Form and general features as in Eugaster,
but differing in the plates of the rays. Rays composed of an ambu-
Bull. 88 15 16
240 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
lacral, adambulacral and marginal series, which are united by their
edges, and apparently not imbricating, the projecting or oblique
anterior face of the marginal plates bearing spines which are inclined
toward the extremity of the ray. Margins of the rays alated by the
extension of the disk.
"I had originally united this form with Eugaster, but further ex-
amination has shown the ray to have a range of marginal plates
outside of the adambulacral plates, and I am unable to discover any
such feature in Eugaster logani.
"This and the preceding genus [EugastereTla] belong to the Ophiu-
ridae, to which may also be referred the genus Protaster."
" Note. It is only as these pages are going to press that my atten-
tion has been directed to the similarity of structure in the ray of
Palseocoma of Salter with that of the proposed genus Ptilonaster.
In this illustrated species of the former genus (Palseocoma marstoni)
the disk is proportionately larger, and the rays much shorter, while
the outer range of plates is represented as imbricating; but being
characterized by a double row of plates bordering the ambulacral
area, it must be regarded as very nearly related if not generically
identical with Ptilonaster. 11
Gregory comments on Ptilonaster as follows: "This genus is an ally
of Eugaster; * * * it is, however, generically distinct. I only
know it from Hall's figures, and therefore prefer to leave the prep-
aration of a formal diagnosis to an American palaeontologist."
GenoJwlotype and only species. P. princeps Hall.
PTILONASTER PRINCEPS Hall.
Ptilonaster princeps HAT.?., Twentieth Hep. N. Y. State Cab. Nat. Hist., 1868,
p. 292; rev. ed., 1868=1870, p. 334, pi. 9, fig. 9.
Palseocoma princeps MILLER, N. Amer. Geol. Pal., 1889, p. 267.
Original description. "The specimen consists of the impression of
the greater part of one ray, with parts of two others, and intervening
portions of the disk. The disk has extended along the ray nearly an
inch from the center of the body; the plates are small, and have been
furnished with slender spinules. The ray is strong and extremely
elongate, having been at least 4 inches in length; its greatest width
is outside of the disk, where it measures seven-sixteenths of an inch.
The ventral side of the ray shows three series of plates ambulacral,
adambulacral, and marginal on each side of the center; of these at
least seven ranges have been included within the disk. The ambula-
cral plates are a little wider than long, arranged in alternating order.
The pores penetrate the interstices near the outer extremity of the
plates, while the partial or obsolete pores are obscure, becoming
deeper and more conspicuous toward the extremity of the ray. Near
the base of the ray there are 12 plates in the length of an inch,
REVISION OF PALEOZOIC STELLEROIDEA. 241
and 14 plates in the same distance in the central portion, while
toward the extremities there are 20 or more in the same space.
The marginal plates are subhexagonal, a little longer than wide, and
ornamented by long slender spines on their outer margins.
"This is a much larger and more robust species than the Eugaster
logani, and in its entire condition it is the largest star-fish known in
our palaeozoic rocks.
" Geological formation and locality. In the Chemung group. The
specimen is in a brownish-gray sandstone, which weathers to a very
light ashen color. It was received from Henry S. Randall, Esq., of
Cortlandville [New York], in the vicinity of which place it was ob-
tained."
The holotype, an imprint, is in the American Museum of Natural
History, No. 4472.
Family ENCRINASTERID^E, new name.
Aspidosomatidse GREGORY, Geol. Mag., dec. 4, vol. 6, 1899, p. 351; Treat. Zool.,
vol. 3, Echinoderma, 1900, p. 250. SCHONDORF, Palseontographica, vol. 57,
1910, pp. 55-61.
Palxgoniasteridce STURTZ (part), Palseontographica, vol. 36, 1890, p. 246 (contains
Aspidosomella and Palxaster).
Lysophiurae with boot-shaped ambulacralia ventrally, and large
disks that are bordered with simple marginalia. Apparently only
the inframarginals are present.
As Aspidosoma Goldfuss is preoccupied, this name can no longer
be used for the family name. Encrinaster Haeckel replaces it and
therefore becomes the base of the family designation.
The family has the single genus Encrinaster.
Genus ENCRINASTER Haeckel.
Plate 35, text fig. 28, 29.
Aspidosoma GOLDFUSS (not Aspidosoma Fitzinger 1843, Keptilia), Verh. naturh.
Ver. preuss. Rheinl., etc., vol. 5, 1848, pp. 145-146. SIMONOVITSCH, Sitzb.
d. mat.-naturw. Classe Akad. Wiss., Wien., vol. 64, Abt. 1, 1871, p. 103.
ZITTEL, Handb. Pal., vol. 1, 1879, p. 451, fig. 322. STURTZ, Palseonto-
graphica, vol. 32, 1886, p. 92; N. Jahrb. fiir Min., etc., 1886, vol. 2, p. 151;
Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 38, 55. SCHON-
DORP, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909, p. 35;
Palseontographica, vol. 57, 1910, pp. 1-63. SPENCER, Mon. Brit. Pal. Aste-
rozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, pp. 23, 24, 49.
Encrinaster HAECKEL, Gen. Morphologie, vol. 2, 1866, p. Ixvii (genolectotype
Aspidosoma arnoldi Goldfuss).
Diagnosis. Animals attaining large size, with a comparatively
large disk. Rays five, more or less petaloid, slender and long, with
wide and open ambulacral furrows.
242 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
"Ambulacrals alternating in adjoining columns, while the neigh-
boring ambulacrals and adambulacrals are directly opposite each
other. The margin of the disk is concave and limited by a row
of more or less conspicuous simple [not two rows of superposed]
marginalia restricted to the disk. The sides of the free rays are
bounded by adambulacrals. These and the ambulacrals are [common
to the two sides of the animals, and abactinally are] roughly sculp-
tured, usually naked, and only
in rare cases were adambulacral
spines observed. Madreporite
actinal, situated near the peri-
stome in one of the interradial
~ I " c ' areas nearest to the left adam-
FIG. 28. AMBULACKALIA AND ADAMBULACRALIA OF bulacrals of a radius. 7 ' (Sch6n-
ENCRINASTER ROEMERI, AFTER SCHONDORF. a, AP- A^^t lain "\
- ' '
PEARANCE OF THE PLATES WHEN CRUSHED OR OB-
SCTTRED BY CLAY. EACH AMBULACRAL HERE APPEARS
AS IF OF TWO PIECES; b, SAME BETTER PRESERVED -,-j- (^ O l r lf 11 ^ G. prm q
OR MORE CLEANED; c, THE ACTUAL FORM OF THE ossi- noLai ^ ss - ^ e
CLES WHEN WELL PRESERVED AND COMPLETELY to the Lower Devonic of Rhenish
Germany.
Remarks. This well-known genus has been referred to the asterids
and to the ophiurids, and though it has most often been placed
among the true starfishes, it now appears to belong to neither of
these orders. Schondorf has restudied the material with great
care and has made out the detail of the ambulacralia, with the
result that he erects for Aspidosoma the order Auluroidea. In
the work of 1910 cited above the matter is set forth in great detail,
and all that can be done here is to direct atten-
tion to the more important of his illustrations,
with their descriptions, reproduced in part here
and on pages 213 and 214.
As Aspidosoma was first used for a genus of
reptiles, it can not, under the rules of nomen-
clature, be used again for another group of
animals. It is possible, however, under the FIG. 29. ORAL SKELETON OF
rules to make use of Enerinaster, but in doing ^7*^"!
SO the proceeding should first be explained. MUCH ENLARGED. AFTER
Encrinaster was proposed by Haeckel as fol- SCHONDOEF -
lows : In defining his sixth order of asterids, the Crinastra or lily stars,
forms that he said already had crinoid characters, he cited as exam-
ples of it Protaster sedgwickii and P. = Aspidosoma arnoldi. These
forms, he added, are so different from Protaster miltoni ( i that we here
separate the former as Encrinaster (E. sedgwickii, E. arnoldi, etc.)."
Under ordinary circumstances it would be best to disregard the
name because of the inadequate definition, but as Aspidosoma is
preoccupied, we are obliged to propose another name to take its
EEVISION OF PALEOZOIC STELLEKOIDEA. 243
place. The first mentioned genotype of Encrinaster is Protaster
sedgwickii, but as this is also the type-species of Protaster, a genus
in good standing, we may accept the second named species, Aspido-
soma arnoldi. Therefore it is here proposed to replace Aspidosoma
Goldfuss (not Fitzinger) by Encrinaster Haeckel, with Aspidosoma
arnoldi Goldfuss as the genolectotype.
ENCRINASTER ARNOLDI (Goldfuss).
Aspidosoma arnoldi GOLDFUSS, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 5,
1848, pp. 145-146, pi. 5. MULLER, Verh. naturh. Ver. preuss. Rheinl.,
etc., vol. 12, 1855, p. 4, pi. 1, figs. 1-3. STURTZ, Palaeontographica, vol. 32,
1886, p. 93. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 62, 1909, p. 38; Palseontographica, vol. 57, 1910, pp. 4, 62, pi. 1, fig. 2;
pi. 2, figs. 6, 7; pi. 3, figs. 7-9; Jahrb. preuss. geol. Landesanst., Berlin, vol.
29, 1908, p. 706. SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr.
Soc. for 1913), 1914, p. 40.
Encrinaster arnoldi HAECKEL, Gen. Morphologie, vol. 2, 1866, p. Ixvii.
Formation and locality. This species is common in the grauwacke
of the Lower Coblenzian at Winningen, on the Mosel, Germany.
ENCRINASTER GOLDFUSSI (Schbndori).
Text fig. 29.
Aspidosoma goldfussi SCHONDORF, Palseontographica, vol. 57, 1910, pp. 11, 62,
pi. 3, figs. 1, 6.
Formation and locality. A common form found associated with
E. arnoldi about Winningen, Germany.
ENCRINASTER PETALOIDES (Simonovitsch).
Plate 35, figs. 5, 6.
Aspidosoma petaloides SIMONOVITSCH, Sitzb. mat.-naturw. Classe Akad. Wiss.,
Wien., vol. 64, Abt. 1, 1871, p. 106, pi. 4, figs. 1-3. ZITTEL, Handb. Pal.,
vol. 1, 1879, p. 453, text fig. 322. STURTZ, Palseontographica, vol. 32, 1886,
p. 92. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62,
1909, p. -37, pi. 4, figs. 4, 5; Palaeontographica, vol. 57, 1910, pp. 14, 63, pi. 1,
figs. 3, 4; pi. 2, fig. 4; pi. 3, figs. 10-16 (complete synonymy given here).
This species is described in great detail by its author and by
Schondorf, and is well illustrated. Here as elsewhere the abactinal
skeleton is that of the actinal area seen from the dorsal side.
Four specimens of this species were found in grauwacke of Upper
Coblenzian age, in the old quarry back of the Hohenrheiner furnace
at Niederlahnstein, Rhenish Germany.
ENCRINASTER PETALOIDES GOSLARIENSIS (Halfar).
Aspidosoma petaloides, var. goslariensis HALFAR, Jahrb. preuss. geol. Lande-
sanst., Berlin, vol. 13, 1893, p. 186, pi. 10, figs. 1, la. SCHONDORF,
Palaeontographica, vol. 57, 1910, pp. 21, 63, pi. 2, fig. 5; Jahrb. preuss. geol.
Landesanst., Berlin, vol. 29, 1908, p. 707.
Formation and locality. The holotype is from the Lower Devonic
of the Harz. Germany.
244 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
ENCRINASTER TISCHBEINIANUS (Roemer) emend. Sttirtz.
Aspidosoma tischbeinianum ROEMER, Palaeontographica, vol. 9, 1863, pp. 144-146,
pi. 23. QUENSTEDT, Petrefactenkunde Deutschlands, vol. 4, 1876, p. 76,
pi. 92, fig. 39. STURTZ, Palaeontographica, vol. 32, 1886, p. 92, pi. 12, figs.
1, 2. SCHONDORF, Palaeontographica, vol. 57, 1910, pp. 23, 62, pi. 1, fig. 1;
pi. 3, figs. 4, 5 (complete synonymy given here).
Formation and locality. Common in the roofing slates of the
Rhenish Lower Devonic at Bundenbach, near Birkenfeld, Germany.
As many as 60 individuals have been found on a slab.
Cat. No. 35117, U.S.N.M.
ENCRINASTER ROEMERI (SchSndorf).
Text fig. 28.
Aspidosoma tischbeinianum ROEMER (part), Palaeontographica, vol. 9, 1863, pi. 25,
fig. 11.
Aspidosoma roemeri SCHONDORF, Palaeontographica, vol. 57, 1910, pp. 28, 62,
pi. 2, fig. 3; pi. 3, figs. 2, 3.
Formation and locality. Found associated with E. tiscTibeinianus
at Bundenbach, Germany.
ENCRINASTER SCHMIDTI (SchSndorf).
Aspidosoma schmidti SCHONDORF, Palaeontographica, vol. 57, 1910, pp. 31, 63,
pi. 1, fig. 6; pi. 3, figs. 17, 18 (complete synonymy given here).
Formation and locality. A slab with six specimens from the Lower
Devonic (Herdorfer zone of the Siegen) at Oberdielfen ; Germany.
ENCRINASTER EIFELENSIS (SchOndorf).
Aspidosoma eifelense SCHONDORF, Palaeontographica, vol. 57, 1910, pp. 34, 63,
pi. 1, fig. 5.
Formation and locality. Known by several examples from the
Lower Coblenzian at Oberstadtfeld, Eifel, Germany.
ENCRINASTER PONTIS (Clarke).
Aspidosoma ? pontis CLARKE, Monog., I, Serv. Geol. Min. Brasil, 1913, p. 312,
pi. 27, figs. 1-8; Bull. N. Y. State Mus., No. 164, 1913, pi. 35, figs. 1-4.
Original description. "This very graceful starfish occurs in abund-
ance in the soft gray shales of Ponta Grossa, sometimes in clustered
groups with its long and slender arms curled and intertwined, more
often as single individuals lying in beds with shells of Orbiculoidea
and various lamellibranchs. I have counted 20 individuals among
the specimens at my command and, singularly, every one of these is
preserved with its ambulacra! face up. The removal of all calcareous
matter has left only internal casts of this surface from which it has
been possible to gain an idea of external structures from carefully
REVISION OF PALEOZOIC STELLEEOI0EA. 245
made impressions. Na specimen gives any clue to the dorsal exterior
save one arm which has been turned sidewise showing a part of that
surface.
"The striking character of this starfish is the unusual delicacy of
its arms, which are very slender and long for a true asteroid and
present an appearance very suggestive of an ophiuran. These slender
arms are usually so exposed as to show only the alternating am-
bulacral plates which increases the effect of slenderness; but the
adambulacrals are occasionally present and give the arms increased
width. These arms are all clearly sulcate, the median groove being
sharply marked in all casts by a thin elevated ridge running the
entire length of the arm. The soft shale holding these specimens is
not a very satisfactory matrix from which to take the special struc-
tures of the animal but in good part these can be made out. The
ambulacral plates are quadrate in outline, alternate in position along
the arm groove, and together are arched so that normally the am-
bulacral surface is an arched ridge rising above the rest of the surface.
These plates, in their arched position, are slightly spaced or dis-
placed along adjoining edges leaving passages from the ambulacra.
They are usually broken down or slipped over each other in com-
pression so that the arm seldom* shows them in the true position.
The surface of these plates is finely granular. The adambulacrals
constitute one (or more ?) lateral rows and are so seldom well defined
that I am not able to represent them with precision but each of them
seems to carry one short spine and all to be granulated on the surface.
The body of the animal is deeply incut by the arms, which radiate
from a narrow center, sharply quinquepartite by the buccal cavity.
The oral apparatus is pretty well made out. The oral frames, one
for each arm, are very conspicuous, though varying in size in different
individuals. Each is broadly saddle-shaped, split medially, the distal
parts joined by a suture, the proximal ends spreading apart at a
sharp angle which opens into the oral cavity. Their surface is
rounded and granulate, the distal ends elevated, and often each half
is inclined downward toward the other along the median suture.
At each proximal angle lies a sharp jaw or tooth. The whole of the
body surface is occupied by the oral rosette composed of the five
Jbuccal divisions."
Formation and locality. From the Devonic of Ponta Grossa and
Jaguariahyva, Brazil.
ENCRINASTER (?) GRAY^E (Spencer).
Aspidosoma grayse. SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc.
for 1913), 1914, pp. 23, 24, 33, 38, text figs. 22, 24, pi. 1, fig. 8.
Occurs in the Caradoc of Great Britain.
246 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Order STREPTOPHIUR^ Bell.
Streptophiurse BELL, Proc. Zool. Soc. London for 1891, 1892, p. 179. GREGORY,
Proc. Zool. Soc. London for 1896, 1897, p. 1036; Treat. Zool., vol. 3, Echino-
derma, 1900, p. 274.
Protophiurex STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 200. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, pp. 234, 239, 251.
Ophiurasterise SCHONDORF, Palseontographica, vol. 57, 1910, p. 61.
Diagnosis by Gregory (1897). Auluroidea "in which the ambu-
lacral ossicles are opposite and are generally fus.ed into vertebral
ossicles. In such cases the vertebral ossicles articulate by a more
or less simple ball-and-socket joint."
"Remarks. The main character of this order is that the ambu-
lacral ossicles are paired, but primitive. The order differs from the
preceding [Lysophiurse] by having the vertebral ossicles always
opposite instead of alternate. In some of the simplest members of
the order, as Ophiurina, the ossicles are not fused, there are no
ventral arm shields, and an ambulacral furrow runs along the ven-
tral side of the arm. In the next higher family, as in the genus
Lapworihura, the ambulacral ossicles are fused, but have plain
articulating surfaces, and there is* an ambulacral furrow."
Contains the f amilies :
Ophiurinidae.
Lapworthuridas.
Eoluidiidae.
Aganasteridae.
Cholasteridae.
Family OPHIUKINIIXE Gregory.
Ophiurinidse GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1037; Treat.
Zool., vol. 3, Echinoderma, 1900, p. 275. STURTZ, Verh. naturh. Ver. preuss.
Bheinl., etc., vol. 56, 1900, p. 201. SCHONDORF, Jahrb. nassauisch. Ver.
Naturk., Wiesbaden, vol. 63, 1910, p. 234.
Original diagnosis. " Strep tophiurae with ambulacral ossicles,
only slightly united, and without ventral arm plates."
Contains the genera:
Ophiurina Sttirtz.
Tremataster Worthen and Miller.
Genus OPHIURINA Sturtz.
Ophiurina STURTZ, Palaeontographica, vol. 36, 1890, p. 232. GREGORY, Proc.
Zool. Soc. London for 1896, 1897, p. 1037.
Gregory's diagnosis reads as follows: "Disk circular, with mar-
ginal plates. Ambulacral ossicles long, narrow bars. Syngnaths
rod-shaped. Adambulacral plates absent or altogether lost from
the fossil."
KEVISION OF PALEOZOIC STELLEEOIDEA. 247
Genoliolotype and only species. 0. lymani Stiirtz (citations as
above). Lower Devonic, Bundenbach, Germany.
Genus TREMATASTER Worthen and Miller.
Tremataster WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883, p. 330.
MILLER, N. Amer. Geol. Pal., 1889, p. 286. GREGORY, Proc. Zool. Soc.
London for 1896, 1897, p. 1037.
Original description. "Body stellate, central part discoid, rays
long, flexuous, and gradually tapering. Rays consisting of a double
series of ambulacral plates, forming, together, a subcuneiform se-
ries, with the tapering ends directed toward the apices of the rays
and uniting with the larger ends of the succeeding double plates,
upon each side of which there is a series of curved adambulacral
plates, which form the margins of the rays. The pores are large
and situated between the contracting sides of the ambulacral plates
and the concave sides of the curving adambulacrals. Four plates
border upon each pore though nearly surrounded by two of them.
Plates 10. Dorsal surface unknown.
"This genus is distinguished by the parallel arrangement of the
ambulacrals and adambulacrals, instead of the alternate order, and
by the large pores."
Gregory's diagnosis is as follows: "Disk with concave inter-
brachial outlines. No marginal plates. Ambulacral ossicles short.
Adambulacrals present."
Genoholotype. T. difficilis Worthen and Miller. T. (?) decheni
(Dewalque) may also belong here.
TREMATASTER DIFFICILIS Worthen and Miller.
Tremataster diffidlis WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883,
p. 330, pi. 31, figs. 3fl, 36.
Original description. "Body stellate, disk small, rays long,
flexuous, and gradually tapering. Ambulacral plates subtrigonal,
elongated, and united upon their straight faces in parallel order,
so as to form a series of subcuneiform sections at the bottom of
the ambulacral groove, with the tapering end of each directed toward
the apex of the ray, and uniting with the larger end of the next
succeeding double plate.
"Adambulacral plates parallel with the ambulacrals, and each
forms a curve from the larger end of an ambulacral toward the
apex of the ray, to unite with the next succeeding adambulacral as
it curves away from its attachment to the ambulacral. By this
order of construction the adambulacrals are fixed, at one end, to
the ambulacrals, while the other end moves upon the curved surface
of an adambulacral so as to allow the same lateral flexibility of
the rays, secured in other genera by the alternate arrangement of
248 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
the plates. The pores are large and situated between the contract-
ing sides of the ambulacral plates, the concave sides of the adam-
bulacrals, and the attaching ends of the succeeding plates, so that
they are bounded by four plates while nearly surrounded by two.
"The oral plates form, in pairs, elongated triangles, with the
acute angle directed toward the center of the visceral cavity. The
apices are joined by a suture with the basal portions, which are
partly separated by a dividing furrow. A single genital pore per-
forates each basal section of the oral plates. Dorsal side unknown.
"Locality and position: Prairie du Long Creek, Monroe County
[Illinois], Chester limestone.
"No. 2479, Illinois State collection of 1878."
.' TREMATASTER (?) DECHENI (Dewalque).
Protaster decheni DEWALQUE, Ann. Soc. ge"ol. Belgique, vol. 8, 1880, pp. 52-54,
pi. 3, figs. 1, 2.
Tremataster (?) decheni GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1037,
footnote.
Formation and locality. Upper Devonic at Walcourt (Namur)
and Tohogne (Luxemburg), Belgium.
Family LAPWORTHUKID^E Gregory.
Lapworthuridse GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1037.
STURTZ, Verb, naturh. Ver. preuss. RheinL, etc., vol. 56, 1900, p. 203.
GREGORY, Treat. Zool., vol. 3, Echinoderma, 1900, p. 275. SCHONDORF,
Palseontographica, vol. 57, 1910, p. 58; Jahrb. nassauisch. Ver. Naturk.,
Wiesbaden, vol. 63, 1910, pp. 235, 239.
Furcasteridx STURTZ, Verb, naturh. Ver. preuss. RheinL, etc., vol. 56, 1900, p.
203. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910,
p. 235.
Palxospondylidsd STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 201. SCHONDORP, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 235.
Original diagnosis. "Streptophiuroida without ventral armplates
or buccal shields; ambulacral ossicles fused, but their articulating
surfaces are plain. Madreporite dorsal." Schondorf states that this
plate is on the ventral side, and that the ambulacrals are not fused
(p. 239).
Contains the genera:
Squamaster Ringueberg.
Lapworihura Gregory.
Sturtzaster Etherrdge.
Bdellacoma Salter.
RTiopalocoma Salter.
Hallaster Stiirtz.
Sympterura Bather.
Fur caster Stiirtz.
Palastropecten Sttirtz.
REVISION OF PALEOZOIC STELLEROIDEA. 249
Sturtz proposes that the family Lapworthuridse be divided into
three families. In the original family will remain Squamaster, Lap-
worthura, HaTLaster, and Sympterura. The family Furcasteridse has
Furcaster, while the Palaeospondylidae embraces Palseospondylus, the
latter name being later changed to Palastropecten by Sturtz, as it
was preoccupied; the family name will, therefore, have to be
changed to Palastropectinidse, as was done by Bather in 1905 (p. 168).
% Genus SQUAMASTER Ringueberg.
Squamaster RINGUEBERG, Bull. Buffalo Soc. Nat. Sci., vol. 5, 1886, p. 5.
Original description. "Body small. Rays long, slender, tapering,
flexible, and divided into annular segments.
"Ventral side of ray with two series of paired, opposite ambulacral
plates, divided by a straight median line; they are irregularly sub-
quadrilateral, with one short side, and are arranged in pairs in each
series, so that two long and two short sides are joined alternately;
consequently there is a long transverse suture regularly alternating
with a short one. This disposition gives dentate lateral margins to
the ambulacral series. The lateral ends of the long transverse
sutures curve away from each other, leaving a small notch between
the inferior plates of each pair and the superior plates of the adjoining
pairs. This notch receives the inner lateral corners of the squamose
dorsal plates. Between the dentate elevations of the ambulacral
plates and the lateral margins of the dorsal plates there are large
triangular openings, which constituted, or in which were placed, the
pores.
"Dorsal side composed of large, quadrangular, squamose plates,
that encircle the arm as far as the ambulacral groove of which they
form the margin. They are divided in the upper portion of the
ray by a median suture.
"Dorsal, ray plates fimbriated on the free margins.
"All plates are perforated by minute, closely arranged pores."
Genoholotype and only species. 8. echinatus Ringueberg.
SQUAMASTER ECHINATUS Ringueberg.
Squamaster echinatus RINGUEBERG, Bull. Buffalo Soc. Nat. Sci., vol. 5, 1886,
p. 6, pi. 1, fig. 1.
Original description. ' ' Body small. Rays long, slender, not widen-
ing at the base ; regularly tapering to a subobtuse point. Dorsal side
covered by thin imbricating plates, with free external margins which
are edged on the outer and lateral sides with closely arranged, long,
slender, filiform spinules, that articulate into minute rounded sockets,
which are scalloped out of the margin of the plate; they are slightly
longer than the exposed portion of the plate, and are placed about
their own diameter apart.
250 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
''Ray divided into about 40 segments or articulations by the
encircling, dorsal plates; these grow rapidly narrower near the body,
and are divided by a dorsal suture running along a raised median
ridge which extends about two-fifths of the length of the ray from
the body out, where it gradually disappears with the last divided
plate; the rest are entire. These plates are quadrangular, and have
then- inner corners articulated into the socket formed in the dentate
lateral -extensions of the ambulacral series. They are ornamented
on each side by two or three closely arranged, subparallel, very fine
ridges, that commence near the dorsal line at the free margins and
extend diagonally backward across the plate toward the inner lateral
articulated corner. Ambulacral groove furnished with opposite,
irregularly quadrilateral, ambulacral plates, divided by a median
suture which is crossed alternately by one long and one short suture.
"The superior plate of each pair has the greatest extension later-
ally, while in the inferior it is the direction of the median line, it
being about twice as long as the superior, which, however, is but
slightly broader.
"Between the lateral margins of the dorsal plates and the dentate
elevations of the ambulacral plates are large sub trigonal open spaces.
"The plates are punctured by very closely arranged minute
perforations."
Formation and locality. In the Rochester shale, at Lockport, New
York. The holotype is in the collection of Dr. Ringueberg. At
least one (No. 1110) and probably two other specimens (No. 465)
are in the Walker collection of the University of Toronto. They are
from the Rochester shale, at Grimsby, Ontario.
Genus LAPWORTHURA Gregory.
Text fig. 30.
Lapworthura GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1037, fig. 5.
SCHONDORF, Palseontographica, vol. 57, 1910, p. 58; Jahrb. nassauisch. Ver.
Naturk., Wiesbaden, vol. 63, 1910, p. 208. SOLLAS and
SOLLAS, Philos. Trans. Roy. Soc. London, ser. B, vol.
202, 1912, p. 213. SPENCER, Mon. Brit. Pal. Asterozoa,
pt> l (Pa 186011 * ^- Soc. for 1913), 1914, pp. 25, 32, 40, 49.
Original diagnosis. "Disk circular, well
marked. Arms very flexible, broad; at first
FIG. SO.-VENTRAL STRUCTURE uniform in width and then tapering slowly.
OP ARM OF LAPWORTHURA, Ambulacral ossicles with the distal and prox-
AFTER GREGORY. . . .
imal margins parallel; with lateral wings curv-
ing round the podial pores. Madreporite large," dorsal.
Genoholotype. Protaster miltoni Salter.
Schondorf remarks as follows (1910&:58):
The ossicles of Lapworthura have been described several times,
but never correctly. The ambulacrals are undoubtedly opposite,
REVISION OF PALEOZOIC STELLEROIDEA. 251
and both dorsally and ventrally have the structure of Encrinaster.
The adambulacrals are also opposite to the ambulacrals, and have
long forward directed spines. In regard to the dorsal position of
the madreporite, he states that the abactinal area has been mistaken
for the actinal, and that therefore the madreporite lies upon the
ventral side.
The disk is not circular in outline, but concave between the inter-
radii.
LAPWORTHURA MILTONI (Salter).
Protaster miltoni SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 330, pi. 9,
fig. 4. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palse-
ontogr. Soc. for 1861), 1862, p. 32, fig. 18. QUENSTEDT, Petrefactenkunde
Deutschlands, vol. 4, 1876, p. 75, pi. 92, fig. 36. STURTZ, N. Jahrb. fur Min.,
etc., 1886, vol. 2, p. 148; Palseontographica, vol. 32, 1886, pp. 79, 94, pi. 8,
fig. 6.
Lapworthura miltoni GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1037,
fig. 5. SCHONDORF, Palseontographica, vol. 57, 1910, p. 58; Jahrb. nassau-
isch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 208. SOLLAS and SOLLAS,
Philos. Trans. Roy. Soc. London, ser. B, vol. 202, 1912, p. 214, text figs. 1,
2, 4 D, pi. 9, figs. 1, 2; pi. 10, figs. 1-4. SPENCER, Mon. Brit. Pal. Aster-
ozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, pp. 25, 32, 35, 36, pi. 1, fig. 9.
Formation and locality.- Abundant in the Lower Ludlow of the
Siluric at Leintwardine, England. See elsewhere in this work
(p. 237) under Sturtzura leptosoma, which Schondorf states is but
the young of L. miltoni. The same author states further that the
latter species either embraces several species or the form has a long
geologic range, for a number of species and genera appear to be
identical with L. miltoni. .
Cat. No. 59394, U.S.N.M.
LAPWORTHURA SOLLASI Spencer.
Lapworthura sollasi SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc.
for 1913), 1914, pi. 1, fig. 10.
Occurs in the Siluric of Great Britain.
LAPWORTHURA (? "probably n. gen.") species (Miller and Gurley).
Aganaster (?) sp., MILLER and GURLEY, Sixteenth Rep. Geol. Surv. Indiana,
1891, p. 372, pi. 9, figs. 12, 13; authors' extracts, 1890, p. 58. GREGORY,
Proc. Zool. Soc. London for 1896, 1897, p. 1038. '
Original description. "We have figured this fragment to show the
anchylosis of the oral plates, and the pores passing through the
ambulacral plates, and also those passing between them. It will
be observed that one pore passes through each plate near the marginal
end, while a double row of pores passes between the plates in the
central part of the arm furrow. We believe this is an Aganaster,
from the <Jepth of the central part of the disk, and the arrangement
of the plates in the arm furrows; but the specimen is larger than any
252 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Aganaster gregarius we have seen and the rays are wider, and as we
do not know the position of the pores in the rays of Aganaster, it
may be our specimen does not belong to this genus."
Remarks. " Seems to me to be allied to Lapworfhura, and at least
a member of the same family. It apparently has no ventral arm-
plates, but an open furrow, and thus differs widely from Aganaster.
It is probably a new genus." (Gregory.)
Formation and locality. Not given. The specimen may be in
the Gurley collection of the University of Chicago.
Genus STURTZASTER Etheridge.
Palaeocoma SALTER (not D'Orbigny 1850), Rep. Brit. Assoc., 1857, Trans, of sec-
tions, p. 76; Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, pp. 324, 327. WRIGHT,
Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palseontogr. Soc. for 1861),
1862, pp. 23, 29. QUENSTEDT, Petrefactenkunde Deutschlands, vol. 4, 1876,
p. 81, pi. 92, fig. 43. ZITTEL, Handb. Pal., vol. 1, 1879, p. 453. STURTZ,
N. Jahrb. fur Min., etc., 1886, vol. 2, p. 152; Verb, naturh. Ver. preuss. Rbeinl.,
etc., vol. 50, 1893, pp. 45, 62.
Sturtzaster ETHERIDGE, Rec. Australian Mus., vol. 3, 1899, p. 129. SCHONDORF,
Jabrb. nassauiscb. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 217.
Genoholotype. Palseocoma marstoni Salter.
Remarks. Palseocoma = Sturtzaster, Bdellacoma, and Rhopalocoma,
all of Salter, are ver}^ poorly known, due to the fact that the specimens,
though not rare, occur in a calcareous shale and are pressed flat till
they have become a thin film of plates greatly obscured by an abun-
dance of spines. Of P. marstoni, the genotype, the writer has seen
five examples, presented to the United States National Museum by
W. R. Billings. On the basis of the characters revealed by these
specimens, he was inclined to leave the genus among the Asteroidea
and in close association either with the Mesopalaeasterinse (nearest
Mesopalseaster, a phanerozonian) , or the Schuchertiidse (near Schuch-
ertia, a cryptozonian) .
The long, slender and serrated spines of Sturtzaster are peculiar to
the genus. In. their abundance and character they remind one more
of ophiurids than of asterids.
The figures of Salter and Wright, if correct, recall the large-disked
cryptozonian genus Schuchertia, and it was this character that prob-
ably also led Sttirtz to refer Sturtzaster to the same group in asso-
ciation with the genera Palasteriscus, Loriolaster, and OTieiropteraster.
Etheridge in 1899 called attention to the fact that D ' Orbigny pro-
posed in 1850 * the new generic name Palseocoma for OpJiiura mulleri
Phillips of the Lias. A name once proposed can not be used again
in another sense, and therefore Mr. Etheridge is within the rules of
nomenclature in substituting for Salter 's second usage of Palseocoma
1 Prodome, vol. 1, p. 240.
REVISION OF PALEOZOIC STELLEROIDEA. 253
the name of Sturtzaster , with Palseocoma marstoni SaJter as the
genotype.
In 1910 Schondorf was able to study "several right good examples"
of S. marstoni, and strangely the form turns out not to be an asterid
but an aulurid, with the essential ambulacra! characters of Lap-
wortJiura miltoni. He states that the ambulacrals are not alternate,
as stated by Salter, but opposite one another. Therefore the
ambulacrals and the ad ambulacrals are also opposite. The ad am-
bulacrals bear two or three spines on then" outer edges, and Salter's
statement and figures showing two columns of ossicles outside of the
ambulacrals appear to be very wrong. Schondorf states that the
peculiar preservation of the arm skeleton has given rise to this state-
ment of Salter, and that there are only ambulacrals and adambula-
crals present. The oral skeleton is distinctly adambulacral. A
madreporite is not determinable.
On the dorsal side, all that can be clearly made out is the presence
of ambulacrals and adambulacrals, all of which are arranged in trans-
verse rows.
STURTZASTER MARSTONI (Salter).
Palseocoma marstoni SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 328,
pi. 9, figs. 3a-3c. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2,
pt. 1 (Palseontogr. Soc. for 1861), 1862, p. 30, fig. 17. QUENSTEDT, Petrefacten-
kunde Deutschlands, vol. 4, 1876, p. 81, pi. 92, fig. 43.
Sturtzaster marstoni SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 63, 1910, p. 217.
Formation and locality. Common in the Lower Ludlow of Church
Hill, Leintwardine, England. Specimens are in many museums, the
United States National Museum and the Yale Museum both having
individuals. See S. colvini and S. cygnipes.
Cat. Nos. 60624, 60625, U.S.N.M.
STtfRTZASTER COLVINI (Salter).
Palseocoma colvini SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 328.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palseontogr. Soc.
for 1861), 1862, p. 30.
Sturtzaster colvini SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol.
63, 1910, p. 219.
Formation and locality. In the Lower Ludlow, Leintwardine, Eng-
land. "Probably completely identical with 8. marstoni." (Schondorf.)
STtJRTZASTER CYGNIPES (Salter).
Palseocoma cygnipes SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p. 329.
WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palaeontogr. Soc.
for 1861)', 1862, p. 30.
Sturtzaster cygniceps (QIC) SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wies-
baden, vol. 63, 1910, p. 219.
Formation and locality. Lower Ludlow, Leintwardine, England.
" Probably completely identical with S. marstoni." (Schondorf.)
254 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
STtJRTZASTER (?) MITCHELLI Etheridge, jr.
Sturtzaster (?) mitchelli ETHERIDGE, jr., Rec. Australian Mus., vol. 3, 1899, p. 128.
Formation and locality. A single specimen from the Upper Trilo-
bite bed of the Siluric (Wenlock series) at Bowning, New South
Wales.
Genus BDELLACOMA Salter.
Palseocoma (Bdellacoma) SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857, p.
324. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palee-
ontogr. Soc. for 1861), 1862, p. 31. ZITTEL, Handb. Pal., vol. 1, 1879, p.
453. STURTZ, Palseontographica, vol. 32, 1886, p. 96.
Bdellacoma STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893, pp. 46,
63.
Genoholotype and only species. P. (B.) vermiformis Salter. Lower
Ludlow, Leintwardine, England.
Remarks. Until this form is restudied and brought up to date the
supposed subgenus of Paldeocoma has no value.
Genus RHOPALOCOMA Salter.
Palseocoma (Rhopalocoma} SALTER, Ann. Mag. Nat. Hist., ser. 2, vol. 20, 1857,
p. 329. WRIGHT, Mon. British Foss. Echinod., Oolitic, vol. 2, pt. 1 (Palae-
ontogr. Soc. for 1861), 1862, p. 31. ZITTEL, Handb. Pal., vol. 1, 1879, p. 453.
STURTZ, Palaeontographica, vol. 32, 1886, p. 96.
Rhopalocoma STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893,
pp. 46, 63.
GenoJiolotype and only species. P. (R.) pyrotechnica Salter. Lower
Ludlow, Leintwardine, England.
Remarks. This form also requires modernization, and until then
the supposed subgenus has no value.
Genus HALLASTER Sturtz.
Protaster HALL (not Forbes), Nat. Hist. N. Y., Pal., vol. 3, 1861, p. 134;
Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 293; rev. ed., 1868=
1870, p. 336.
Hallaster STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 150.
Original description (Sturtz). "Ambulacral plates alternating
[they are opposite]; the podial pores lie between these and the side
plates. The dorsal skeleton consists of two columns of dorsal plates
and two of lateral plates, of which the former adjoin medially along
the rays. For P. forbesi Hall of the Silurian of North America I
propose the generic name Hallaster, because on the one hand Forbes
did not sufficiently characterize his Protaster, and on the other the
above species in the sense of Salter's Protaster is not congeneric
with it."
GenoJiolotype and only species. Protaster forbesi Hall.
Remarks. This genus appears to be most closely related to Lap-
worihura, with the most obvious difference in the shape of the am-
REVISION OF PALEOZOIC STELLEROIDEA.
255
bulacral ossicles. In Hallaster the lateral wings of the plates curve
round the podial openings proximally, while in Lapworthura they do
so distally. Further, the side plates of the latter genus bear spines
along their sides, while in the former genus they are restricted to the
distal ends of the plates.
HALLASTER FORBESI (Hall).
Text fig. 31.
Protaster forbesi HALL, Nat. Hist. N. Y., Pal., vol. 3, 1861, p. 134, pi. 7 A, figs.
8-10; Twentieth Rep. N. Y. State Cab. Nat. Hist., 1868, p. 293, pi. 9, figs. 5,
6; rev. ed., 1868=1870, p. 336, pi. 9, figs. 5, 6. QUENSTEDT, Petrefacten-
kunde Deutschlands, vol. 4, 1876, p. 134, pi. 95, fig. 13. STUKTZ, Palaeonto-
graphica, vol. 32, 1886, pp. 78, 83.
Hallaster forbesi STURTZ, N. Jahrb. fur Min., etc., 1886, vol. 2, p. 150.
The original description will not be repeated here because it is
faulty and was later corrected by Hall.
Hall's description of 1868. P. forbesi "has a circu-
lar disk, composed of squamif orm spiniferous plates
and five long flexuous rays. These rays I have
[originally] represented as composed, on the lower
side, of a double range of [adambulacral] plates, as
described and represented by Professor Forbes, but
finding outside of these a range of small ossicles
[side plates] to which are attached the spine bases,
these have been shown as a part of an articulating
spine (in the illustration, plate 7 A 1 ), an unnatural representation,
which I am now able to correct.
"In the species from the Lower Helderberg group, Protaster forbesi ,
the ventral surfaces of the rays are composed of an ambulacral and
adambulacral series of plates on each side [there are no ventral ray
plates]. The ambulacral plates are obliquely quadrangular and alter-
nating in a slight degree [in the type they are opposite] ; the adambu-
lacral plates as seen from the lower side are narrow, elongate, oblique,
and laterally imbricating, presenting the appearance of an oblique
ridge with the anterior extremity projecting, and forming the point
of attachment for the spines, with which each one is furnished. [These
spines are striated longitudinally.] When the ray is abruptly curved,
these plates project outward, sometimes almost rectangularly; and
when at the same time the ambulacral area is obscured by aclhering
matrix, these plates might readily be mistaken for appendages of
the inner ranges. The pores are comparatively large, truncating
the outer adjacent angles of the ambulacral plates, while the base
of one adambulacral plate and the side of another form the exterior
margin. The centers of the upper sides of the rays are composed of
FIG. 31. ACTINAI, BAT
PLATES OP HALLASTER
FORBESI (HALL). OP-
POSITE AMBULACRALS,
AND THE SIDE PLATES.
* Nat. Hist. New York, Pal., vol. 3, 1861.
50601 Bull. 8816 17
256 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
two ranges of subimbricating [dorsal] plates, which are closely joined
along the median line; the marginal plates are the upper edges of the
adambulacral plates, which bear on their anterior ends one, two, or
three short spines."
Formation and locality. From the Coeymans member of the Hel-
derbergian series of the Lower Devonic, at Jerusalem Hill, near
Litchfield, New York. The holotype is in the American Museum of
Natural History, No. 2302. There are two good specimens and a
single arm in the Beecher collection at Yale University.
Genus SYMPTERURA Bather.
Text fig. 32.
Sympterura BATHER, Geol. Mag., dec. 5, vol. 2, 1905, pp. 161-169, pi. 6, figs. 1,
2, 3, 6. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 216.
Original diagnosis. "A Lapworthurid with spinulose disk ex-
tending to second arm segment, with oral skeleton of teeth, long
A 3
rame
t&etk
FIG. 32. SYMPTERURA MINVERI, AFTER BATHER. A. SEMIDIAGRAMMATIC RECONSTRUCTION OF SYN-
GNATHS, X 10. B. RECONSTRUCTION OF SEVENTH AND EIGHTH ARM SEGMENTS, x 18: a, DISTAL, AND ft,
PROXIMAL PORTION OF VERTEBRA; C, 'ADAMBULACRALIAJ p, PODIAL DEPRESSION; q, INTERSPACE, OCCU-
PIED, AT LEAST IN PART, BY INTERVERTEBRAL MUSCLES.
jaws, and short mouth frames (torus not seen), with free arm seg-
ments containing a vertebral ossicle, possibly compound, grooved
ventrally and provided on each side with two wings, to the distal of
which is attached an adambulacral spiniferous element."
GenoJiolotype and only species. S. minveri Bather (same references
as above). The holotype was found in the Devonic (?Lower) at
Epphaven, near Padstow, North Cornwall, England.
As this specimen presents in the ambulacralia characters of great
value in the morphology of the Ophiurid vertebrse, it is advisable
to quote here somewhat extensively from Bather's statements.
"The median body of each segment is undoubtedly the equivalent
of the normal Ophiurid vertebra. On the accepted theory that
this -vertebra was evolved by the fusion of a pair of ossicles originally
alternating, then opposite, and finally joined along the middle or
REVISION OF PALEOZOIC STELLEROIDEA. 257
radial line, the present specimen may be described as in a stage of
development later than the fusion, but possibly retaining hi the
median groove a trace of the primitive distinctness of the ossicles.
Those ossicles are usually described as ambulacra!, and that term
may be accepted so far as it denotes homology with the ossicles
so named in an Asterid. * * *
"As a rule among Ophiurids in this stage of evolution, each ver-
tebra has on each side a single side process on which abuts an outer
ossicle, the so-called adambulacral, and these adambulacrals, which
bear the lateral spines, alternate with the ambulacral ossicles or
vertebrae. It is a little difficult to interpret the appearances of the
present specimen in harmony with such an arrangement. On the
assumption that each segment should contain one ambulacral pair
(here fused) and one adambulacral pair bearing spines, ray i has been
assigned nine segments. But each such segment is seen to have two
processes on each side, and the distal one of these bears the spines.
For this reason, and because of the appearances in segment 8, it may
be supposed that the outer spiniferous portion of the distal process
(c in fig. 1) is an adambulacral element. What, then, is the proxi-
mal side process? To this question two possible answers present
themselves.
"The appearance of a transverse division in the vertebra, between
the proximal and distal processes, suggests that each segment really
consists of two ambulacral pairs, of which only the distal one is
associated with adambulacrals, the proximal pair of adambulacrals
being suppressed. Each of these supposed ambulacral pairs pos-
sesses a single pair of lateral processes, but those of the distal pair
are partly overlaid by the adambulacral elements. In the distal
region of the ray the proximal and distal halves of the vertebra are
distinguishable; but nearer the disk and within it they have come
closer together, the transverse division has disappeared, and the
side processes are reduced to structures which appear as bars, but
are probably flattened laminae. Thus the vertebrae seen in ray v are,
on this explanation, of compound origin. * * *
"Now, comparison of the vertebrae of recent Ophiurids with the arm
segments, especially the proximal ones, of our fossil leaves no room for
doubt that the depression marked p is for the reception of a podium,
while that marked q is for the insertion of the ventral intervertebral
muscles. A single arm segment of a recent Ophiurid, such as OpTii-
amchno, or Amphium, is therefore homologous with any one of the
whole segments here numbered 1-8. If one of these is really com-
posed of a distal and a proximal hah 7 (lettered a and & respectively
in figs. 1 and 2) the same statement would apply to the recent Ophi-
urid, and some trace of this origin might be seen in the development.
258 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
The observations of Prof. Ludwig 1 and Mr. J. W. Fewkes 2 on
Amphiura squamata have shown that the two lateral halves of each
vertebra unite first at the proximal and distal ends, leaving a space
between. This is suggestive of an origin from proximal and distal
elements. On the other hand each lateral half springs from a
single center of calcification, from which it stretches out in a distal
direction, so that the incipient ambulacral element is thicker at its
proximal end. This last-mentioned feature is paralleled by the
boot-shaped proximal portions of the ambulacral ossicles in the Prot-
asteridas. At a slightly later stage the ambulacral of Amphiura
broadens at the distal end, which thus comes to resemble the distal
portion of the Protasterid ambulacral. If this distal portion in
Amphiura were, as indicated by Prof. Ludwig's observations, merely
an extension of the proximal portion, then it might be inferred
that the whole structure, both here and in the various Paleozoic
forms alluded to, represented a single ambulacral pair and not two
successive pairs. In such case one would accept Dr. Gregory's
suggested explanation 3 'that the smaller pieces are only triangular,
distal portions of the ambulacral ossicles, apparently separated from
the proximal portion[s] by a groove.' Dr. Gregory further sug-
gests that these grooves or depressions 'were for the lodgment of
the ventral muscles which moved the arms. 7 A far more probable
explanation surely is that they were for the reception of the branches
from the radial water- vessel to the podia. This explanation, how-
ever, consonant as it is with the facts of development of Amphiura,
reminds one of another difficulty. Prof. Ludwig 4 has pointed out
that, whereas the so-called ambulacrals of Asterids He at right angles
to the perradius and between the successive branches and podia,
those of Ophiurids lie parallel to the perradius and across the branches
to the podia. Now, if, as we have every reason to believe, the Ophi-
urids were derived from Asterids, it is hard to see why this change
should have taken place. That is the difficulty, but it is a difficulty
that disappears if we suppose that the vertebra is really compounded
of two successive ambulacral pairs in the manner outlined above.
The remaining objection to this latter ' hypothesis is that, as Dr.
Gregory has pointed out, it postulates the suppression of alternate
podia. Nevertheless, similar fusion and concomitant suppression
are not so unknown among Echinoderms as to render this objection
a fatal one.
"The suggestion that the vertebrae of Ophiurids are composed of
two successive ambulacral pairs certainly can not be proved with
1 Zeits. wiss. Zool., vol. 36, 1881, p. 181.
2 Bull. Mus. Comp. ZooL, vol. 13, No. 4, 1887.
Proc. Zool. Soc. London for 1896, 1897, p. 1032.
* Op. cit., p. 185.
REVISION OF PALEOZOIC STELLEROIDEA. 259
the evidence at our disposal. But it is doubtful whether it can
be disproved, and I therefore make it to promote a search for
further evidence pro or con.
"Two other features in the present specimen need discussion,
and may perhaps throw light on the problem just posed. The
first of these is the fact that the broadest and stoutest portion of
each arm segment appears to be the distal half (marked a in fig. 1),
whereas in the developing Ampliiura, and in the adult Protasteridse
the reverse is the case. This character is also found in Furcaster
palseozoicus as figured by Mr. Sttirtz, in Ptilonaster princeps of
J. Hall, and in Lapworthura Miltoni, as interpreted by Dr. Gregory.
(See our fig. 4 [30].) In the Epphaven fossil the feature is to some
extent accounted for by the supposed fusion of a spiniferous adam-
bulacral element with the distal wing of the ambulacral; but in
Lapworthura and Furcaster this whig, while unconnected with the
adambulacral, is still more preponderant. In some other early
Palaeozoic genera, e. g. Eugaster, J. Hall, and EopJdura, Jaekel,
the wing is neither distal nor proximal, but arises half way up the
ambulacral, so that one can not say to which of two successive seg-
ments the podium belongs; in fact, the structure is in this respect
absolutely that of an Asterid. 1 These differences call for some
explanation. Starting from the indifferent Ordovician type of
EopMura, we may suppose that in one series of genera (e. g. Protas-
teridse) the whig moved toward the mouth, while in another series
(e. g. Lapworthuridse) it moved away from the mouth. In both
series the podium ultimately became inclosed within the substance
of (apparently) a single ossicle; but, if this took place by a sub-
sequent lateral outgrowth of stereom, then in the former case the
vertebra must have inclosed a podium distal to itself, and in the
latter case a podium proximal to itself. Thus the relations of the
vertebras to the podia would be fundamentally different in the
two lines of descent. On the other hand, alternate podia may,
as suggested on a previous page, have been inclosed by the union
of successive pairs of vertebrae, and this type of structure may
be the one seen in process of development in the arm segments of
our fossil, and the one that persisted to the present day.
"The second feature worthy of attention is the considerable
space that seems to intervene between the distal wing of one seg-
ment and the proximal wing of the next. This is conspicuous in the
distal region of ray i, where, further, the depression (g in fig. 1 [32 B])
markedly resembles the podial depression (p in fig. 1), and differs
from it mainly in being rather nearer the axis of the arm. The
theory that the Ophiurids are descended from Asterids does not
i See Jaekel, Zeits. geol. Ges., vol. 55, Protok., 1904, pp. 106-113.
260 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
explain the evolution of such an interspace, and the general tendency
to shortening and approximation of the vertebrae, shown in the
history of the Ophiurids, leads one to inquire why there should
ever have been introduced a stage in which, as in the Devonian
OpJiiurina, the recent deep-sea Ophwhelus, and the young AmpTiiura,
the vertebrae have so far departed from the primitive type as to
become axially elongate. This stage has been regarded as primi-
tive by Lyman, Ludwig, and other zoologists, but in theory and
fact it is ' primitive' only so far as the later forms are concerned,
while with reference to the complete series it is 'intercalated.'
"Recurring to the hypothesis that the vertebrae are compounded
of successive pairs of ambulacrals, we find in it an explanation
not merely of this curious elongation of the arm segments in general,
but of the interspaces between them in this genus (lettered q). An
objection to the hypothesis was the necessity for assuming the sup-
pression of alternate podia. Here, it may be, is evidence of a
stage in which those podia were not yet suppressed, though perhaps
somewhat atrophied. If both depressions (p and q) were for the
reception of podia, then their alternate approximation to and re-
moval from the perradius may be compared with the similar phe-
nomenon in Asterids.
" Whether the features just discussed be or be not accepted as
evidence in favor of the compound origin of the Ophiurid vertebra,
they demand some explanation; and it may be added that the same
hypothesis will perhaps furnish an equally needed explanation for
the hitherto unexplained differences that obtain between Palaeo-
zoic genera of Ophiurids in the relations of the adambulacrals to the
ambulacrals.
"The origin of the Ophiurid mouth-skeleton also may be illu-
minated by the preceding hypothesis, which is in full harmony with
the opinion that many more arm segments enter into the composition
of that structure than is the case in Asterids, an opinion based on
embryological research by Dr. O. zur Strassen, 1 and on palaeonto-
logical investigation by Dr. O. Jaekel in the paper quoted above.
In this region of the ray, at any rate, adambulacrals and podia must
have been suppressed in all Ophiurids. We may here note the
apparent absence from our specimen of the adambulacral elements
known as side mouth-shields, as well as of the peristomial plates.
The latter are wanting in most early Ophiurids, a fact confirmatory
of Dr. zur Strassen's conclusion that they are not ambulacral elements,
but ' secondary calcifications confined to the interradial region.' '
Schondorf states that the structure of the arms is like that in
LapwortJiura miltoni and the oral armature as in Encrinaster.
i Zool. Anzeiger, vol. 24, 1901, p. 609.
REVISION OF PALEOZOIC STELLEROIDEA. 261
Genus FURCASTER Stlirtz.
Furcaster STURTZ, Palaeontographica, vol. 32, 1886, p. 79; N. Jahrb. fiir Min.,
etc., 1886, vol. 2, p. 148; Verh. naturh. Ver. preuss. Eheinl., etc., vol. 50,
1893, p. 25. GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1038;
Treat. Zool., vol. 3, Echinoderma, 1900, p. 275, fig. 31.
Disk small, circular. Rays five, short, narrow, tapering gradually,
and slightly flexible. Actinally flat and abactinally rounded. There
is a dorsal median column of slightly projecting plates and two lateral
ranges of side plates. The dorsal plates are medially depressed, with
their outer sides extended into ears, into which fit the spine-bearing
side plates.
"Ambulacral ossicles [opposite] of a long central body and two
short wings, which are attached only to the anterior corner of the
ossicle." (Gregory.) Buccal scutes small.
GenoJiolotype. F. palseozoicus Stlirtz. F. (?) daoulasensis (Davy)
may also belong here.
Remarks. "In Stiirtz's description he refers to the existence in
this genus of ventral arm plates, and even figures them. His type-
specimens are now in the British Museum, but I can not verify the
existence of any ventral arm plates." (Gregory.)
FURCASTER PALvEOZOICUS Stiirtz.
Furcaster palseozoicus STURTZ, Palaeontographica, vol. 32, 1886, p. 79, pi. 8,
figs. 4, 5; N. Jahrb. fiir Min., etc., 1886, vol. 2, p. 148; Palaeontographica,
vol. 36, 1890, p. 214. GREGORY, Proc. Zool. Soc. London for 1896, 1897,
p. 1038.
Formation and locality. In the Lower Devonic roofing slates,
Bundenbach, Germany. There are three specimens of this species
at Yale University.
Cat. No. 59383, U.S.N.M.
FURCASTER (?) DAOULASENSIS (Davy).
Protaster daoulasensis DAVY, Bull. Soc. geol. France, ser. 3, vol. 14, 1887, pp. 182-
187, text figs. GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1038,
footnote ("is an ally of Furcaster").
Formation and locality. From the Lower Devonic, Rumguen, near
Brest, France.
Genus PALASTROPECTEN Sturtz.
Palastropecten STURTZ, Palseontographica, vol. 32, 1886, p. 88, pi. 10, figs. 3, 4;
vol. 36, 1890, p. 213, pi. 26, figs. 12, 13. GREGORY, Proc. Zool. Soc. London
for 1896, 1897, p. 1038.
Palsespondylus STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 190.
Palxospondylus STURTZ, Verh. naturh. Ver. preuss. Kheinl., etc., vol. 56, 1900,
pp. 191, 202.
Diagnosis ly Gregory. "Disk circular, large (badly preserved in
the specimens; probably originally soft and irregular). Arms thick,
262 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
broad, and somewhat lanceolate in shape. Ambulacral ossicles
narrow, the lateral wings resting on the whole body of the ossicle.' 7
Genoholotype and only species. P. zitteli (Stiirtz) (citations as above) .
In the Lower Devonic, Bundenbach, Germany.
Family EOLUIDIIME Gregory.
Eoluidiidx GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1038.
Eophiuridss STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 203. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 235.
Original diagnosis. " Strep tophiuras with the ambulacra! ossicles
united to form vertebral ossicles. Ventral arm plates present, but
there are 310 buccal shields. (Dorsal arm plates present only in
the highest genus.)
"Remarks. This family includes three Devonian genera, which
differ from the previous families of this order by the presence of
ventral arm plates and by having vertebral ossicles, which articulate
( ? always) by simple rounded pits and processes. The family differs
from living Streptophiurse by the absence of buccal shields and
the simplicity of the oral armature."
Contains the genera:
Eoluidia Stiirtz.
Eospondylus Gregory.
Miospondylus Gregory.
Genus EOLUIDIA Sturtz.
Asterias asperula ROEMER (part), Palaeontographica, vol. 9, 1863, p. 146, pi. 24,
figs. 1-5; pi. 26, -fig. 6; pi. 27.
Eoluidia STURTZ, Palseontographica, vol. 32, 1886, p. 89, pi. 11, figs. 1, 2; vol. 36,
1890, p. 211, pi. 26, figs. 10, 11. GREGORY, Proc. Zool. Soc. London for
1896, 1897, p. 1038. SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London,
ser. B, vol. 202, 1912, p. 226, fig. 4E.
EopMurites STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900, p. 190.
Diagnosis. "Disk rather large; the interbrachial outlines are
deeply concave. [Kays very long, slender, and fragile.] Each
syngnath consists of pairs of mouth frames and jaws; a jaw plate
is present. The vertebral ossicles are small and the union of the
two lateral elements incomplete; the lateral wings are thin. The
adambulacral plates are triangular and each of them bears several
spines. The pores for the podia occur at the middle of the lateral
margin of the ventral arm shields. " (Gregory.)
Genoholotype and only species. E. decheni Sturtz (citations as
above). Lower Devonic, Bundenbach, Germany.
Cat. No. 35116, U.S.N.M.
REVISION OF PALEOZOIC STELLEROIDEA. 263
Genus EOSPONDYLUS Gregory.
Ophiurella STURTZ (not Agassiz, 1834), Palseontographica, vol. 32, 1886, p. 77.
Eospondylus GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1039.
Original diagnosis. "Disk circular. Ambulacral ossicles com-
pletely fused into vertebral ossicles, each of which, however, is trav-
ersed by a pore. The adambulacral ossicles are somewhat pear-
shaped. The podial pores are at the posterior angles of the ventral
arm plates."
Genoholotype and only species. Ophiurella primigenia Stiirtz.
EOSPONDYLUS PRIMIGENIUS (Stiirtz).
Ophiurella primigenia STURTZ, Palseontographica, vol. 32, 1886, p. 77, pi. 8,
figs. 1, 2; vol. 36, 1890, p. 210, pi. 26, figs. 6, 7.
Eospondylus pnmigenia GREGORY, Proc. Zool. Soc. London for 1896, 1897,
p. 1039.
Formation and locality. Lower Devonic, Bundenbach, Germany.
Genus MIOSPONDYLUS Gregory.
Ophiura rJienana STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 50, 1893,
p. 29, pi. 1, figs. 1-3.
Miospondylus rhenanus GREGORY, Proc. Zool. Soc. London for 1896, 1897, p.
1039. SOLLAS and SOLLAS, Philos. Trans. Koy. Soc. London, ser. B, vol.
202, 1912, p. 226.
Onginal diagnosis (Gregory). "Disk circular. Ambulacral ossi-
cles completely united; each half of the vertebral ossicle is boot-
shaped. The oral angles each consist of a pair of syngnaths without
jaw plate. The ventral arm plates are small and are not notched
by podial pores."
Genoholotype and only species. Opliiura rhenana Stiirtz (citations
as above). Lower Devonic, Bundenbach, Germany.
Remarks. Eospondylus and Miospondylus " agree in family char-
acters with Eoluidia, but differ from it in the structure of both the
ambulacral and adambulacral plates." (Gregory.)
Family AGANASTERID.^ Stiirtz.
Aganasteridse STURTZ, Verb, naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900,
p. 203. SCHONDORP, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 63,
1910, p. 235.
Genus AGANASTER Miller and Gurley.
Aganaster MILLER and GURLEY, Sixteenth Rep. Geol. Surv. Indiana, 1891, p.
372; authors' extracts, 1890, p. 57. GREGORY, Proc. Zool. Soc. London for
1896, 1897, p. 1039.
^ Ophiopege BOHM, Zeit. geol. Gesell., vol. 45, 1893, p. 159 (same genoholotype as
for Aganaster).
Original description. "In 1869 Meek and Worthen described an
Ophiuroidea, in the Proceedings of the Academy of Science of
264 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
Philadelphia, page 169, under the name of Protaster gregarius, which
they redefined and illustrated in the Geological Survey of Illinois,
volume 5, page 509, under the name of Protaster ( ?) gregarius. They
had numerous specimens 'in the condition of casts and molds, in
a very fine, somewhat granular matrix, that did not show the details
of its structure very clearly/ but they said, 'It will probably be
found to be generically distinct from the Silurian typical forms of
Protaster, but we prefer to place it provisionally in that genus for the
present.' In the collection of Mr. Gurley there are several specimens
belonging to this species, and they show a few characters not ob-
served by Meek and Worthen, and demonstrate very clearly this
species does not belong to the genus Protaster; we therefore propose
to include this species in a new genus, Aganaster, and describe the
characters, so far as known, as follows:
" General outline, a central circular disk with five long, narrow
rays ; the circular disk on the dorsal side is covered with small polygo-
nal plates which are not interrupted by the presence of the rays,
thus showing the disk had a depth greater than the depth of the
rays; rays very narrow and convex or half cylindrical, spine bearing,
gradually tapering, and from the dorsal side appear as if composed
of plates arranged exactly opposite each other; the ventral side shows
a rather deep central disk with marginal plates. There are 10 oral
plates in the central part of the disk."
Genoholotype and only species. Protaster gregarius Meek and
Worthen.
Remarks. Aganaster "has nothing to do with Protaster and is
clearly a member of the Streptophiurse. As far as its characters are
known to me it must be included among the Eoluidiidae. It differs
from the rest of this family by the presence of dorsal arm plates.
* * * Bohm has founded the genus OpJiiopege on the type species
of Aganaster." (Gregory.)
AGANASTER GREGARIUS (Meek and Worthen).
Protaster f gregarius MEEK and WORTHEN, Proc. Acad. Nat. Sci. Philadelphia,
vol. 21, 1869, p. 169; Geol. Surv. Illinois, vol. 5, 1873, p. 509, pi. 16, fig. 5.
Alepidaster gregarius MEEK, Amer. Journ. Sci., ser. 3, vol. 4, 1872, p. 275 (gen.
ref. only).
Aganaster gregarius MILLER and GURLEY, Sixteenth Rep. Geol. Surv. Indiana,
1891, p. 372, pi. 9, figs. 10, 11; authors' extracts, 1890, p. 57, pi. 9, figs. 10,
11. MILLER, N. Amer. Geol. Pal., 1st. App., 1892, p. 673, fig. 1208.
Description of 1873. "The disk of this species is circular in out-
line, slightly convex above, and measures from 0.20 to 0.30 inch in
diameter. In most cases it looks as if merely covered by a smooth,
membranaceous integument. Some casts of its external surface,
however, seem to show traces of flat, nearly smooth, imbricating
scales above. The five arms are slender, flexible, and rather long
KEVISION OF PALEOZOIC STELLEROIDEA. 265
in proportion to breadth. In a specimen with a disk measuring
0.25 inch in breadth, the diameter of the arms near the disk is only
0.05 inch. None of the specimens show the entire length of the arms,
though some fragments of them were seen lying detached in the
matrix, about 0.55 inch in length, without being complete at either
end. From the breadth and gradual taper of these, it would seem
probable that when entire they may have been 0.75 to 1 inch in length.
Their impressions in the matrix give no indications of a longitudinal
furrow along the under side, but show that there were about six
pairs of arm pieces in a length of 0.16 inch. These pieces appear
to be nearly though not exactly opposite, and each one provided
below with a comparatively large, round, deep pit, or pore, near
the middle of its anterior side. Along their lateral margins there
appear to be impressions in the matrix of very small spines (one to
each arm piece), though if such, they must have been extremely
short. Impressions of the upper side of the slender arms show
them to have been somewhat rounded above, with the nearly square
arm pieces slightly alternating. Some of the impressions seem to
show traces of central pores or pits, one at the middle of each pair
of pieces, though in others no traces of these are visible/ 7
Formation and locality. Common as molds in a fine impure
sandstone and in the famous crinid bed at Crawfordsville, Indiana,
in the Keokuk formation of the Mississippic.
There are two specimens (No. 6) of this species in the Wachsmuth
collection at Harvard University. At Yale University there are
four specimens.
Family CHOLASTERID^ Worthen and Miller.
Cholasteridse WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883, p. 328.
Original deHnition. Cholaster "is so widely different from those
heretofore discovered that we think it belongs to a distinct family,
which might very appropriately be designated the Cholasteridse,
but until other specimens have been found showing other parts
of the body, we prefer to let the family remain undefined."
Contains the genus :
Cholaster Worthen and Miller.
Genus CHOLASTER Worthen and Miller.
Cholaster WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883, p. 328.
GREGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1040.
Original description. Dorsal side alone known. "Body trun-
cated pentagonal, central area circular, deep and large in propor-
tion to the rays; rays distant, smaU, short and abruptly truncated.
266 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
" Dorsal side possessing a rather large centro-dorsal plate, which
is surrounded by five plates occupying the position of radials, while
the other part is covered with numerous small disk plates. "
Gfenoholotype and only species.- C. peculiaris Worthen and Miller.
Gregory states that CJiolaster "appears to be allied to Aganaster,
but the structure of the ambulacral ossicles is unknown."
CHOLASTER PECULIARIS Worthen and Miller.
Cholaster peculiaris WORTHEN and MILLER, Geol. Surv. Illinois, vol. 7, 1883,
p. 329, pi. 31, figs. 4a, 4b.
Original description. "The general form of the body is that of a
truncated pentagonal star, with a large circular disk. A centro-
dorsal plate is surrounded by five others, all of which are much larger
than other plates of the body, and occupy a centro-dorsal depres-
sion. The position of these five plates is that of radials, resting
upon a basal, and therefore all other plates of the body might be
regarded as radials and interradials, the latter being very numer-
ous, because of the wide separation of the rays at their junction
with the body. The interradials are polygonal, convex, or sub-
spinous plates, without much variation in size whether filling the
larger or smaller interradial spaces. The plates following the larger
radial plates, as above defined, in the direction of the rays, are not,
however, by their form or arrangement to be distinguished from
the interradials, though differing from those forming the rays.
The plates covering the dorsal side are thus divided into three
kinds, viz.: First, the larger plates filling the centro-dorsal depres-
sion; second, the disk plates which cover all other parts of the body
except the rays, and, third, the plates which cover the rays proper,
which in this species are easily distinguished from the central disk
plates.
"The rays are short, abruptly truncated, and slightly expanded
at the apices by reason of an enlargement of the terminal plates.
They are widely separated from each other, though not at uniform
distances, and present the appearance of having been stuck on the
central disk, instead of having grown from it, an appearance more
marked, by reason of the change in the form of the plates, from the
disk to the rays.
"One of the interradial spaces is much greater than the others,
so that a line may be drawn across the disk, leaving three entire
rays upon the smaller half. The back of each ray is covered by a
series of transversely elongated plates separated from the side
series by a longitudinally impressed line. A single series of plates
covers either side of a ray, interlocking with the transverse dorsal
series and directed backward toward the disk.
"The oral plates are unknown, and no madreporiform tubercle
has been detected."
REVISION OF PALEOZOIC STELLEBOIDEA,
267
Formation and locality. Okaw Bluffs, between Chester and
Kaskaskia, Illinois, in the Chester formation of the Upper Missis-
sippic. The holotype is in the Illinois State collection, No. 2480.
Subclass OPHIUROIDEA.
These animals are not present in the older Paleozoic, may have
appeared in the late Devonic, and <Jo not seem to have been abundant
FIG. 33. OPHIOTERESIS. AFTER BELL FROM GREGORY.
ABORAL SURFACE OF AN ARM OSSICLE: a, ARTICULAR
CAVITIES; d, THE DOUBLE DORSAL SHIELDS; I, LATERAL
ARM PLATES.
FIG. 34. SYNGNATHS OF OPHIURA CIUARIS.
AFTER MULLER FROM GREGORY. ;, JAW;
m.f., MOUTHFRAME; n, g., GROOVE FOR CIR-
CUMCESOPHAGEAL NERVE-RING; p. d., PORE
AND DEPRESSION FOR ORAL TENTACLE.
before the Triassic, since which time they occur more and more com-
monly. In the present oceanic waters they are popularly known as
sand-stars, brittle-stars, branching-stars, or basket starfish. They
range from shallow and estuarine waters to abyssal depths. Typical
Ophiuroidea differ from typical Asteroidea in having the arms sharply
marked off from the disk as appendages, and in the absence of
grooves along the actinal side of the arms. This means that the body
cavity which in the Asteroidea extends out into the rays is restricted
in the Ophiuroidea to the disk.
The subclass Ophiuroidea is defined by Schondorf as follows:
"Ainbulacral water-vascular system situated in a small groove
at the base of the ray ossicles, and ventrally covered by a single
column of ventral shields. From the radial canal outside of the
ambulacrals arise simple lateral branches that never have ampullae,
as a rule curve upward, pass into and through the substance of the
ossicles, and finally open out laterally between the ventral and side
shields as the ambulacral podia. Ambulacrals opposite, each right
and left piece coossified into a vertebra with complicated articular
surfaces [see figs. 34-36]. Adambulacrals transformed into lateral
shields. Vertebrae dorsally covered by a single column of dorsal
shields." Disk circular in outline, lt without marginal plates, and
sharply separated from the rays, that as a rule are rounded. There
is no typical madreporic plate. One of the ventrally situated mouth
shields serves as madreporite" (1910a: 246).
268 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
In ophiurids the oral skeleton, or syngnaths, is composed of three
pairs of adambulacrals and ambulacrals. Of these the two first
pairs alone are prominent and preserved. The third pair remains
internal, small, and rudimentary.
Family ONYCHASTERID^ Miller.
Onychasteridae MILLER, N. Amer. Geol. Pal., 1889, p. 216. GREGORY, Proc. Zool.
Soc. London for 1896, 1897, p. 1040. STURTZ, Verh. iiaturh. Ver. preuss.
Kheinl., etc., vol. 56, 1900, p. 202. SCHONDORF, Jahrb. nassauisch. Ver.
Naturk., Wiesbaden, vol. 63, 1910, p. 235.
Original diagnosis. " Strep tophiurse with well-developed vertebral
ossicles, and with very flexible, contorted, unbranched arms; there
U.S. "
u. ^
FIGS. 35 AND 36. VERTEBRAL OSSICLE OF OPHIURA CILIARIS. AFTER MULLER FROM GREGORY. 35;
ABORAL SURFACE; 36, ADORAL SURFACE, c, CANAL FOR THE PODION (SHOWN BY REMOVAL OF A PART OF
MUSCLE FIELD ON RIGHT SIDE); J.W., LOWER MUSCLE FIELD; l.f., LOWER AND ./., UPPER CANAL FURROWS,
., UMBO; U. *., UMBONAL SOCKET.
are no external arm plates, the integument containing granules
only.' 7
Contains the genus :
Ony chaster Meek and Worthen.
Genus ONYCHASTER Meek and Worthen.
Onychaster MEEK and WORTHEN, Geol. Surv. Illinois, vol. 3, 1868, p. 526; Proc.
Acad. Nat. Sci. Philadelphia, vol. 21, 1869, p. 82; Geol. Surv. Illinois, vol. 5,
1873, p. 474. ZITTEL, Handb. Pal., vol. 1, 1879, p. 443. MILLER, N. Amer.
Geol. Pal., 1889, p. 264. STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc.,
vol. 50, 1893, p. 30. GREGORY, Proc. Zool. Soc. London for 1896, 1897, p.
1040. STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900, p. 183.
SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909,
pp. 47-61, vol. 63, 1910, p. 240; Palaeontographica, vol. 57, 1910, p. 59.
SPENCER, Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914,
pp. 26, 50. I. B. J. SOLLAS, Philos. Trans. Hoy. Soc. London, ser. B, vol.
204, 1913, p. 51. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden,
vol. 66, 1913, pp. 97-114.
REVISION OF PALEOZOIC STELLEEOIDEA.
269
Genoholotype. 0. flexilis Meek and Worthen.
For generic description, see under 0. flexilis.
Remarks ly Gregory.- "This interesting genus has hitherto been
placed among the Euryalidse, of which it has been regarded as the best
known fossil representative. As Prof. Bell, however, has remarked,
Meek and Worthen's clear figures of the vertebral ossicles show that
the articular surfaces are Strep tospondy line and not Cladophiuroid."
Schondorf in 1909 restudied Ony chaster hi detail, and came to the
following conclusions :
u Ony chaster flexilis shows plainly in the structure of its arm ossicles
that it belongs to the Ophiuroidea. As in this subclass, so also in
Onychaster the arm ossicles originated from two halves (ambulacrals)
that grew tightly together. When compared with living Ophiurids, this
40
41
FIGS. 37-41 .ONYCHASTER FLEXILIS. AFTER SCHONDORF. 37 AND 38, PROXIMAL VERTEBRA SEEN FROM
THE ABORAL AND ADORAL SURFACES. 39, VENTRAL VIEW OF TWO PROXIMAL VERTEBRA, ORIENTED
WITH THE ABORAL SIDE UP AND THE ADORAL DOWN. 40, SIDE VIEW OF SAME. ADORAL SIDE ON LEFT,
ABORAL ON RIGHT. 41, VENTRAL VIEW OF SEVERAL SUCCESSIVE VERTEBRA WITH THE ABORAL SIDE
AT THE TOP. O, UPPER LATERAL ARTICULAR KNOB; O', SOCKET FOR KNOB OF ADORAL SIDE; C, INSERTION
FIELD FOR THE UPPER, AND 0, FOR THE LOWER INTERMEDIARY VERTEBRAL MUSCLES; gr, MEDIAN UN-
PAIRED DEPRESSION; gr\, SMALL DEPRESSION FOR THE KNOB, fci, OF THE ADORAL SIDE; gr, LARGE DE-
PRESSION OF THE CENTRAL AREA OF VERTEBRA; TO, BARRIER SEPARATING THE UPPER AND LOWER
INTERMEDIARY VERTEBRAL MUSCLES; T, LATERAL GROOVE (? FOR LATERAL BRANCHES OF WATER-
VASCULAR SYSTEM); S, VENTRAL PROJECTION OF THE LATERAL KNOBS OF ADORAL SIDE; t AND t' , UNPAIRED
MEDIAN ARTICULAR KNOB AND SOCKET.
amalgamation of the two arm ossicles in Onychaster is still plainly to be
seen. The equivalent skeleton of the adambulacrals in the asterids is
not yet completely separated from the arm ossicles nor developed into
independent side or lateral shields ; on the contrary, they still remain
attached and in articulation with the arm ossicles on their ventral outer
margin. On their somewhat fluted outer margin they bear a few spines.
The arm ossicles are not externally naked but are covered by smaller
additional plates that bear flat calcareous particles. Individual large
dorsal shields do not appear to be present, for on the large Berlin speci-
men the arm ossicles lie directly beneath the small flat dorsal plates.
Only in the distal part of the rays does one observe some rows of
270 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
apparently independent dorsal plates, that overlie one another in
a scalelike manner, but this determination needs to be substantiated
on better preserved material. Along the median line of the arm
ossicles there lies ventrally a single column of ventral plates, that
bear spines and externally cover the radial canal of the water vas-
cular system. The lateral branches from the radial canal to the
ambulacral podia do not penetrate the substance of the arm ossicles
as in living ophiurids, but lie between each pair of vertebrae. The
oral skeleton consists of five interradially placed mouth-corner
pieces. Each one of these is made up of three elements, the two
outer of which are united into pairs" (pp. 60-61).
Contains the following species :
0. flexilis Meek and Worthen. Keokuk.
0. asper Miller. Keokuk.
0. barrisi (Hall). Burlington.
0. confragosus Miller. Keokuk.
0. demissus Miller. Keokuk.
ONYCHASTER FLEXTLIS Meek and Worthen.
Text figs. 37 to 42.
Onychastei* flexilis MEEK and WORTHEN, Geol. Surv. Illinois, vol. 3, 1868, p. 526,
figs. A-D; Proc. Acad. Nat. Sci. Philadelphia, vol. 21, 1869, p. 83; Geol. Surv.
Illinois, vol. 5, 1873, p. 510, pi. 16, figs. 3a-3Z. ZITTEL, Handb. Pal., vol. 1,
1879, p. 444, fig. 314. MILLER, N. Amer. Geol. Pal., 1889, p. 264, fig. 374.
SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 62, 1909,
pp. 47-61; vol. 63, 1910, p. 240. I. B. J. SOLLAS, Philos. Trans. Roy. Soc.
London, ser. B, vol. 204, 1913, pp. 51-62, text figs. 1-4, pi. 8, figs. 1-6.
SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wiesbaden, vol. 66, 1913, pp.
97-114, text figs. 1, 2, pi. 3, figs. 3-9. SPENCER, Mon. Brit. Pal. Asterozoa,
pt. 1 (Palseontogr. Soc. for 1913), 1914, pp. 26, 50.
Original description. "The interesting fossil on which we propose
to found this genus and species seems to differ so widely, in some of
its characters, from the true starfishes, as well as from the Ophi-
urians, as to leave doubts whether it can be properly placed in either
of these groups as now understood. * * * In habit and general
appearance it most nearly resembles the Ophiurians, from which,
however, it differs widely in structure. It is composed of a rather
small subdiscoid body, and five long, slender, rounded, flexible arms
or rays. In nearly all of the specimens yet found the arms are folded
together like the claws of a bird when grasping some small object.
A few of them, however, have the arms opened out more or less, so
as to show that they were very flexible, or capable of being moved
about in all directions. They usually increase slightly in thickness
for a short distance from the body, then taper very gradually to their
extremities, being about 2.50 inches in length, and 0.22 inch in
breadth, at the widest part.
KE VISION OF PALEOZOIC STELLEKOIDEA. 271
"On the dorsal side of the body * * * there is seen a compar-
atively large circular area or disk, composed of an outer circle of ten
rather prominent pieces, united together in five pairs by close-fitting
sutures, each piece being pierced by a round ovarian ? pore. Imme-
diately within this circle there is, apparently, another circle of ten
smaller pieces, also united in five pairs, but without pores; and within
this latter circle there is a third range of five still smaller, nonporif-
erous pieces, surrounding a central anal? opening; the whole re-
minding one of the apical disk of an Echinoid, though differing in
structure from this part of the known types of that group. It is
also worthy of note that there is some analogy between this disk and
the body of a crinoid, excepting that there is a central opening, and
thnt the first division of the radial series takes place immediately on
the inner range of pieces corresponding to the basal pieces of a crinoid,
while all of the third range of pieces are pierced by pores. * * *
"Immediately outside of the circle of ten pore pieces, mentioned
above, each pair of these pieces is succeeded by two or three pairs of
differently formed, interlocking, transverse pieces, in direct range, con-
necting them with the dorsal side of each of the five rays. A little far-
ther out the dorsal side of the rays, these transverse pieces are seen
to become disconnected by more or less wide spaces, and gradually
pass into pairs of lanceolate pieces, deeply furrowed longitudinally,
while between the inner ends of the two pieces of each pair there
appears to be a porelike opening. These latter disconnected pieces
continue all the way out to the extremities of the rays, and, with
numerous smaller intervening ossicles, form together, as it were, the
skeleton or framework of the long flexible rays. It is only, however,
when an outer granular integument has been removed that this
skeleton structure can be seen. In some parts of some of our specimens
this outer granular covering remains, and is seen to be composed of
numerous small, rounded, rather prominent ossicles, regularly
arranged in qui[n]cunx, so as to give the surface a chagreenlike rough-
ness. These ossicles were doubtless attached to, and secreted by, a
soft dermal envelope, covering the whole surface, while the larger
pieces within formed the frame, as it were, of the whole structure,
and probably furnished points of attachment for the muscles that
moved the rays.
"None of our specimens show clearly the inner side or ambulacral
furrows of the rays, nor the under side of the body consequently
we know nothing of the nature or position of the mouth or of the
ambulacra. In several instances, however, we have seen the remains
of one or more rows of small, short, longitudinally striated spines
along the inner side of the arms.
50601 Bull. 8815 18
272 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
"The entire breadth of a mature individual, across between the
extremities of the rays on opposite sides, if these rays were straight-
ened out, would be about 5 to 6 inches."
Formation and locality. This species occurs quite commonly in
the famous crinid bed at Crawfordsville, Indiana, in the Keokuk
formation of the Mississippic. Specimens are in several American
museums; two are at Yale University, others in the United States
National Museum.
Cat. No. 59392, U.S.N.M.
ONYCHASTER ASPER Miller.
Onychaster asper MILLER,. Seventeenth Rep. Geol. Surv. Indiana, 1892, p. 684,
pi. 12, figs. 3-5; advance extras, 1891, p. 74, pi. 12, figs. 3-5. MILLER, N.
Amer. Geol. Pal., App. 1, 1892, p. 680, fig. 1240; App. 2, 1897, p. 749, fig*.
1366. KEYES, Missouri Geol. Surv., vol. 4, 1894, p. 131.
Original description. " Dorsal side covered with an integument of
small plates and numerous short spines. Central disk rather large,
circular, convex, inflated from the point of contact with the arms.
The outer integument covers the whole surface of the central disk,
leaving no orifice exposed. Where the outer integument is worn off
the disk is composed of rather large, polygonal spine bearing plates.
The spines do not arise from the center of the plates, but laterally
from little pits or sockets at the sutures. These sockets give the
plates a somewhat sculptured appearance. The spines have a bulb
at the base and taper to an obtuse point above.
"The arms are long, rounded on the dorsal side, and very flexible.
Figure 4 shows the arms abruptly folded from the middle over the
ventral part, while figures 3 and 5 have the arms folded like the claws
of a bird grasping some small object. There is a row of spines on each
side of the arm furrows.
"This species is so different from the type that the generic reference
is very doubtful."
Formation and locality. In the Keokuk formation of the Missis-
sippic, at Boonville, Missouri. The specimens are said to be in the
Miller and F. A. Sampson collections.
ONYCHASTER BARRISI (Hall).
Protester f barrisi HALL, Desc. N. Sp. Crinoidea, 1861, p. 18.
Onychaster ? barrisi MEEK and WORTHEN, Proc. Acad. Nat. Sci. Philadelphia,
vol. 21, 1869, p. 83; Geol. Surv. Illinois, vol. 5, 1873, p. 476, pi. 10, figs. la-Id
Hall's original description. " A fragment of an Asterias } among the
Burlington fossils, presents, in the rays and in the oral plates, some
characters in common with Protaster; but I can discover no evidence
of a central disk. The fragment preserves the center of the lower
side and parts of four rays. All that remain of the rays are two
distinct ranges of plates, which, near the base, are separated by a
REVISION OF PALEOZOIC STELLEROIDEA. 273
wide groove, but farther on appear to come in contact on their inner
margins. These plates consist of joints closely articulated together,
with a longitudinal foramen: their lower sides are marked lengthwise
by a comparatively wide but not deep groove. The body of the plate
has, on each side, a lateral arching process which is jointed at the
two extremities, and separated in the center by a pore-like perfo-
ration. There are ten oral plates, two from each division of the ray:
these plates are expanded vertically; their extreme points have the
inner edges slightly curving; the lower external faces are slightly
indented, or crenulate ; the surface of attachment is wide and strong,
and constricted at the base by a distinct groove, beyond which it
again expands.
"This form, if really without a disk, differs essentially from Prot-
aster; and there are also other differences, which may make it
necessary to constitute a distinct genus when better specimens shall
be obtained."
Formation and locality. In the Burlington limestone at Burlington,
Iowa. There are three specimens of this species in the Wachsmuth
collection (Nos. 10, 11, and 12) at Harvard University.
ONYCHASTER CONFRAGOSUS Miller.
Onychaster confragosus MILLER, Seventeenth Rep. Geol. Surv. Indiana, 1892,
p. 684, pi. 12, figs. 6, 7; advance extras, 1891, p. 74, pi. 12, figs. 6, 7.
Original description. "This species is so different from the one ^ast
described [0. asper], and also from the type of this genus, that it may
not be congeneric with either of them. I have no doubt that it
belongs to the same family. I have seen only the dorsal side of the
disk and the dorsal and lateral sides of the rays. There is no such
integument of small plates covering the dorsal side as belongs to
0. asper, but the surface is covered with short spines.
"The central part of the disk is somewhat injured in our specimens,
and its character can not be definitely determined. It is surrounded
by large, centrally convex, six-rayed plates. These rays seem to be
the elevated ridges that separate the sockets for the insertion of the
spines. Outside of this circle of ten six-rayed plates, there is a circle
of quadrilateral plates more or less sculptured by the sockets for the
spines, and here the radials may be said to commence. The dorsal
side of each ray consists of three series of plates, and there is one
series upon each side, or five series in an arm. The first plates are
connected laterally by smaller ones, in the angular depressions,
between the commencement of the rays, which form part of the disk*
All of the radial plates are more or less sculptured by the depressions
for the insertion of the spines. The spines are longer than the diam-
eter of a plate, bulbous at the lower end, and taper to an obtuse point.
The arms are long, round on the dorsal side, flexible and capable of
being rolled up on the ventral side or being twisted laterally. No
274 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
such ovarian pores piercing the plates of the disk, as described and
illustrated in O.flexilis, have been discovered."
Formation and locality. In the Keokuk formation, at Boonville,
Missouri. The cotypes are said to be in the Miller collection.
ONYCHASTER DEMISSUS Miller.
Ony chaster demissus MILLER, Seventeenth Rep. Geol. Surv. Indiana, 1892, p. 685,
pi. 12, figs. 8-10; advance extras, 1891, p. 75, pi. 12, figs. 8-10; N. Amer.
Geol. Pal., App. 1, 1892, p. 680, fig. 1241.
Original description. "This species hangs its arms down and folds
them like the claws of a bird grasping some small object, and in these
respects is more like 0. flexilis than either of the preceding species.
The central disk is slightly concave, subpentagonal in outline, and
the rays drop down at right angles to the circumference of the disk.
"In the center of the disk there is a low, subcircular elevation, in
which I have been unable to find any sutures, or to determine whether
or not there is an opening of any kind ; it appears to consist of a single
plate. It is surrounded by a series of ten plates that form the sub-
pentagonal rim of the central disk. These plates are large, very
convex, radiately sculptured, and bent down in the direction of the
radial series as well as curving in to unite with the central plate. The
sculpturing is due to the sockets for the insertion of the rays. The
radial series commence from this circle of plates. The rays are
angular on the dorsal side or obtusely rounded. The dorsal side of
each ray consists of three series of plates, and there is one series on
each side, or five series in an arm. The first two plates in each ray
are connected laterally by smaller ones, which form part of the disk,
in the angular depressions, between the commencement of the rays.
All of the radial plates are more or less sculptured or pitted by the
depressions for the insertion of the spines.
"The arms are longer, more angular, and have rather smaller spines
than either of the preceding species. I have been unable to detect
any ovarian pores, but spine sockets very much resembling pores are
indicated in figure 8, but they occur in the sutures and are readily
distinguished from pores that pierce the plates."
Formation and locality. In the Keokuk formation, at Boonville,
Missouri. The three cotypes are said to be in the Miller collection.
There are two good specimens (No. 10994) in the Gurley collection at
the University of Chicago, and another specimen, from Crawfords-
ville, Indiana, in Yale University.
FORMS WHOSE RELATIONSHIPS ARE UNKNOWN.
CRIBELLITES CARBONARIUS Tate.
Cribellites carbonarius TATE, Rept. Brit. Assoc. Adv. Sci., for 1863, 1864, Notices
and abstracts, p. 88.
Original description. "This Asteroid, the first recorded from the
Mountain Limestone, is an impression of the upper surface, in a
REVISION OF PALEOZOIC STELLEROIDEA. 275
fine-grained micaceous sandstone. It is named CribeUites carbo-
narius; and the following characters are observable: Rays five,
rounded, lanceolate, five times as long as the disk, ridged in the center,
covered with longitudinal rows of reticulating tubercles; disk small
and tuberculated. The disk is only 0.3 of an inch in diameter,
while the rays are 1.5 inches in length. A circular impression in the
disk may be the impression of the Madreporiform nucleus. In the
form of this Asteroid, and in the characters observable, it is similar
to Oribella rosea, Muller; but the rays are proportionally longer, the
disk smaller, and the tubercles much nearer to each other than in
the recent analogue. The sandstone from which the fossil Sea-star
was obtained lies 20 feet above the Shilbottle coal, and about 600
feet below the base of the millstone grit, being in the upper part of
the Mountain Limestone formation, which, in Northumberland, is
about 3,000 feet in thickness. In this sandstone there also occur
Strophomena crenistria and the remains of carboniferous plants."
Order OPHIOCISTIA Sollas.
Ophiocistia SOLLAS, Quart. Journ. Geol. Soe. London, vol. 55, 1899, pp. 692, 700.
SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. London, ser. B, vol. 202,
1912, pp. 214, 222.
An anomalous order of free Echinoderma.
"The Ophiocistia are Ophiuroidea with five paired series of append-
ages, proceeding from the ventral surface of a plated test; and in
which vertebral ossicles are absent or insignificant" (p. 700).
Contains but one family, the Eucladiidse.
Remarks. That Eudadia and Euthemon, the genera belonging to
this order, are ophiurids is said to be apparent from the following:
"The absence of any openings on the dorsal surface, and of any indi-
cation of an anus, the ventral position of the madreporite, and the
sharp distinction of the arms from the test." On the other hand,
they differ from all known ophiurids in several important particu-
lars:
"The structure and disposition of the arms is unlike anything
known among either the Ophiurse or the Euryalse, and finds no
parallel among any group of fossil Ophiuroidea. If we consider the
disposition of the arms first, we find as a constant character among the
rest of the Ophiuroidea the extension of five of these appendages over
the ventral surface of the disk as far as the buccal aperture; the
vertebral ossicles of the arms are also serially represented in the
buccal armature. In the Euciadiidae also the arms are given off
from the- ventral surface of the test, and the first pair have their
origin in the outer distal angle of the jaws; if, however, we are to
regard the serial arms of Eucladiidse as the free extremities of lateral
276 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
branches given off in pairs from a median hypothetical arm, then we
must admit that the branching takes place within the test to an extent
otherwise unknown within the class. If we turn next to the structure
of the arms, we find no less striking peculiarities. No decisive
evidence exists to prove that vertebral ossicles are present, but if
they are they must be out of all proportion small compared with the
lumen of the arm. Since the cavity of the arm is almost entirely
unoccupied by skeletal structures, the question naturally arises as
to the nature of the soft parts which it contained. The distal arms
are so large that they might well have afforded room for extensions
of the digestive viscera from the test. The absence of visible aper-
tures in the arms is another very puzzling feature, and one is almost
tempted to inquire whether the paired appendages are to be compared
with arms at all.
"The nature of the buccal armature is very different from that
of any other Ophiuroid, though there is a certain amount of corre-
spondence in the paired structure of its five pieces. The absence of
any opening that could be taken for bursal apertures is noteworthy."
Family EUCLADIID^E Gregory.
Eudadiidse GEEGORY, Proc. Zool. Soc. London for 1896, 1897, p. 1040. SOLLAS,
Quart. Journ. Geol. Soc. London, vol. 55, 1899, p. 692. STURTZ, Verh.
naturh. Ver. preuss. Rheinl., etc., vol. 56, 1900, p. 204. SCHONDORF, Jahrb.
nassauisch. Ver. Naturk., Wiesbaden, vol. 63, 1910, p. 236.
Contains the genera:
Eudadia Woodward.
Euthemon Sollas.
Genus EUCLADIA Woodward.
Plates 37, 38.
Eudadia johnsoniH. WOODWARD, Geol. Mag., vol. 6, 1869, p. 241, pi. 8. ZITTEL,
Handb. Pal., vol. 1, 1879, p. 443. STURTZ, Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, 1893, p. 30. GREGORY, Proc. Zool. Soc. London for
1896, 1897, p. 1040, fig. 6 on p. 1041. SOLLAS, Quart. Journ. Geol. Soc. Lon-
don, vol. 55, 1899, p. 692. STURTZ, Verh. naturh. Ver. preuss. Rheinl., etc.,
vol. 56, 1900, p. 183. SCHONDORF, Jahrb. nassauisch. Ver. Naturk., Wies-
baden, vol. 62, 1909, p. 47; vol. 63, 1910, p. 240. SOLLAS and SOLLAS, Philos.
Trans. Roy. Soc. London, ser. B, vol. 202, 1912, pp. 214, 222. SPENCER,
Mon. Brit. Pal. Asterozoa, pt. 1 (Palseontogr. Soc. for 1913), 1914, p. 50.
This genus has seven pairs of arms in each radius, or 35 in all.
These are covered with finely scaly integument.
Genoholotype. E. johnsoni Woodward (citations as above), Lower
Ludlow formation, at Sedgley, near Dudley, England. Other species
are E. woodwardi Sollas, also from the Lower Ludlow, at Leintwar-
dine, England, and E. f leecheri, new species, from the Lower Devonic
Coeymans limestone, New York.
REVISION OF PALEOZOIC STELLEEOIDEA. 277
EUCLADIA WOODWARDI Sollas.
Plate 37, figs. 1, 2.
Eucladia woodwardi SOLLAS, Quart. Journ. Geol. Soc. London, vol. 55, 1899, p.
695, figs. 1 and 2 on p. 694.
Original description. Ten specimens " agree in presenting five
paired series of appendages, proceeding from the ventral surface
of the body, which now possesses a more or less oval outline. The
specimens are all of nearly the same size, the central body measur-
ing about 2 by 3 cm., the longest arms 2.5 cm. in length, and about
3 mm. in breadth where broadest. Not more than four, possibly
only three, pairs of arms can be traced in connection with each
radius, but an additional pair may have existed close to the buccal
armature, and have since become crushed out of recognition.
" TTie dorsal surface [fig. 1]. This is completely covered by nu-
merous rounded polygonal or irregular scales, about 0.2 mm. thick
and not exceeding 5 mm. in diameter; their surface is richly granu-
lated. They are not arranged according to any discoverable law,
though there may be a tendency to run parallel with the ambitus.
In their present state they overlap each other to such an extent
that one plate may be half concealed by another; no doubt they
were imbricated during life, but the excessive overlap now presented
is probably due in part to crushing. The direction of the imbrica-
tion is upward, that is, in the direction opposite to that of tiles on
a roof, and thus resembles the imbrication of the dorsal surface of a
recent Ophiuroid. The plates are all of the same nature, and there
are no openings on the dorsal surface.
" The ventral surface [fig. 2]. In the center the powerful buccal
armature is a very conspicuous object. It consists of five pairs
of strong plates or ossicles, precisely similar in their form and
arrangement to those of Eiicladia johnsoni. Around the armature
are numerous small plates, irregularly disposed; from their form
and size these may be regarded as elements of the test, though they
may possibly include remains of crushed arms. If, as judging
from analogy we might suppose, minute arms proceeded from the
outer angles of the jaws, they have since disappeared.
" Outside the irregularly scattered small plates larger ones are
seen arranged; along the five radii these are escutcheon-shaped,
imbricated, and form a single series of three. Distally, each plate
is produced into three processes, a single median and two lateral;
the angle formed by the side of the plate and each lateral process
is rounded and thickened to form one side of a circular aperture
for the passage or insertion of the arm, and the other half of the
aperture is completed by the thickened margin of a similarly exca-
278 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
vated adjacent or adradial plate. The adradial plates form a pair,
which, meet in the interradius. The arrangement is similar to that
which occurs in the genus EutTiemon, to be next described; in the
latter, however, the plates are not overlapping, but tesselated.
The overlapping of the plates is less on the ventral than on the
dorsal side, a difference which is probably due to the less amount
of displacement suffered by the ventral plates; and this, again, is
explicable on the assumption that the ventral surface was flatter
than the dorsal.
" The arms. The arms of this species do not exhibit that marked
increase in size, as their position in the series becomes more distal,
which is characteristic of Eucladia johnsoni. Their average length
is 25 mm. Near their tapering extremities the arms present on both
dorsal and ventral surface three plates, two of which are lateral
and one median, recalling the characteristic plating of an Ophiuroid
arm; but nearer the origin the number of conspicuous plates on
either surface is increased to four or even more, and smaller supple-
mental plates are inserted between them in a manner precisely
similar to that already described hi the case of Eucladia johnsoni.
The larger plates, swollen at first and becoming mucronate finally,
are produced into a short awn-like termination.
"A search, which proved unsuccessful, was made for some trace
of vertebral ossicles; had these structures been present originally,
they must either have been very small or some traces would still
be discernible. In Lapworihura, which occurs in the same rocks,
the vertebral ossicles are the most obvious elements in the brachial
skeleton.
"The distinction of Eucladia woodwardi from E. johnsoni rests
on the smaller number of arms possessed by the former and the
closer approach of these to equality in dimensions. The speci-
men selected as the 'type' is exhibited in the Oxford University
Museum, and bears a label stating where its description may be
found."
Formation and locality. In the Lower Ludlow, Leintwardine,
England.
EUCLADIA (?) BEECHERI, new species.
Plate 38, fig. 1.
As this problematic fossil is of considerable interest, it is thought
best to make it known in this work, even though the preservation
is not good. The general characters are easily seen in the specimen,
however, and much better than can be shown by photography, but
in detail very little can be made out. The fossil lies in a dense
dark limestone beneath the surface of a parting between two beds
EEVISION OF PALEOZOIC STELLEEOIDEA. 279
of stone. Originally but a few of the arms showed, and it was at
once seen by the late Professor Beecher that he had found some-
thing very different from a crinid. He bestowed a great deal of
skill and labor on the fossil later on, bringing it out to its present
relief, and in this condition it has remained since 1901, the year
of its collection. It seemed to both Professor Beecher and the writer
that it was an early form of branching or basket star, recalling the
living AstropJiyton. Recently, since reading the work of Sollas on
Eudadia, it became clear that this view could not be maintained.
The specimen seemingly shows the ventral side.
The disk is large and distinctly pentagonal, and from its five cor-
ners radiate the more or less coiled rays. Disk composed of de-
cidedly thick plates, apparently closely adjoining, but then- arrange-
ment is too much disturbed and crystalline to make out. R = ? about
13 mm., r ) =10.5 mm., B, to edge of rays averaging 22 mm.
Bays apparently completely circular in outline and covered by
an abundance of imbricating scales, of which there are about five
across the diameter of a ray. The average diameter of the rays
is between 1.5 and 2 mm. As the rays are crystalline the nature
of the internal skeleton can not be made out further than that
there are vertebrae present. Each radius appears to have 4 pairs
of arms, there being therefore 40 in all, but then* situation in the
disk and how they appear during growth cannot be made out. Three
pairs are of the same size throughout, while the fourth pair, which
lies upon the others, seems to be about half grown, and if there is no
decided distortion of the disk plates, these appeared alternately,
first on one side and then on the other. In Eudadia the 30 rays
are practically all alike, but in EutJiemon the 20 arms are equally
divided between full and half grown ones.
Formation and locality. From about the middle of the Coeymans
limestone of the Helderbergian (Lower Devonic), at Jerusalem Hill,
Litchfield, Herkimer County, New York. The holotype is hi the
Peabody Museum of Yale University.
Genus EUTHEMON Sollas.
Euthemon igerna SOLLAS, Quart. Journ. Geol. Soc. London, vol. 55, 1899, p. 696,
figs. 3 and 4 on p. 698. SOLLAS and SOLLAS, Philos. Trans. Roy. Soc. Lon-
don, ser. B, vol. 202, 1912, p. 222.
This genus is very similar to Eudadia , but differs at once in having
but 4 arms in each radius, or 20 in all; 10 of these are more than
twice as long (12 to 13 mm.) as the others (5 mm.).
GenoJiolotype and only species. E. igerna Sollas, from the Wenlock
limestone, Croft farm, Malvern, England.
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284 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
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Nat. Hist., vol. 5, pp. 34-44, pis. 1, 2. (Tseniaster elegans.)
18826. Description of three new species, and remarks upon others. Journ. Cin-
cinnati Soc. Nat. Hist., vol. 5, pp. 116-117, pi. 5. (Protaster miamiensis.)
1884. Description of a beautiful star fish and other fossils. Journ. Cincinnati Soc.
Nat. Hist., vol. 7, pp. 16-20, pi. 4. (Palseaster magnificus.)
1889. North American geology and palaeontology. First appendix, 1892; Second
appendix, 1897.
1892. Palaeontology. Seventeenth Rept. Geol. Surv. Indiana, pp. 611-705, pis.
1-20; Advance extras, 1891. (Onychaster asper, 0. confragosiis, and 0.
demissus.}
MILLER, S. A., and DYER, C. B.
1878a. Contributions to palaeontology. Journ. Cincinnati Soc. Nat. Hist., vol. 1,
pp. 24-39, pis. 1-2. (Palseaster simplex, P. spinulosus, Palseasterina
approximata, P. speciosa, Protaster flexuosus.)
18786. Contributions to palaeontology, No. 2. Privately printed. (Palseaster
dubius.)
MILLER, S. A., and GURLEY, W. F. E.
1891. Description of some new genera and species of Echinodermata from the
Coal Measures and Subcarboniferous rocks of Indiana, Missouri, and
Iowa. Sixteenth Rept. Geol. Surv. Indiana, pp. 327-373, pis. 1-10;
Advance extras, 1890. (Schcenaster legrandensis, Aganaster.)
REVISION" OF PALEOZOIC STELLEROIDEA. 285
MILLER, S. A., and GURLEY, W. F. E. Continued.
1897. New species of crinoids, cephalopods, and other Palaeozoic fossils. Bull.
Illinois State Mus. Nat. Hist., No. 12. (Palseaster wykoffi.)
MtJLLER, J.
1855. In ZEILER and WIRTGEN, Bemerkungen uber die Petrefacten der altern
devonischen Gebirge am Rheine, etc.; Verh. d. naturh. Ver. preuss.
Rheinl., etc., vol. 12, pp. 1-28, pis. l-9a. (Asterias rhenana.)
MURCHISON, R. I.
1854. Siluria.
NICHOLSON, H. A., and ETHERIDGE, R., JR.
1880. A monograph of the Silurian fossils of the Girvan District in Ayrshire,
fasc. 3. (Tetraster, T. wyville-thomsoni.}
D'ORBIGNY, A. D.
1849. Prodrome de Pal6ontologie, vol. 1. (Ccelaster americanus.}
PARKS, W. A.
1908. Notes on the ophiuran genus, Protaster, with description of a new species.
Trans. Canadian Inst., vol. 8, pp. 363-372, 1 pi. (Gives summary of
history of genus. P. whiteavesianus.)
QUENSTEDT, F. A.
1876. Petrefactenkunde Deutschlands, vol. 4, Asteriden und Encriniden.
RAYMOND, P. E.
1912a. On two new Paleozoic starfish (one of them found near Ottawa), and a new
crinoid. Ottawa Naturalist, vol. 26, pp. 77-81, pi. 5, 2 text figs.
(Palxaster f wilsoni, Schcenaster f montanus.)
19126. On the nature of the so-called "covering plates" in Protopalseaster narra-
wayi. Ottawa Naturalist, vol. 26, pp. 105-108, pi. 6.
RlNGUEBERG, E. N. S.
1886. New genera and species of fossils from the Niagara shales. Bull. Buffalo
Soc. Nat. Sci., vol. 5, pp. 5-22, pis. 1,2. (Squamaster, S. echinatus,
Protaster stellifer, Eugaster condnnus.)
ROEMER, F.
1863. Neue Asteriden und Crinoiden aus devonischem Dachschiefer von Bunden-
bach bei Birkenfeld. Palaeontographica, vol. 9, pp. 143-152, pis. 23-
29. (Aspidosoma tischbeinianum, Asterias asperula, A. spinosissima,
Helianthaster, H. rhenanus.)
RUEDEMANN, R.
1912. The Lower Siluric shales of the Mohawk valley. Bull. N. Y. State Mus.,
No. 162. (Txniaster schoharix.)
SALTER, J. W.
1857. On some new Palaeozoic star-fishes. Ann. Mag. Nat. Hist., ser. 2, vol. 20,
pp. 321-334, pi. 9. (Palseocoma, P. marstoni, P. colvini, P. cygnipes,
Bdellacoma. B. vermiformis, Rhopalocoma, R. pyrotechnica, Palseaster
asperrimus, P. coronella, Palasterina, Protaster miltoni, P. leptosoma.)
1861. Additional notes on some new Palaeozoic star-fishes. Ann. Mag. Nat.
Hist., ser. 3, vol. 8, pp. 484-486, pi. 18, figs. 9-11.
1866. On the fossils of North Wales. Mem. Geol. Surv. Great Britain, vol. 3,
App., pp. 240-381, pis. 1-28. (Palseaster imbricatus.)
SALTER, J. W., and SOWERBY, J. de C.
1845. In SEDGWICK, On the Older Palaeozoic (Protozoic) rocks of North Wales.
Quart. Journ. Geol. Soc., London, vol. 1, pp. 5-22. (Ophmra salteri
and Asterias primseva listed in table, without description.)
SANDBERGER, G., and SANDBERGER, F.
1850-1856. Die Versteinerungen des rheinischen Schichtensystems in Nassau.
(Ccelaster, C. latiscutatus.)
286 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
SCHONDORF, F.
1907a. Ueber einen fossilen Seestern Spanister latiscutatus Sandb. spec, aus
dem Naturhistorischen Museum zu Wiesbaden. Jahrb. nassauisch.
Ver. f . Naturk. , Wiesbaden, vol. 60, pp. 17XM76, 3 text figs. (Spaniaster. )
19076. Ueber Archseasterias rhenana Joh. Muller und die Porenstellung palao-
zoischer Seesterne. Centralb. f. Mineral., etc., pp. 741-750, 6 text
figs.
1909a, Palaozoische Seesterne Deutschlands. I. Die echten Asteriden der
rheinischen Grauwacke. Palseontographica, vol. 56, pp. 37-112, text
figs., pis. 7-11. (Agalmaster, A. grandis, A. intermedius, A. miellensis,
Rhenaster, R. schwerdi, Trimeraster, T. parvulus, Eifelaster, E. foll-
manni, Xenaster dispar, X. elegans.)
19096. Die Asteriden des russischen Karbon. Palseontographica, vol. 56, pp.
323-338, text fig., pis. 23, 24.
1909c. Die fossilen Seesterne Nassaus. Jahrb. nassauisch. Ver. f. Naturk.,
Wiesbaden, vol. 62, pp. 7-46, text figs., pis. 2-5. (Discusses skeletal
structure of asterids; Miomaster, M. drevermanni.)
1909rf. Organization und Aufbau der Armwirbel von Ony chaster. Jahrb. nassau-
isch. Ver. f. Naturk., Wiesbaden, vol. 62, pp. 47-63, pi. 6.
1910a. Ueber einige "Ophiuriden und Asteriden" des englischen Silur und ihre
Bedeutung fiir die Systematik palaozoischer Seesterne. Jahrb. nassau-
isch. Ver. f. Naturk., Wiesbaden, vol. 63, pp. 206-256, text figs. (Im-
portant for classification.)
19106. Palaozoische Seesterne Deutschlands. II. Die Aspidosomatiden des
deutschen Unterdevon. Palseontographica, vol. 5 7 , pp. 1-63, pis. 1-3.
(Discusses structure and systematics of group in much detail. Erects
a new order, Auluroidea. Aspidosoma goldfussi. A. roemeri. A. eifelense.)
1910c. Aspidosoma schmidti nov. spec. Der erste Seestern aus den Siegener
Schichten. Jahrb. k. preuss. geol. Landesanst. und Bergakad., Berlin,
vol. 29, 1908, pt. 1, pp. 698-708, 1 pi.
1913a. Palaeaster eucharis Hall aus dem nordamerikanischen Devon. Jahrb.
nassauisch. Ver. f. Naturk., Wiesbaden, vol. 66, pp. 87-96, text figs.
1, 2, pi. 3, figs. 1, 2.
19136. Ueber Onychaster, einen Schlangenstern aus dem Karbon. Jahrb. nas-
sauisch. Ver. f. Naturk., Wiesbaden, vol. 66, pp. 97-116, text figs.
1, 2, pi. 3, figs. 3-12.
SCHUCHERT, C.
1914. Fossilium Catalogus, I : Animalia, pars 3, Stelleroidea palaeozoica.
SIMONOVITSCH, S.
1871. Ueber einige Asterioiden der rheinischen Grauwacke. Sitzb. d. mat.-
naturw. Classe d. k. Akad. d. Wiss.. Wien, vol. 64, Abt. 1., pp. 77-122,
pis. 1-4. (Xenaster, X. margaritatus, X. simplex, Asterias acuminatus,
Aspidosoma petaloides.}
SOLLAS, I. B. J.
1913. On Onychaster, a Carboniferous brittle-star. Philos. Trans. Roy. Soc.
London, ser. B, vol. 204, pp. 51-62, text figs. 1-5, pis. 8, 9.
SOLLAS, W. J.
1899. Fossils in the University Museum, Oxford: I. On Silurian Echinoidea and
Ophiuroidea. Quart. Journ. Geol. Soc., London, vol. 55, pp. 692-715,
text figs. (Erects new order, Ophiocistia. Eudadia woodwardi, Euthe-
mon, E. igerna.)
SOLLAS, W. J., and SOLLAS, I. B. J.
1912. Lapworthura: a typical brittle-star of the Silurian age. Philos. Trans.
Roy. Soc. London, ser. B, vol. 202, pp. 213-232, text figs. 1-5, pis. 9, 10.
(Rhodostoma, Protaster groomi.)
KEVISIOK OF PALEOZOIC STELLEROIDEA. 287
SPENCER, J. W.
1884. Niagara fossils, Pt. 3. Bull. Mus. Univ. State Missouri, No. 1, pp. 52-61,
pis. 7, 8. (Palseaster granti.}
SPENCER, W. K.
1914. A monograph of the British Palaeozoic Asterozoa. Pt. 1. Palaeontogr. Soc.
for 1913 ; pp. 1-56, text figs. 1-31, pi. 1. (Eoactis, E. simplex, Uranaster
elizx, Aspidosoma grayse, Lapworthura sollasi.)
STSCHUROWSKY.
1874. Nachr. d. Moskauer Gesell. d. Liebhaber d. Naturges. (Asterias montanus.)
Not seen.
STURTZ, B.
1886a. Ueber palaozoische Seesterne. N. Jahrb. f. Mineral., etc., vol. 2, pp.
142-154. (Hallaster.)
18866. Beitrag zur Kenntniss palaozoischer Seesterne. Palseontographica, vol. 32,
pp. 75-98, pis. 8-14. (Ophiurella, 0. primigenia, Roemeraster, R. asperula,
Astropecten schluteri, Eoluidia, E. decheni, Protester acanthion, P. pri-
mus, Fur caster, F. palseozoicus, Bundenbachia, B. beneckei, B. grandis,
Palastropecten, P. zitteli, Loriolaster, L. mirabilis, Palasteriscus, P.
devonicus.)
1890. Neuer Beitrag zur Kenntniss palaozoischer Seesterne. Palseontographica,
vol. 36, pp. 203-247, pis. 26-31. (Echinasterella, E. sladeni, Cheiropter-
aster, C. giganteus, Medusaster, M. rhenanus, Palseostella, P. solida,
Ophiurina, 0. lymani, Palseophiura, P. simplex, Palasterina follmanni.
Describes additional material of forms dealt with in 1886, revises
other species, discusses structure, systematics, habits, etc., of Paleozoic
ophiurids and asterids.)
1893. Ueber versteinerte und lebende Seesterne. Verh. naturh. Ver. preuss.
Rheinl., etc., vol. 50, pp. 1-92, pi. 1. (Salteraster, Ophiura rhenana,
Palxnectria, P. devonica.)
1900. Ein weiterer Beitrag zur Kenntniss palaozoigcher Asteroiden. Verh. naturh.
Ver. preuss. Rheinl., etc.,- vol. 56, pp. 176-240, pis. 2-4. (Reviews
Gregory's work of 1896 and gives classification of Ophiuroidea with
definition of families. Hisingeraster, Trentonaster, Hudsonaster, Pseudo-
palasterina, Eophiurites, Palseophiomyxa, Palseospondylus, Palxosolaster,
P. gregoryi, Echinasterias, E. spinosus, Echinodiscus, E. multidactylus,
Echinostella, E. traquairi, Jaekelaster, J. petaliformis.)
THORENT.
1838. M^moire sur la constitution ge*ologique de la partie nord du Departe*ment
de 1'Aisne. Mem. Soc. ge"ol. de France, vol. 3, pp. 239-260, pi. 22.
(Asterias constellata.)
TRAUTSCHOLD, H.
1879. Die Kalkbriiche von Mjatschkowa, Theil 3. M^m. Soc. imp. Nat. Moscou,
vol. 14, pp. 101-180, 7 pis. (Calliaster, C. mirus, Stenaster confluens.)
TROOST, G.
1835. Description of a new species of fossil Asterias (Asterias antiqua). Trans.
Geol. Soc. Pennsylvania, vol. 1, pp. 232-235.
ULRICH, E. 0.
1878. Descriptions of some new species of fossils from the Cincinnati group;
Journ. Cincinnati Soc. Nat. Hist., vol. 1, pp. 92-100, pi. 4. (Protasterina,
P.fimbriata.)
1879. Descriptions of new genera and species of fossils from the Lower Silurian
about Cincinnati. Journ. Cincinnati Soc. Nat. Hist., vol. 2, pp. 8-30,
pi. 7. (Palseaster finei.)
50601 Bull. 88 15 19
288 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
VERRILL, A. E.
1914. Monograph of the shallow-water starfishes of the North Pacific coast from
the Arctic Ocean to California. Smithsonian Institution, Harriman
Alaska ser., vol. 14.
WOODWARD, H.
1869. On Eucladia, a new genus of Ophiuridae, from the Upper Silurian, Dudley.
Geol. Mag., vol. 6, pp. 241-245, pi. 8. (Eucladia, E. johnsoni.)
1874. Description of a new species of starfish from the Devonian of Great Ingle-
bourne, Harberton, South Devon. Geol. Mag., dec. 2, vol. 1, pp. 6-10,
96, 238, 432. (Helianthaster filitiformis.}
WORTHEN, A. W., and MILLER, S. A.
1883. Descriptions of new Carboniferous Echinoderms. Geol. Surv. Illinois,
vol. 7, pp. 327-331, pi. 31. (Compsaster, C. formosus, Cholaster, C. pe-
culiaris, Tremataster, T. dijficilis.)
WRIGHT, T.
1862. A monograph on the British fossil Echinodermata from the Oolitic forma-
tions, vol. 2, pt. 1. Palseontogr. Soc. for 1861. (Contains a diagnosis of
nine Paleozoic genera with species.)
1873. On a new genus of Silurian Asteriadse. Quart. Journ. Geol. Soc., London,
vol. 29, p. 421. (Trichotaster, T. plumiformis.)
ZITTEL, K. von.
1879. Handbuch der Palaontologie, vol. 1.
EXPLANATION OF PLATES.
PLATE 1.
FIG. 1. Rudsonaster narrawayi (Hudson) (also see plates 2 and 4).
A greatly enlarged photograph of the genoholotype of Protopalssaster
narrawayi Hudson. The inner ventral side of the specimen is here seen
with all of the dorsal skeleton weathered away except one of the basal
supramarginalia. Note the well-preserved oral armature and in places
the ambulacralia. The latter are the "covering plates" of Hudson.
Photograph by Prof. George H. Hudson.
Middle Ordovicic (Black River). Ottawa, Canada.
Collection of Mr. J. E. Narraway.
PLATE 2.
FIG. 1. Hudsonaster narrawayi (Hudson) (also see plates 1 and 4).
A photograph X 6 of the actinal side. Note the five long spines (tori)
of the oral armature.
Middle Ordovicic (Black River). Kirkfield, Ontario, Canada.
Collection of Peabody Museum, Yale University.
FIG. 2. Hudsonaster matutinus (Hall) (also see plates 3 and 5).
A photograph X 3 of three specimens showing the actinal area as preserved
on a piece of black limestone.
Middle Ordovicic (Trenton). Rathbone Brook, near Newport, Her-
kimer County, New York.
Collection of Museum of Comparative Zoology, Harvard University
(No. 26).
PLATE 3.
FIG. 1. Hudsonaster rugosus (Billings).
A photograph X 2 of the abactinal area of a cotype.
Upper Ordovicic (Richmondian). Charleton Point, Anticosti Island,
Gulf of St. Lawrence, Canada.
Collection of the Geological Survey of Canada (No. 1999).
FIG. 2. Hudsonaster matutinus (Hall) (also see plates 2 and 5).
A retouched photograph X 2 showing the abactinal side.
Middle Ordovicic (Trenton). Trenton Falls, New York.
Collection of the Museum of Comparative Zoology, Harvard University
(No. 3).
FIG. 3. Hudsonaster batheri, new species.
A diagram X 3 of the actinal side made from a wax squeeze by Bather,
now in the United States National Museum (Cat. No. 60601).
Upper Ordovicic. Thraive, Girvan, Scotland.
289
290 ' BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
PLATE 4.
FIG. 1. Hudsonaster narrawayi (Hudson) (also see plates 1 and 2).
A camera lucida drawing X 8 of the actinal side. The granules are
drawn somewhat too strongly.
Middle Ordovicic (Black River). St. Paul, Minnesota.
Ulrich collection of the United States National Museum (Cat. No.
60602).
FIG. 2. Hudsonaster milleri, new species.
A diagram, considerably enlarged, of an axillary area from the actinal
side.
Middle Ordovicic (Trenton). Fayette County, Kentucky.
Collection of the State University of Kentucky.
FIG. 3. Mesopalxaster (?) lanceolatus, new species.
Camera lucida drawing of one ra> from the actinal area, X 2. The
central furrow in the ambulacrum is decidedly V-shaped.
Upper Ordovicic (Utica). Near Rome, New York.
Collection of Peabody Museum, Yale University.
FIG. 4. Australaster giganteus (Etheridge).
An axillary area and part of two rays somewhat reconstructed from the
original figure. Natural size. Note the single very large axillary plate,
the enlarging adambulacrals, and the diminishing inframarginals.
"Permo-Carboniferous" (Lower Marine). Parley, Northumberland
County, New South Wales.
Collection of the Mining and Geological Museum, Sydney.
PLATE 5.
FIGS. 1 and 2. Hudsonaster matutinus (Hall) (also see plates 2 and 3).
1. Camera lucida drawing X 3.5 of the actinal side of one of the speci-
mens photographed on plate 2, fig. 2.
Middle Ordovicic (Trenton). Near Newport, New York.
Collection of Museum of Comparative Zoology, Harvard University
(No. 26).
2. Camera lucida drawing X 4.5 of the abactinal area of a specimen in
which the ossicles are somewhat displaced.
Middle Ordovicic (Trenton). Lachine, near Montreal, Canada.
Collection of the United States National Museum (Cat. No. 60603).
Collected by W. R. Billings.
PLATE 6.
FIGS. 1 and 2. Hudsonaster incomptus (Meek).
1. Camera lucida drawing X 4 of the actinal side.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No. 40882).
2. Camera lucida drawing X 4 of the abactinal area in a well preserved
specimen. The anal opening may have been in the first circle of small
plates adjacent to the centro-dorsal piece and in the same interradius as
the madreporite.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No. 40882).
REVISION OF PALEOZOIC STELLEEOIDEA. 291
PLATE 7.
FIGS. 1-4. Palseaster niagarensis Hall.
1. Camera lucida drawing X 4 of the abactinal side in its present
preservation. Holotype.
2. Abactinal side of theholotype restored, X 4. The drawing probably
has too many accessory disk pieces.
3. Camera lucida drawing X 4 of the a'ctinal side of one of the rays of
the holotype. Since this drawing was made the rock in the ambulacral
furrow has been dug away down to the ambulacral plates, which are
small and deep-seated.
4. A few of the inframarginals and supramarginals from the abactinal
side to show the smooth central areas surrounded by granular borders, X 4.
Siluric (Rochester shale). Lockport, New York.
Collection of Cornell University (No. 7331).
FIG. 5. Mesopalseaster find (Ulrich) (also see plate 9).
Camera lucida drawing X 16 of one of the rays. The ossicles are always
more or less displaced in this species.
Upper Ordovicic (Eden shales). Cincinnati, Ohio.
Ulrich collection of the United States National Museum (Cat. No. 60604).
PLATE 8.
FIGS. 1 and 2. Mesopalseaster shafferi (Hall).
1. Camera lucida drawing X 8 of the abactinal side of a one-third
adult-sized specimen. As the medial disk pieces are displaced in the
individual, they are here drawn in in their probable natural arrangement
all should be pointed like the central disk plate.
Upper Ordovicic (Maysvillian, Corryville member). Cincinnati, Ohio.
Vaupel collection of the United States National Museum (Cat. No.
60605).
2. Camera lucida drawing X 3 of the actinal area of a mature indi-
vidual.
Upper Ordovicic (Kichmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
59391).
FIG. 3. Miomaster drevermanni Schondorf.
Actinal side as reconstructed by Schondorf. Natural size.
Lower Devonic (Upper Coblenzian). Miellen, Germany.
PLATE 9.
FIG. I. Mesopalseaster (?) parviusculus (Billings).
Camera lucida drawing X 5 of a gutta-percha squeeze of the holotype,
actinal side.
Siluric (Lower Arisaig). Arisaig, Nova Scotia.
Original at McGill University; gutta-percha squeeze in United States
National Museum (Cat. No. 60620).
FIG. 2. Mesopalseaster (?) cataractensis, new species.
Photograph X 3 of the well-preserved holotype.
Siluric (Cataract formation). Hamilton, Ontario.
Collection of the Hamilton Natural History Society.
FIG. 3. Mesopalseaster (?) granti (Spencer).
Photograph X 2 of a specimen showing the dorsal side.
Siluric (Cataract formation). Hamilton, Ontario.
Collection of Peabody Museum, Yale University.
292 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
FIG. 4. Mesopalseaster intermedius, new species.
Photograph X 3 of the holotype. Between the two rays lies an arm of
a crinid, giving the impression that this form has a large disk.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Collection of the University of Chicago (No. 9575).
FIG. 5. Mesopalseaster finei (Ulrich) (also see plate 7).
Ulrich's original figures. Fig. 5, one of the cotypes from the abactinal
side X 2; fig. 5a, the madreporite X 6; fig. 56, a ray from the actinal side X 3.
Upper Ordovicic (Eden). Cincinnati, Ohio.
Ulrich collection of the United States National Museum (Cat. No. 60604).
FIG. 6. Mesopalseaster caractad (Gregory) (also see plate 11).
Retouched photograph X 3 of a wax squeeze by Bather, now in the
United States National Museum (Cat. No. 60606). Actinal area.
Ordovicic (Caradoc). Church Stretton, England.
PLATE 10.
FIGS. 1 and 2. Mesopalseaster clarki (Clarke and Swartz).
1. Drawing of the actinal side of the holotype X 1.5.
2. Abactinal area X 1.5. Note the few accessory pieces in the angles
between the radials.
Upper Devonic (Chemung). Near Oakland, Maryland.
Collection of the Maryland Geological Survey.
PLATE 11.
FIG. 1. Mesopalxaster caractad (Gregory) (also see plate 9).
Retouched photograph X 3 of a wax squeeze by Bather, now in the
United States National Museum (Cat. No. 60606). Abactinal area.
Ordovicic (Caradoc). Church Stretton, England.
FIG. 2. Devonaster chemungensis, new species.
Photograph of the natural mold of the actinal side of the holotype,
natural size.
Upper Devonic (Chemung). "Central Pennsylvania."
CoUection of Columbia University (No. 6228G).
PLATE 12.
FIGS. 1 and 2. Spaniaster latiscutatus (Sandberger).
1. Abactinal side.
2. Actinal side.
After Schondorf .
Lower Devonic. Germany.
FIGS. 3-5. Devonaster eucharis (Hall).
3. Abactinal side, natural size, a, the madreporite. After Hall.
4. Actinal side, natural size. After Hall. The tiny ambulacral plates
should have been drawn as opposite one another and not alternate.
5. Hall's diagram (3a) of the actinal plate arrangement. The ambu-
lacrals are incorrectly drawn, as their arrangement is opposite and not
alternate, a, ambulacralia; aa, adambulacralia; m, inframarginalia; o,
pairs of oral armature ossicles; p, podial openings, but drawn a little too
large; tm, the single interbrachial axillaries.
Middle Devonic (Hamilton). Near Hamilton, New York.
REVISION OF PALEOZOIC STELLEROIDEA. 293
PLATE 13.
FIGS. 1 and 2. Promopalseaster wilsoni (Raymond).
1. Photograph X 1.5 of the holotype, showing the abactinal side.
2. One of the rays of the holotype X 3.
Middle Ordovicic (Lowville). City View, Ottawa, Canada.
Collection of Miss A. E. Wilson.
FIG. 3. Promopalseaster prenuntius, new species (also see plate 15).
Photograph, natural size, of the actinal side of the holotype.
Middle Ordovicic (Lower Trenton). Frankfort, Kentucky.
Collection of the State University of Kentucky.
FIG. 4. Anorthaster miamiensis (Miller) (see also plate 20).
Retouched photograph, natural size, of the actinal side of the holotype.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40880).
FIG. 5. Neopalseaster crawfordsvillensis (Miller) (see also plate 23).
Retouched photograph, natural size, of the holotype, showing the abac-
tinal area.
Mississippic (Keokuk). Crawfordsville, Indiana.
Harris collection of the United States National Museum (Cat. No. 60607).
PLATE 14.
FIGS. 1 and 2. Promopalseaster speciosus (Meek)?
Reproduction, natural size, of the original lithograph of "Asterias prir
mordialis." Probably the young of P. speciosus.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
The specimen is now lost.
FIGS. 3 and 4. Promopalseaster speciosus (Meek) (also see plate 15).
Photographs, natural size, of the abactinal and actinal sides of the
holotype.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard
University (No. 22).
PLATE 15.
FIGS. 1-4. Promopal&easter speciosus (Meek) (also see plate 14).
1. Camera lucida drawing X 4 of an interbrachial area.
2. Camera lucida drawing X 4 across a ray at about its mid-length.
From the actinal side, showing the ambulacrals, podial openings,
adambulacrals, inframarginals, and supramarginals.
3. Two adambulacral spines near oral region, X 7.
4. A part of the abactinal area of a ray near its mid-length, X 2.5.
All drawn from the holotype.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard Uni-
versity (No. 22).
FIG. 5. Promopalseaster prenuntius, new species (also see plate 13).
Diagram of the actinal interbrachial plate arrangement.
294 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
FIGS. 6-8. Promopalseaster bellulus, new species (also see plates 16 and 18).
6. Camera lucida drawing X 4 of an interbrachial area. The adambu-
lacrals are in natural position, but the interbrachial axillaries and the
inframarginals are somewhat displaced. Taken from the specimen on
plate 16, fig. 1.
7. Camera lucida drawing X 7 of the abactinal area of a ray. The
ossicles are all displaced, but originally they were united in a spicular
and partially overlapping mesh.
8. Madreporite X 7.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National .Museum (Cat. No.
40879).
PLATE 16.
FIG. 1. Promopalseaster bellulus, new species (also see plates 15 and 18).
Retouched photograph, natural size, of the actinal side. The specimen
is in limestone.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40879).
FIG. 2. Promopalseaster spinulosus (Miller and Dyer) (also see plates 17 and 18).
Retouched photograph, natural size, of the actinal side of the holotype
of Palseaster longibrachiatus Miller. The well-preserved columns are the
inframarginals, but in places some of the adambulacrals are also shown.
Upper Ordovicic (Richmondian). Near Clarksville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40881).
PLATE 17.
FIGS. 1 and 2. Promopalseaster spinulosus (Miller and Dyer) (also see plates 16 and
18).
Retouched photographs X 3 of the actinal and abactinal sides of the
holotype.
Upper Ordovicic (Richmondian). Probably near Waynesville, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard
University (No. 16).
PLATE 18.
FIGS. 1-3. Promopalseaster spinulosus (Miller and Dyer) (also see plates 16 and 17).
1. Camera lucida drawing X 2, somewhat restored, of the holotype
of Palseaster longibrachiatus Miller.
2. Camera lucida drawing X 7 of the abactinal area of a ray near its
base. Shows the large infra- and supramarginals and ambital pieces.
Center of ray has spicular ossicles.
3. Madreporite X 7.
Upper Ordovicic (Richmondian). Near Clarksville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40881).
REVISION OF PALEOZOIC STELLEEOIDEA. 295
FIGS. 4 and 5~Promopalseaster bellulus, new species (also see plates 15 and 16).
4. Adambulacralia and their spines, X 7. The ambulacrum lies to
the left.
5. A few abactinal spines, X 7.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40879).
FIG. G.Promopalseaster wyloffi, (Miller and Gurley) (also see plate 19).
Diagram X 3.5 of one of the interbrachial areas.
From the holotype.
Upper Ordovicic (Richmondian). Near Madison, Indiana.
Collection of the University of Chicago (No. 6066).
FIG. 7. Promo palaeaster exculptus (Miller) (also see plate 20).
Adambulacral and ambulacral plates at about mid-length of a ray, X 4.
From the holotype.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
60608).
FIG. S.Promopalssaster dyeri (Meek) (also see plates 20 and 25).
Five inframarginal plates X 4. To show the articulations for the stout
spines that lie on the right. Also note the smaller spines. The plate
extensions to the left connect with the vertical ridges on the ambulacral
plates.
From the holotype.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard Univer-
sity (No. 13).
PLATE 19.
FIG. 1. Trimeraster parvulus Schondorf.
Reconstruction X 1.75 of the actinal side. After Schondorf.
Lower Devonic (Coblenzian). Near Rhens, Germany.
FIG. 2. Promo palseaster wykoffi (Miller and Gurley) (also see plate 18).
Photograph X 2 of the holotype, showing the actinal side. One of
the finest of Paleozoic specimens.
Upper Ordovicic (Richmondian). Near Madison, Indiana.
Collection of the University of Chicago (No. 6066).
PLATE 20.
FIG. 1. Anorthaster miamiensis (Miller) (also see plate 13).
Camera lucida drawing X 5.5 of one of the interbrachial areas of the
holotype. The only Paleozoic form known with the area wholly made
up of adambulacrals.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40880).
FIG. 2.Promopalseaster exculptus (Miller) (also see plate 18).
Camera lucida drawing X 4 of an interbracbial area. Note the modified
ambulacral plates in connection with the oral armature. Holotype.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
296 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
FIGS. 3-6. Promopalseaster dyeri (Meek) (also see plates 18 and 25).
3. One of the interbrachial areas with the ossicles displaced and the
adjoining adambulacral plates with their spines, X 4.
4. Madreporite X 4. The rest of the plate is covered.
5. Two abactinal spines, X 7.
6. Five different abactinal ossicles, X 7.
All drawn from the holotype.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard Uni-
versity (No. 13).
PLATE 21.
FIG. 1. Promopalseaster magnificus (Miller) (also see plates 22 and 23).
Photograph, natural size, of the actinal side of one of the cotypes. This
speci nen is probably the best preserved of Paleozoic starfishes and is one
of the largest species.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No. 40883).
PLATE 22.
FIG. 1. Promopalseaster magnificus (Miller) (also see plates 21 and 23).
Abactinal side of the same specimen illustrated on plate 21.
PLATE 23.
FIGS. 1-3. Promopalseaster magnificus (Miller) (also see plates 21 and 22).
1. Camera lucida drawing X 3.5 of an interbrachial area. Additional
tube-feet probably also came out at the junction of the forked crests.
2. Camera lucida drawing X 4 of the actinal area of one of the rays
toward the distal ends. Large adambulacrals bound the rays distally,
while the smaller pieces outside are the inframarginals.
3. A distal portion of the abactinal area, X 3.5.
All drawn from the cotypes.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
40883).
FIG. 4. Neopalseaster crawfordsvillensis (Miller) (also see plate 13).
Camera lucida drawing X 4 of the abactinal area of the holotype. Ac-
cessory disk pieces are drawn in; in the specimen they are all displaced.
The madreporite should be more finely striate. In places the dorsal
skeleton is lost, exposing the ambulacrals.
Mississippic (Keokuk). Crawfordsville, Indiana.
Harris collection of the United States National Museum (Cat. No.
60607).
FIGS. 5-7. Petraster spedosus (Miller and Dyer) (also see plates 26 and 27).
5. The ambulacral, adambulacral and inframarginals of a ray at its
mid-length, X 4. Seen from the actinal side.
6. Inframarginal, accessory interbrachial, and adambulacral pieces, X 4.
Seen from the dorsal side.
7. Ambulacral and adambulacrale, X 4. Seen from the dorsal area.
REVISION OF PALEOZOIC STELLEROIDEA. 297
PLATE 24.
FIGS. 1-2. Xenaster margaritatus Simonovitsch.
1. Reconstruction, somewhat enlarged, of the abactinal side. The
structure of the medial portion of the disk is based upon Agalmaster
miellensis.
2. Reconstruction, somewhat enlarged, of the actinal area.
After Schondorf.
* Lower Devonic (Upper Coblenzian). Niederlahnstein, Germany.
PLATE 25.
FIG. 1. Promopalssaster dyeri (Meek) (also see plates 18 and 20).
Retouched photograph, natural size, of the actinal side of the holotype.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Dyer collection, Museum of Comparative Zoology, Harvard University
(No. 13).
FIG. 2. Lepidasterella babcocki, new species.
Photograph X 1.5 of the holotype. A natural mold of the abactinal
area in sandstone.
Upper Devonic (Lower Chemung). Near Ithaca, New York.
Collection of Mrs. G. W. Babcock.
PLATE 26.
FIG. 1. Petraster spedosus (Miller and Dyer) (also see plates 23 and 27).
Retouched photograph, natural size, of the abactinal area of the holotype.
Upper Ordovicic (Richmondian). Near Winchester, Ohio.
Dyer collection of the Museum of Comparative Zoology, Harvard Uni-
versity (No. 14).
FIG. 2. Petraster (?) americanus (D'Orbigny).
Reproduction, natural size, of Meek's original figure. The specimen is
seen from the abactinal side, exposing, however, the inner surface of the
actinal skeleton, which is deeply embedded in limestone.
Upper Ordovicic (Maysvillian). Cincinnati, Ohio.
Collection of the University of Chicago.
PLATE 27.
FIGS. 1-4. Petraster spedosus (Miller and Dyer) (also see plates 23 and 26).
1. Camera lucida drawing X 4 of the abactinal side of a ray. Holotype
The inframarginals border the animal, with the supramarginals imme-
diately inside of them. The radial column is readily distinguished
along the center of the rays.
2. Camera lucida drawing X 6 of an interbrachial area in a young
individual (Harris collection, Cat. No. 60609, U.S.N.M.). The inframar-
marginals are above and the adambulacrals below to the right. The rest
rest of the plates are accessory interbrachials.
3-4. Madreporite from the side and from the lower or under surface, X 8.
FIG. 5. Petraster rigidus (Billings).
Outline tracing of the actinal skeleton from a photograph X 2 of the
holotype.
Middle Ordovicic (Trenton). Ottawa, Canada.
Collection of the Geological Survey of Canada (No. 1401a).
298 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
FIGS. 6-8. Urasterella grandis (Meek) (also see plates 28 and 30).
6. Camera lucida drawing X 8 of part of the abactinal area of a ray proxi.
mally. Note how the ossicles are drawn out into nonarticulating blunt
spines.
7. Camera lucida drawing X 4 of the actinal side about the mouth. The
complete oral armature is preserved.
8. Madreporite in outline, X 7.
From specimens in the Harris collection of the United States National
Museum (Cat. No. 40885).
PLATE 28.
FIGS. 1 and 2. Urasterella grandis (Meek) (also see plates 27 and 30).
1. Retouched photograph, natural size, of a large and folded specimen.
Both ventral and dorsal areas are shown.
2. Retouched photograph, natural size, of an average individual seen
from .the actinal side.
Upper Ordovicic (Richmondian). Near Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No. 40885).
FIGS. 3 and 4. Urasterella pulchella (Billings) (also see plate 30).
3. Photograph, natural size, of an electrotype, seen from the actinal side.
Middle Ordovicic (Trenton). Hull, Ottawa, Canada.
Original in collection of the Geological Survey of Canada; electrotype in
United States National Museum (Cat. No. 60610).
4. Retouched photograph, natural size, of the abactinal area.
Middle Ordovicic (Trenton). Trenton Falls, New York.
Collection of the Museum of Comparative Zoology, Harvard University
(No. 31).
FIG. 5.' Urasterella girvanensis, new species.
Reproduction X 2 of the original figure by Nicholson and Etheridge,
somewhat altered.
From a wax squeeze made by Bather, now in the United States National
Museum (Cat. No. 60611).
Upper Ordovicic. Thraive, Girvan, Scotland.
PLATE 29.
FIG. 1. Urasterella ulrichi, new species (also see plate 30).
Retouched photograph, natural size, of the abactinal side of a cotype.
The three short blunt rays are interpreted as healed stumps.
Middle Ordovicic (Black River). Minneapolis, Minnesota.
Ulrich collection of the United States National Museum (Cat. No. 60612).
FIG. 2. Urasterella huxleyi (Billings).
Photograph X 2 of the holotype. Abactinal view.
Middle Ordovicic (about Chazy). Point Rich, Newfoundland. .
Collection of the Geological Survey of Canada (No. 554.)
PLATE 30.
FIGS. 1-4. Urasterella grandis (Meek) (also see plates 27 and 28.)
1-2. Camera lucida drawing X 16 of the adambulacrals with their articu-
lar spines and probable paxillse.
3. Camera lucida drawing X 8 of the two columns of ambulacra!
plates a little displaced. The podial openings are situated laterally
between the thinner ends of the ossicles.
4. Camera lucida drawing X 7 of the adambulacrals with their spines.
From specimens in the Harris collection of the United States National
REVISION OF PALEOZOIC STELLEROIDEA. 299
FIG. 5. Urasterella pulchella (Billings) (also see plate 28).
Camera lucida drawing X 8. The large plates to the left are the coin-
shaped adambulacrals (some have tipped over and show their articular
faces), next are the small elongate inframarginals, and outside of these
are the spinose ambitals.
FIGS. 6 and 7. Urasterella ulrichi, new species (also see plate 29).
6. Camera lucida drawing X 4 of the abactinal side of a young
individual. The medial ray plates should be more tumid than here
represented. Note the primitive structure of the disk.
7. Camera lucida drawing X 6 of the actinal side of a young indi-
vidual. The outer dark margin is adhering rock, through which some
of the abactinal ambital nonarticulating spines project. Note the
phanerozonian interbrachial characters.
Middle Ordovicic (Black River). Minneapolis, Minnesota.
Ulrich collection of the United States National Museum (Cat
No. 60612).
PLATE 31.
FIGS. 1 and 2. Compsaster formosus Worthen and Miller.
1. Actinal view, natural size, of the holotype. Reproduction of the
original illustration.
2. Part of a ray, X 2. After Worthen and Miller.
Upper Mississippic (Chester). Okaw Bluffs, Illinois.
" Illinois State collection of 1880, No. 2476."
FIG. S.Calliasterella mira (Trautschold).
Reproduction, natural size, of Schondorf's reconstruction.
Upper Carboniferous (Moscovian). Mjatschkowa, near Moscow, Russia.
PLATE 32.
FIG. 1. Stenaster salteri Billings.
Retouched photograph X 3 of a cotype. Actinal view.
Middle Ordovicic (Trenton). Belleville, Ontario, Canada.
Collection of the Geological Survey of Canada.
FIG. 2. Schuchertia stellata (Billings) (also see plate 33).
Photograph X 4 of the actinal side of the holotype.
Middle Ordovicic (Trenton). Ottawa, Canada.
Collection of the Geological Survey of Canada (No. 1399).
FIG. 3. Schuchertia laxata, new species (also see plate 33).
Retouched photograph X 2 of the holotype from the abactinal side.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No. 60613).
FIGS. 4 and 6. Schcenaster (?) legrandensis Miller and Gurley.
4-5. Actinal and abactinal views, natural size.
6. Part of actinal side X 6.5. After Miller and Gurley.
Lower Mississippic (Kinderhookian). Le Grand, Iowa.
?Gurley collection of the University of Chicago.
PLATE 33.
FIG. 1. Schuchertia stellata (Billings) (also see plate 32).
Camera lucida drawing X 8 of the abactinal surface. Somewhat
restored.
Middle Ordovicic (Trenton). Near New Edinburgh, Canada.
Collection of the Geological Survey of Canada.
FIGS. 2 and 3. Schuchertia laxata, new species (also see plate 32).
2. Some of the outer ambital plates, all disjointed, X 8.
3. An interbrachial area and ambulacrum, X 4.
Upper Ordovicic (Richmondian). Waynesville, Ohio.
Harris collection of the United States National Museum (Cat. No.
300 BULLETIN 88, UNITED STATES NATIONAL MUSEUM.
FIG. 4. Tetraster wyville-thoTtisoni Nicholson and Etheridge.
Diagram X 4 of the actinal skeleton. From a wax squeeze by Bather,
in the United States National Museum (Cat. No. 60614).
Upper Ordovicic. Thraive, Girvan, Scotland.
FIG. 5. Schcenaster (?) montanus Raymond.
Reproduction X 2 of the original figure. Actinal side.
Mississippi (Madison). Spring Canyon, near Alder, Montana.
Collection of the Carnegie Museum.
FIG. 6. SJicenaster (?) wachsmuihi Meek and Worthen.
Reproduction, natural size, of the original figure. Actinal view.
Lower Mississippi (Burlington). Burlington, Iowa.
Collection of the Museum of Comparative Zoology, Harvard University
(No. 7).
PLATE 34.
FIG. 1. Palseosolaster (?) gyalum (Clarke).
Actinal view of one of the cotypes, natural size.
Upper Devonic (Portage). Ithaca, New York.
Collection of Cornell University.
PLATE 35.
FIGS. 1-4. Schcenaster fimbriatus Meek and Worthen.
1. View of the abactinal side, natural size, of one of the cotypes.
2. Enlarged view of one of the rays, showing the arrangement of the
ossicles and the pores, as seen when the surface is ground down.
3. Actinal side of a ray, enlarged, showing only the adambulacrals.
4. Actinal side of another cotype, natural size.
The original figures of plate 19, figs. 7 a to 7d.
Upper Mississippi (St. Louis). St. Clair County, Illinois.
FIGS. 5 and 6 Encrinaster petaloides (Simonovitsch).
5. Abactinal view, about X 2.
6. An arm from the actinal side, about X 2.
After Schondorf.
Lower Devonic (Upper Coblenzian). Niederlahnstein, Germany.
PLATE 36.
FIG. 1. Tseniaster spinosus (Billings).
Photograph X 2 showing the actinal side of one of the cotypes.
Middle Ordovicic (Lower Trenton). Montmorency Falls, Quebec,
Canada.
Collection of the Geological Survey of Canada (No. 1404).
FIGS. 2 and 3.Taeniaster cylindricus (Billings).
2. Abactinal view X 2 of one of the cotypes.
3. Actinal view X 2 of another cotype. Both specimens are on the
same slab.
Middle Ordovicic (Trenton). Ottawa, Canada.
Collection of the Geological Survey of Canada (No. 1405a).
FIG. 4. Alepidaster flexuosus (Miller and Dyer).
An unpublished photograph X 3.5, made many years ago, of the holo-
type of Protasterina fimbriata Ulrich.
Upper Ordovicic (Lower Edenian). Covington, Kentucky.
Ulrich collection of the United States National Museum (Cat. No. 60615).
REVISION OF PALEOZOIC STELLEKOIDEA. 301
PLATE 37.
FIGS. 1 and 2. Eucladia woodwardi Sollas.
Dorsal and ventral reconstructions X 5/3 of this remarkable fossil. On it
is based the order Ophiocistia Sollas. "One arm, the most proximal of
the radius on the lower right-hand corner, is omitted, so as to show the
aperture from which it proceeds." After Sollas.
Siluric (Lower Ludlow). Leintwardine, England.
PLATE 38.
FIG. l.Eucladm (?) beecheri, new species.
Photograph X 2 of the holotype from the actinal side. The specimen
itself is difficult to make out, hence the indistinctness of the photograph.
Lower Devonic (Coeymans). Jerusalem Hill, Litchfield, Herkimer
County, New York.
Collection of Peabody Museum, Yale University.
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HUDSON ASTER NARRAWAYI.
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HUDSONASTER INCOMPTUS.
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MESOPAL/E ASTER CLARKI.
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, MESOPAL/EASTER CARACTACI; 2, DEVONASTER CHEMUNGENSIS.
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1 , 2, SPANIASTER LATISCUTATUS; 3-5, DEVONASTER EUCHARIS.
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1-3, PROMOPAMEASTER; 4, ANORTHASTER; 5, NEOPAL/EASTER.
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Asterlas prtmorci tails
PROMOPALXEASTER SPECIOSUS.
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1-4, PROMOPAL/EASTER SPECIOSUS; 5, P. PRENUNTIUS; 6-8, P. BELLULUS.
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PROMOPAL/CASTER SPINULOSUS.
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4
DETAIL OF FIVE SPECIES OF PROMOPAL/EASTER.
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1 , TRIMERASTER PARVULUS; 2, PROMOPAL/EASTER WYKOFFI.
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PROMOPAL/EASTER MAQNIFICUS.
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XENASTER MARQARITATUS.
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1, PROMOPAL/EASTER DYERI; 2, LEPIDASTERELLA BABCOCKI.
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BULLETIN 88 PL. 26
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M/7-? 4- > y . ' - l -4- >r r^T > -
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1, PETRASTER SPECIOSUS; 2, P. (?) AMERICANUS.
U. S. NATIONAL MUSEUM
BULLETIN 88 PL. 27
1-5, DETAIL OF PETRASTER; 6-8, URASTERELLA.
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1, 2, URASTERELLA GRANDIS; 3, 4, U. PULCHELLA; 5, U. QIRVANENSIS.
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DETAIL OF URASTERELLA.
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1, STENASTER; 2, 3, SCHUCHERTIA; 4-6, SCHCENASTER (?).
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^K^fflK^
1-3, DETAIL OF SCHUCHERTIA; 4, TETRASTER; 5, 6, SCHCENASTER (?).
U. S. NATIONAL MUSEUM
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.~~
Kg^K, ji ^ .^
"^l^te'^
PAL/EOSOLASTER (?) GYALUM.
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1-4, SCHCENASTER FIMBRIATUS; 5, 6, ENCRINASTER PETALOIDES.
FOR EXPLANATION OF PLATE SEE PAGE 300.
U. S. NATIONAL MUSEUM
BULLETIN 88 PL. 36
1, T>ENIASTER SPINOSUS; 2, 3, T. CYLINDRICUS; 4, ALEPIDASTER FLEXUOSUS.
FOR EXPLANATION OF PLATE SEE PAGE 300.
U. S. NATIONAL MUSEUM
BULLETIN 88 PL. 37
EUCLADIA WOODWARDI.
FOR EXPLANATION OF PLATE SEE PAGE 301.
U. S. NATIONAL MUSEUM
BULLETIN 88 PL.
EUCLADIA (?) BEECHERI.
FOR EXPLANATION OF PLATE SEE PAGE 301.
INDEX.
[The important references In each case are in heavy-faced type. Synonyms are in italics.]
Page.
abactinal 13
abnormal development 101
aboral, see abactinal.
accessory actinals 13
accessory interbrachials 13, 15,45,46
accessory plates 13, 47, 51
Acroura ( Ophiura) schlotheimii 18
actinal 13
acuminatus, Asterias. ., 93
acuminatus, Mesopalseaster (?) 75, 93
acuminatus, Roemeraster (?) 93
adambulacralia, see adambnlacrals.
adambulacral oral skeleton 16
adambulacral spines 13, 43
adambulacrals (see also lateral shields) 13,
34,43,162,213,214,222
Agalmaster 40, 49, 128, 13 1 , 133
A.grandis 132
A. intermedium 133
A.miellensis 132
Aganaster 263
A . gregarius 264
A. (?) sp. Miller and Gurley 251
Aganasteridse 246, 263
Alepidaster 216,228
A. flexuosus 230, 231
A. granuliferus 229, 230
A. gregarius 264
A. miamiensis '. 230, 238
A. n. sp 230
ambital and ambital plates 13, 46, 174
ambulacralia, see ambulacrals.
ambulacral oral skeleton 16
ambulacrals 13,33,34,41,213,235,258
alternate arrangement 13,
33, 41 , 152, 172, 174, 213, 214, 215, 222, 226
opposite arrangement 13,
41,163,213,214,224,246
americanus, Cadaster 146
americanus, Petraster (?) 139, 146
ampullae 213
anal opening 13,89,54,61
Anorthaster 41,42,43,50, 125
A. miamiensis 127
Anorthasterinse 52, 74, 126
anthonii, A sterlas 146
antiqua, Argaster 86
antiqua, Asterias, Hisinger 149
antiqua, Asterias, Troost 86
antiqua, Hisingeraster 149
antiqua, Palseaster (Argaster) 80
antiqua, Palxasterina 149
50601 Bull. 8815 20
Page.
antiqua, Palmipes 149
antiqua, Petraster (?) 86
antiquata, Asterias 109
antiquata, Palxaster 109
antiquus, Asteriscus 149
antiquus, Lindstromaster 149, 153
antiquus, Mesopalseaster (?) 75, 86
antiquus, Palxaster 86
anus, see anal opening.
approximata, Palxasterina 142
A rchxasterias 129, 130
arcs, see axil.
? Argaster 74, 77, 87
A . antiqua 86
arnoldi, Aspidosoma 243
arnoldi, Encrinaster 241, 248
asper, Onychaster 270, 2 72
asperrima, Palxaster. 187
asperrima, Urasterella (?) 175, 18 7
asperrimus, Salteraster 187
asperrimus, Tetraster 187
asperula, A sterias 188, 262
asperula, Roemeraster 188
asperula, Urasterella 175, 188
Aspidosoma 241
A . arnoldi 243
A.eifelense 244
A. goldfussi 243
A . gray se 245
A. petaloides 243
A. petaloides goslariensis 243
A . ? pontis 244
A . roemeri 244
A. sehmidti 244
A. tischbeinianum 244
Aspidosomatidx 241
Asterias 33
A sterias Graham, Anthony and James 146
A . acuminatus 93
A . anthonii 146
A . antiqua Hisinger. 149
A . antiqua Troost 86
A . antiquata 109
A, asperula 188,262
A.constellata 187
A. matutina 57
A. montanus 189
A . primxva 167
A . primordialis 108
A. (Archxasterias) rhenana 131
A. spinosissima 19
asterid radicle 34
ysterina 33
303
304
INDEX.
Page.
Asteriscus antiquut 149
Asteroidea 41, 42, 43, 51
evolution of 11, 30, 48, 52, 106
number of, in Paleozoic 2 7, 28
terminology 13
Astropecten 161
A. (?) schliiteri 161
Ataxaster 161
A. pygmseus 161
Auluroidea 80,38,40,41,42,43,163,212,213
geological distribution of 28
number of, in Paleozoic 27,28
Australaster 44,45,49,66,72, 171
A.giganteus 73, 171
A. (?) stutchburii 73
australis, Txniaster 236
axil 14
axillary inframarginal 14
axillary interbrachial 14
axillary marginal 15
axillary ossicles 34, 49
babcocki, Lepidasterella 160
barrisi, Onychaster 270, 272
barrisi, Protester (?) 272
bathed, Hudsonaster J 55, 65, 167
Bdellacoma 248,252,254
B. vermiformis 254
beecheri, Eucladia (?) 278
bellulus, Mesopalseaster 75, 91
bellulus, Petraster 91
bellulus, Promopalseaster 104, 105, 1 13
beneckei, Bundenbachia 234
bibliography 281-288
biforis, Protaster 226, 227
bivium 14
body-wall 14
bohemica, Eophiura 222
Bohemura 216, 22 8
B.jahni 223
bonneyi, Palxasterina 153
bonneyi, Palasterina 151, 152, 168
brisingoides, Protaster 236
brisingoides, Stiirtzura 236
buccal processes 225
Bundenbach 29
Bundenbachia 216, 284
B . beneckei 234
B. grandis 235
caecal pores 14
Calliaster 190
C. mirus 190
Calliasterella 39, 47, 49,50, 190
C. mira 191
Calliasterellidse 162, 163, 190
Calliasteridx 190
caractaci, Mesopalseaster 75, 92
caractaci, Palseaster 92
caractaci, Protopalxaster 92
carbonarius, Cribellites 274
carinals, see radials.
catalogue of Paleozoic Stelleroidea 51
cataractensis, Mesopalaeaster (?) 75, 89
central disk, see disk.
central disk plate, see centrodorsal plate.
central plate, see centrodorsal plate.
centrodorsal plate 14, 34,35,48
Page.
Cheiropteraster 40, 199, 202, 252
C. giganteus 202
chemungensis, Devonaster 101
Cholaster 265
C. peculiaris 266
Cholasteridse 246, 265
ciliaris, Ophiura 268
clarkana, Palseaster 61,63
clarkei, Etheridgaster 172
clarkei, Monaster 171, 1 7 2
clarkei, Palseaster, De Koninck 172
clarkei, Palseaster, Miller 61
clarki, Mesopalaeaster (?) 75, 94
clarki, Palxaster 94
Classification 51
Cadaster 95
C. americanus 146
C. latiscutatus 96
C. tenuiradiatus 19
colvini, Palzocoma 253
colvini, Stiirtzaster 253
Compsaster 192
C. formosus 193
C. n.sp 194
Compsasteridse 162, 163, 191
concinna, Eugasterella (?) 238, 289
concinnus, Eugaster 239
confluens, Stenaster (?) 165,167
confragosus, Onychaster 270, 273
constellata, Asterias 187
constellata, Urasterella (?) 175, 187
coronella, Palxaster 167
coronella, Stenaster (?) 165, 167
crawfordsvillensis, Neopalseaster 135, 186
crawfordsvillensis, Palaeaster 136
Cribellites carbonarius 274
Crossaster 38
Cryptozonia.... 80,33,38,40,41,43,44,45,46,47,162
cygnipes, Palseocoma 253
cygnipes, Sturtzaster. 253
cylindrica, Lapworthura 220
cylindrica, Palseocoma 220
cylindrica, Txniura. 220
cylindricus, Tseniaster 216,217,218,219,220
daoulasensis, Furcaster (?) 261
daoulasensis , Protaster 261
darwini, Echinasterella (?) 200
decheni, Eoluidia 262
decheni, Eophiurites 262
decheni, Protaster 248
decheni, Tremataster (?) 247, 248
demissus, Onychaster 270,274
development, abnormal 101
geological, of Stelleroidea 27, 28
Devonaster 40, 44,
46, 47, 49, 50, 74 , 76, 77, 9 7 , 129, 130, 133
D. chemungensis 101
D.eucharis 29,37,98
devonica, Palxnectria 155
devonicus, Palasteriscus 200
difflcilis, Tremataster 247
disk 14,213,225
how enlarged 32
disk accessory plates 34, 46
disk ambitals 13
disk inframarginals 15
INDEX.
305
disk marginals 14
disk plates 14
primary 49,179,184,190,191
disk supramarginals 15
dispar, Xenaster 130, 131
dorsal, see abactinal.
dorsals, see radials.
dorsal shield 213
dorso-laterals, see accessory plates.
drevermanni, Miomaster 96
dubius, Mesopalseaster (?) 75, 85
dubius, Palxaster 85
dyeri, Palxaster 120
dyeri, Petraster 121
dyeri, Promopalaeaster 104, 105, 120
Echinaster 45
Echinasterella 199, 200
E. (?) darwini 200
E . sladeni 200
EcMnasterias 38,207,209, 211
E. spinosus 211
echinatus, Squamaster 249
Echini, radii of 35
Echinodiscaster 38,40,207,209,211
E. multidactylus 211
Echinodiscites 211
Echinodiscus 211
E. multidactylus 211
Echinostella 38,207,209,212
E. traquairi 212
Eifelaster 128, 133
E. follmanni 134
eifelense, Aspidosoma 244
eifelensis, Encrinaster 244
elegans, Protaster 221
elegans, Tseniaster 219,221
elegans, Xenaster 130, 131
elizae, Uranaster 155
Encrinaster (part) 224
Encrinaster 43, 45, 226, 241
E. arnoldi....; 241,243
E. eifelensis 244
E.goldfussi 243
E. (?)grayae 245
E.petaloides 243
E. petaloides goslariensis 243
E.pontis 244
E. roemeri 242, 244
E. schmidti 244
E. tischbeinianus 244
Encrinasterix (part) 215
Encrinasteridse 215, 241
Eoactis 173,178
E. simplex 186
Eoluidia 262
E. decheni 262
Eoluidiidse 246,262
Eophiura 216,222,223,259
E . bohemica 222
Eophiuridx 262
Eophiurites 262
E. decheni 262
Eospondylus 262, 2 63
E . primigenius 263
Etheridgaster 170
Page.
E. clarkei 172
eucharis, Devonaster 29,37, 98
eucharis, Palxaster 93
eucharis, Xenaster 93
Eucladia 275, 276
E. (?)beecheri 278
E. johnsoni 276
E. woodwardi 276, 277
Eucladiidge 276
Eugaster 237
E. concinnus 239
E. logani 238
Eugasterella 235, 237, 259
E. (?) concinna 238, 239
E . logani 237, 2 3 8
Euophiuroidea 213
EuryalQ annulatum 20
Euthemon.....' 275,276, 279
E. igerna 279
evolution of Asteroidea .% 30
exculptus, Palxaster 117
exculptus, Promopalseaster 104, 105, 117
eye-plate, see ocular plate.
ferox, Palaeodiscus 23
filiciformis, Helianthaster 159, 1 60
fimbriata, Palxasterina 204
fimbriaid, Protaster 231
fimbriata, Protasterina 231
fimbriatus, Schoenaster 204
flnei, Mesopalseaster 75, 81
fine i Palxaster 81
flexilis, Onychaster 269, 270
flexuosus, Alepidaster 230, 231
flexuosus, Protaster 231
flexuosa, Protasterina 231
follmanni, Eifelaster 134
follmanni, Palasterina 157
follmanni, Pseudopalasterina 156, 157
forbesi, Hallaster 254, 2 66
forbesi, Protaster 255
formosus, Compsaster 193
Furcaster 248,259, 261
F. (?) daoulasensis 261
F. palseozoicus 259, 261
Furcasteridx 248
genitals, see interradial plates.
genitals in Echini 35
geological development of Stelleroidea 27, 28
geological distribution of Stelleroidea 28
giganteus, Australaster 73, 171
giganteus, Cheiropteraster 202
giganteus, Monaster 73
giganteus, Palxaster ( Monaster) 73
girvanensis, Urasterella 167, 175, 186
goldfussi, Aspidosoma 243
goldfussi, Encrinaster 243
Goniaster 61
grandis, Agalmaster 132
grandis, Sundenbachia 235
grandis, Palseophiomyxa 234, 286
grandis, Stenaster 180
grandis, Urasterella 174, 175, 180
granti, Mesopalaeaster 75, 89
granti, Palxaster 89
granuliferus, Alepidaster 229, 280
306
INDEX.
Page.
granuliferus, Protester (f) 230
granulosus, Palxaster, Hall 112
granulosus, Palxaster, Meek 109
granulosus, Promopalaeaster 104, 1 1 2, 113
graspers 33
grayx, Aspidosoma 245
grayse, Encrinaster (?) 245
grayi, Lepidast*r 158
gregarius, Aganaster 264
gregarius, Alepidaster 264
gregarius, Protester (/) 264
Gregoriura 216,233
G. spryi 234
gregoryi, Palseosolaster 209
groomi, Protaster 226, 227
gyalum, Helianthaster 160,210
gyalum, Palseosolaster (?) 210
Hallaster 248, 254
H. forbesi 254, 255
Harris, I. H., gift oietarflshes by 9
Jiarrisi, Palxaster 180
Helianthaster Clarke (part) 209
Helianthaster Roemer 38,
40,45,157,158,159,209,211
H. flliciformis 159,160
H. gyalum (part) 160, 210
H. rhenanus 159
H.roemeri 209,211
H. n. sp . Clarke 160
Helianthasteridx 157
Helianthasterinse 157
Heliaster 38,208
hirudo, Palxaster 188
Urudo, Uraster 188
hirudo, Urasterella 175,188
Hisingeraster 148
H, antiqua 149
Hudsonaster 31,32,34,36,
38, 39, 40, 41, 42, 44,45, 47, 48, 49, 52, 68,
69, 75, 105, 135, 140, 158, 165, 179, 194, 196
H.batheri 55,65,167
H. incomptus. 34,36,40,55,61
H. matutinus 34,55, 57, 139, 141, 166
H. milleri ..... 55,60
H.narrawayi 43,50,55,59
H. rugosus 36, 55, 64
Hudsonasteridse] 52, 53,162
huxleyi, Stenaster 182
huxleyi, Urasterella 175, 182
igerna, Euthemon 279
imbricatus, Palxaster 169
imbricatus, Tetraster (?) 168, 169
incomptus, Hudsonaster 34,36,40,55, 61
incomptus, Palxaster. 61
inframarginalia 16, 34, 48, 152, 194
inframarginal plates 16, 33, 174, 179, 185
Interact] nals 13
interbrachial adambulacrals 1 5, 126
interbrachial areas and arcs.. . . 14, 60, 138, 185, 194
interbrachial marginals 15
intennarginal plates, see ambital plates,
intermarginals, see ambital.
intermedius, Agalmaster 133
intermedius, Mesopalseaster 75, 79
interadial plates 15, 32
Jaekelaster 40, 192
Page.
J. petaliformis 192
jahni, Bohemura 223
jamesi, Palxaster 146
jamesi, Palasterina (t) 146
johnsoni, Eucladia 276
kinahani, Palasterina 155
kinahani, Uranaster 154, 155
Labidaster 208
lanceolatus, Mesopalaeaster (?) 75, 82
Lapworhtura 248, 250, 254
L. cylindrica 220
L. miltonl 250,261,253
L. sollasi. 251
L. (?)sp 251
Lapworthuridae 246, 248
lateral shields 214
latiscutetus, Coelaster 96
latiscutatus, Spaniaster 96
laxata, Schuchertia 196, 198
legrandensis, Schoenaster (?) 206
Lepidaster 38,40,158,160
L . grayi 158
Lepidasteracea 53
Lepidasterella 38, 40, 1 60
L.babcocki 160
Lepidasteridae 52,53,157
leptosoma, Protaster 237
leptosoma, Rhodostoma 237
leptosoma, Sturtzura 236, 237
leptosomoides, Sturtzura 236, 237
Lindstromaster 138, 140, 148, 154, 156
L. antiquus 1 4 9 , 153
Lindstromasterinx 138
logani, Eugaster 238
logani, Eugasterella 237, 288
longibrachiatus, Palxaster 115
Loriolaster 199,200, 201, 202,252
L. mirabilis 201
lymani, Ophiurina 247
Lysophiurae 215
madreporite 15,39,40,209,211,214,222,223,226
magniflcus, Promopalaeaster 42,
104,105,106,113,122
margaritetus, Xenaster 131,132
margaritatus, Xenaster 129, 130, 181
marginalia, see marginal plates.
marginal plates 1 6,33, 43, 135, 152, 213
marstoni, Palxocoma 253
marstoni, Sturtzaster 253
matutina, Asterias 57
matutinus, Hudsonaster 34, 55, 6 7,139, 141, 166
matutinus, Palxaster 67
meafordensis, Tseniaster 221
measurements 15
median dorsals, see radials.
Medusaster 38,207,212
M. rhenanus 212
Mesopaleeaster 28, 39, 41, 44, 45, 46, 47, 49,
50, 56, 74,95,98, 105, 126, 130, 135, 158
M.(?) acuminatus 75, 93
M. (? ) antiquus 75, 8 6
M.beUulus 75,91
M. caractaci 75, 92
M.(?) cataractensis 75,89
M.(?)clarki 75,94
M.(?)dubius 75,85
INDEX.
307
Page.
M.finei 75,81
M. grant! 75,89
M. intermedius 75, 79
M.(?) lanceolatus 75, 82
M.(?) parviusculus 75, 87
M. proavitus 75, 83,85
M.shafferi 44,75,77
M.(?) wilberanus 75, 84
Mesopalseasterinae 52, 74,162
miamiensis, Alepidaster 230, 238
miamiensis, Anorthaster 127
miamiensis, Palseaster 127
miamiensis, Protaster 233
miellensis, Agalmaster 132
milleri, Hudsonaster 55, 60
miltoni, Lapworthura 250, 251, 253
mittoni, Protaster 251
minveri, Sympterura. . : 256
Miomaster 43,50, 74, 96
M. drevermanni 96
Miospondylus 262, 263
M. rhenanus 263
mira, Calliasterella 191
mirabilis, Loriolaster 201
mirus, Calliaster 190
mitchelli, Sturtzaster (?) 254
Monaster Gregory 49, 72
Monaster Etheridge 40, 1 70
M. Clarkei 171, 172
M. giganteus 73
M . stutchburii 73
Monasteridae 162, 163, 1 70
montana, Urasterella 175, 189
montanus, Asterias 189
montanus, Palseaster 189
montanus, Schoenaster (?) 43, 207
mouth angle plate, see oral armature.
multidactylus, Echlnodiscaster 211
multidactylus, Echinodiscus 211
multlrayed starfishes 38, 40, 157, 207
narrawayl, Hudsonaster 43, 50, 55, 69
narrawayi, Protopalseaster 59
neglecta, Palseura 223
Neopalseaster 40, 43, 48, 49, 57, 69, 98, 134
N. crawfordsvillensis 135, 136
Neopalseasteridse 52,53, 184
niagarensis, Palaeaster 68, 69
obtusus, Ophiura 25
obtusus, Palssaster 167
obtusus, Protaster (Ophiura) 25
obtusus, Stenaster (?) 165, 167
obtusus, Uraster 167
occurrence of starfishes 28
ocularplate 16,34,48,134,135
oculars in Echini 35
ontogeny of Stelleroidea 36
Onychaster 268
O.asper 270,272
O. barrisi 270, 272
O. confragosus 270, 273
O. demissus 270, 274
O.flexllis 269,270
Onychasteridse 268
Ophiocistia 275
Ophio-Encrinasterix .' 215
OpJiiopege 263
Page.
Ophiura ciliaris 268
O. obtusus 25
O. (?)ramosa 22
O . rhenana 263
O . salteri 227
O. schlotheimii 13
Ophiurasterix 215,246
Ophiurella 263
0- primigenia 263
Ophiurina 246, 260
O. lymani 247
Ophiurinidse 246
Ophiuroidea 41, 42, 43, 2 1 2 , 258, 267
geological distribution of 28
number of, in Paleozoic 27, 28
oral, see actinal.
oral angles 16
oral armature 16, 43, 162, 172, 210, 242
oral skeleton, see oral armature.
ordinaria, Schuchertia 196, 199
ossicles, see plates.
Palseaster 10,
40, 41, 44, 49, 56, 66, 67 , 75, 98, 104, 135, 158
Palseaster (Monaster) Etheridge (part) 72,170
P. (Argaster) antiqua 86
P. antiquata 109
P. aniiquus 86
P. asperrima 187
P. caractad 92
P. clarkana 61,63
P. clarke i De Koninck .- 172
P. clarkei Miller 61
P.darki : 94
P. coronella 167
P. crawfordsvillensis 136
P.dubius 85
P.dyeri 120
P. eucharis 98
P.exculptus 117
P. fin* 81
P. (Monaster) giganteus 73
P. grant i 89
P. granulosus Hall 112
P. granulosus Meek 109
P.Mrrisi 180
P.Mrudo 188
P. imbricatus 169
P. incomptus 61
P.jamesi 146
P. longibrachiatus 115
P. matutinus 57
P. miamiensis 127
P. montanus 189
P. niagarensis 68, 69
P . obtusus 167
P. parviusculus 87
P.pukJiellus 178
P. pygmaea. 23
P.ruthveni 187
P.sJiafferi - 77
P. simplex. 61, 62
P.speciosus 109
P. spinulosus - ^
P. squamatus 2 s
P. (Monaster) stutchburii 73
P. wilberanus 84
308
INDEX.
P. ? wilsoni 106
P.wykoffi 119
Palaeasteridse 52, 53, 66
Palxasterina 138, 150
P. antiqua 149
P. approximata 142
P. bonneyi. 153
P. fimbriata 204
P. primxva 153
P. speciosa 142
Palxasterinx 66
Palxasterinidx 138
Palxbrisingidx 199
Palxchinasteridx 199
Palxgoniasteridx (part) 66,241
Palxgoniasteridx 128
Palxnectria 155
P. devonica 155
Palxocoma Miller (part) 239
Palxocoma Salter 252
Palxocoma (Bdellacoma) 254
Palxocoma (Rhopalocoma) 254
P. colvin i 253
P. cygnipes 253
P. cylindrica 220
P. marstoni , 253
P. princeps 240
P. pyrotechnica 254
P. spinosa 219
P. vermiformis 254
Palaeodiscus ferox. . 23
Palseophiomyxa. 216, 234
P. grandis. 234, 235
Palxophiomyxidx 215, 234
Paiseophiura 235
P. simplex 235
Palxophiuridx 215,216, 235
Palaeosolaster 38, 40,207, 209, 211
P. gregoryi 209
P. (.)gyalum 210
P.rcemeri 211
Palseosolasteracea 163
Palseosolasteridse 162, 163, 20 7
Palxospondylidx 248
Palxospondylus 261
Palaeostella 51,138,155
P.solida 156
palaeozoicus, Furcaster 259, 261
Palxspondylus 261
P.zitteli 262
Palaeura 216,223
P.neglecta 223
Palasteracanthion 173
Palasterina Billings (part) 195
Palasterina Salter 40,
49, 130, 138, 140, 1 50, 154, 156, 196
P. bonneyi 151,152,153
P.follmanni. , 157
P.(f)jamesi. 146
P. kinahani. 155
P. primaeva 149, 151, 152, 153
P. (?) ramseyensis 152, 154
P. rigidus 141
P. rugosa 64
P. stellata 196
Palasterinacea. . . 53
Page.
Palasterinidae 52,53, 138, 196
Palasteriscidae 162, 163, 199
Palasteriscus 40, 199, 200, 252
P. devonicus 200
Palastropecten 248, 261
P. zitteli 262
Palmipes antiqua 149
papulae 16
parviusculus, Mesopalseaster (?) 75, 87
parviusculus , Palxaster 87
parvulus, Trimeraster 134
paxillae 116,156
peculiaris, Cholaster 266
pedicellariae 16
perfectus, Siluraster 66
perradial 17
petaliformis, Jaekelaster 192
petaloides, A spidosoma 243
petaloides, Encrinaster 243
petaloides goslariensis, A spidosoma 243
petaloides goslariensis, Encrinaster 243
Petraster 39,
44,45,46,47,51,138,149,154,156,196
P. (?) americanus 139,146
P. (?) antiqua 86
P. bellulus 91
P.dyeri 121
P. rigidus (part) 57
P.rigidus 139,141
P. smythi 139,147
P. speciosus 47, 139, 142
P. wilberianus 84
petri, Protaster 25
Phanerozonia 80,
38,39, 40, 41, 43, 44, 45, 46, 48, 51
phylembryo of Stelleroidea 34
Pisaster 38
plates 17
plumiformis, Trichotaster 26, 38
podia and podial openings 18,
17,33,38,39,209,213
pontis, Aspidosoma? 244
pontis, Encrinaster 244
prenuntius, Promopalseaster 104, 1 7
preservation of starfishes 28
primxva, Asterias 167
primxva, Palxasterina 153
primaeva, Palasterina 149, 151, 152, 153
primxvus, Uraster 153
primary inframarginal 15
primary radials 1 7, 32
primary skeleton 13, 15, 1 7 , 31, 32, 34, 56, 191
primary supramarginal 15
primigenia, Ophiurella 263
primigenius, Eospondylus 263
primitive starfish, most 31, 33
primordialis , Asterias 108
primus, Protasteracanthion 188
princeps, Palxocoma 240
princeps, Ptilonaster 240, 259
proavitus, Mesopalaeaster 75, 83, 85
Promopalaeaster 28,
39,41,42,44,45,46,47,50,56,76,98, 102, 126
P. bellulus 104,105,118
P.dyeri 104,105,120
P.exculptus 104,105,117
INDEX.
309
Page.
P. granulosus 104, 112,113
P. magnificus 42, 104, 105, 106, 113, 122
P. prenuntius 104, 107
P. speciosus 104, 105, 109, 113
P. sp. undet 104, 108, 109
P, spinulosus 44, 104, 105, 116
P. wilsoni 104, 106
P. wykoffi 42, 104, 119
Promopalaeasteracea 53
Promopalsesteridse 52, 53, 73
Promopalaeasterinse 52, 74, 102
Protaster Hall (part) 254
Protaster Forbes 216,217, 224, 229
P. (?) barrisi 272
P. biforis 226, 722
P. brisingoides 236
P. daoulasensis 261
P.decheni 248
P.elegans * 221
P. fimbriata 231
P. flexuosus 231
P.forbesi 255
P. (?) granuliferus 230
P. (?) gregarius '264
P.groomi 226,227
P. leptosoma 237
P. miamiensis 233
P. miltoni 251
P. (Ophiura) obtusus 25
P.petri 25
P. (?) salted 226,227
P.sedgwicki 225,226,243
P. (?) steUifer 226, 228
P. (?) whiteavesianus 226, 227
Protasteracanfhion 173, 178
P. primus 188
Protasteridae 215
Protasterina 228
P. fimbriata 231
P.flexuosa 231
Protoeuryale 25
Protopalxaster 53, 56
P. caractad 92
P. narrawayi 59
Protophiurex 246
Protophiuroidea 213
proximal inframarginals 14, 16
proximal radials 17
proximal supramarginal 15
Pseudopalasterina 138, 156
P.foUmanni 156,157
Ptilonaster 235,289
P. princeps 240, 259
pulchella, Urasterella. 175, 178
pulchellus, Palxaster 178
pulchellus, Stenaster 178
Pycnopodia 38, 42, 207, 208
pygmaea, Palaeaster 23
pygmaeus, Ataxaster 161
pyrotechnica, Palxocoma 257
pyrotechnica, Rhopalocoma 254
radial accessory plates 13
radialia 17,34,35,46
radicle of asterids 34
ramosa, Ophiura (?) 22
s, Palasterina (?) 152,154
Page.
ray ambitals 13
rays 17,38,44,48,207,213,223
regeneration of. 185
regeneration 87, 185
respiration, see papulae and csecal pores.
rhenana, Asterias (Archxasterias) 131
rhenana, Ophiura 263
rhenanus, Helianthaster 159
rhenanus, Medusaster 212
rhenanus, Miospondylus 263
rhenanus, Xenaster (?) 130, 1 8 1
Rhenaster 43, 128, 188
R. schwerdi 133
Rhodostoma 236
R. leptosoma 237
Rhopalocoma 248,252, 254
R. pyrotechnica 254
rigidus, Palasterina 141
rigidus, Petraster 57
rigidus, Petraster 139, 141
Roemeraster 173, 177
R. (?) acuminatus 93
R. asperula 188
Roemerasterinx 172
roemeri, Aspidosoma 244
roemeri, Encrinaster 242,244
roer eri, Helianthaster 209, 211
roemeri, Palseosolaster 211
rugosa, Palasterina 64
rugosus, Hudsonaster 36,55, 64
ruthveni, Palxaster 187
ruthveni, Uraster. 187
ruthveni, Urasterella 174, 175, 18 7
SaUeraster 173,178
S. asperrimus 187
salted, Ophiura 227
salted, Protaster (?) 226,227
salted, Stenaster 164, 166
schlotheimii, Acroura (Ophiura) 18
schlotheimii, Ophiura 18
schluteri, Astropecten (?.) 161
schmidti, A spidosoma 244
schmidti, Encrinaster 244
Schcenaster 202
S.fimbriatus 204
S. (?) legrandensis 206
S. (?) montanus 43,207
S. (?)n.sp 206
S. ( ?) wachsmuthi 205
Schcenasteridse 162,163,202
schohariae, Taeniaster 219,220
Schondorf, on revision of German starfishes. . 11
Schuchertia. 51, 140, 152, 194, 195, 252
S. laxata 196, 198
S.ordinaria 196,199
S.stellata 195,196
Schuchertiacea 163
Schuchertiidae 162,163,194
schwerdi, Rhenaster 133
secondary skeleton, see secondary plates.
sedgwicki, Protaster 225, 226, 243
selwyni, Urasterella 175,188
shafferi, Mesopalajaster 44, 75, 77
shafferi, Palxaster 77
Siluraster 49,53 65
S.perfectus 66
310
INDEX.
simplex, Eoactis . -. 186
simplex, Palxaster 61, 62
simplex, Palaeophiura 235
simplex, Xenaster 69
skeleton, secondary, see accessory plates.
sladenl, Echinasterella 200
smythi, Petraster 139, 147
Solaster 38
solida , Palseostella 156
sollasi, Lapworthura 251
Spanlaster 39,43,47,49,50,74,95
S. latiscutatus 96
speciosa, Palxasterina 142
speciosus Palxaster 109
speciosus, Petraster 47, 139, 142
speciosus, Promopalaeaster 104, 105, 109,113
spines, see paxillae and pedicellarise.
spinosa, Palxocoma 219
spinosissima, Asterias 19
spinosity 37,41
spinosus, Echinasterias 211
spinosus, Tgeniaster 216, 217,218, 219
spinulosus, Palxaster 115
spinulosus, Promopalaeaster 44, 104, 105, 115
spryi, Gregoriura 234
Squamaster 248,249
S. echinatus 249
squamatus, Palaeaster 23
stalk of primitive starfishes 83, 34
starfish, most primitive 31 , 33
origin of graspers 33
origin of wriggling type 32
radicle 34
stellata, Palasterina 196
stellata, Schuchertia 195, 1 96
stellata, Trentonaster 196
SteUeroidea
geological development 27,28
geological distribution 28
ontogeny 36
phylembryo 34
typembryo 8 5, 36
stellifer, Protaster (?) 226, 228
Stenaster (part) 173
Stenaster 31,41,42,43,50,68,163,168,212
S. (?) confluens 165,167
S. (?) coronella 165,167
S.grandis 180
S.huxleyi 182
S. (?) obtusus 165, 167
S.pulcJiellus 178
S.salteri 164,165
Stenasteracea 163
Stenasteridae 162, 168
stone-canal, see madreporite.
Strep tophiurse 215, 249
Stiirtzaster 248,252
S.colvini 253
S.cygnipes 253
S. marstoni 253
S. (?) mitchelli 254
Stiirtzura 235,286
S. brisingoides 236
S.leptosoma 236,237
S. leptosomoides 236,237
Page.
stutchburii, Australaster (?) 73
stutchburii, Monaster 73
stutcJiburi i, Palaeaster ( Monaster} 73
supramarginalia 15, 34,35, 46
Sympterura ; 248, 256
S. minveri 256
syngnaths 218,222,224,242,256,267
Tseniaster 216,218
T. australis 236
T. cylindricus 216,217,218,219,220
T.elegans 219,221
T. meafordensis 221
T. schohariae 219, 220
T. spinosus 216,217,218, 219
Txniasteridx 172
Txniura 216
T.cylindrica 220
tenuiradiatus, Coelaster 19
terminals, see ocular plates.
terminology of Asteroidea 13
Tetraster 31,41,42,43,50,164,165,167,212
T. asperrimus 187
T. (?) imbricatus 168,169
T . sp . ind . Nicholson and E theridge 186
T. wyville-thomsoni (part) 65
T. wyville-thomsoni 168
t ischbeinianum, Aspidosoma 244
tischbeinianus, Encrinaster 244
torus 1 6, 43, 54, 174
traquairi, Echinostella 212
Tremataster 246, 247
T. (?) decheni 247, 248
T. difficilis 247
Trentonaster 195
T. stellata 196
Trichotaster plumiformis 26, 38
Trimeraster 44,45,50, 128, 184
T. parvulus 134
trivium 14
Trochitaster, see Trichotaster.
typembryo of Stelleroidea 85, 36
ulrichi, Urasterella 37, 43, 45, 49, 175, 183
Uranaster 61,138,154
U. elizse 155
U. kinahani 154, 155
Uraster hirudo 188
U. obtusus 167
U. primxvus 153
U.ruthveni 187
Urasterella Sturtz 163
Urasterella McCoy 40,
41,42,45,47,49,50,69,164,178,194,212
U. (?)asperrima 175,187
U. asperula 175, 188
U. (?) constellata 175, 187
U. girvanensis 167, 175, 186
U. grandis 174,175,180
U. hirudo 175,188
U.huxleyi 175,182
U. montana 175,189
U. n. sp 189
U.pulchella 175,178
U . ruthveni 174, 175, 1 8 7
U.selwynl 175,188
U. ulrichi 37,43,45,49,175,188
INDEX.
811
Page.
Urasterellacea 163
Urasterellidse 162,163,172
ventral, see actinal.
ventral shield 213
vermiformis, Bdellacoma 254
vermiformis, Palxocoma 254
vertebral ossicles 18,213,257,267,269
wachsmuthi, Schoenaster (?) 205
whiteavesianus, Protaster (?) 226, 22 7
wilberanus, Mesopalseaster (?) 75, 84
wilberanus, Palseaster 84
Wilberianus, Petraster 84
wilsoni, Palseaster (?) 106
wilsoni, Promopalseaster 104, 106
woodward!, Eucladla 276, 27 7
wrigglers 32,33,38
Page.
wykoffi, Palseaster 119
wykoffi, Promopalseaster 42, 104, 1 1 9
wyville-thomsoni, Tetraster 65
wyville-thomsoni, Tetraster 168
Xenaster 40,44,45,46,47,49,50,98,128,129
X. dispar 130,181
X.elegans 130,181
X. eucharis 98
X. margaritatus Follmann 131
X. margaritatus Simonovitsch (part) 131, 132
X. margaritatus Simonovitsch 129,130, 181
X. (?)rhenanus 130,181
X. simplex 96
Xenasteridse 52,53, 128
zitteli, Palseos'pondylus 262
zitteli, Palastropecten 262
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