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UNIVERSITY OF IOWA MONOGRAPHS
Bulletin from the Laboratories of Natural History
VOLUME VII 1915-1918
UNIVERSITY OF IOWA
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CONTENTS
VOLUME VII
VERRILL, A. E. Report on the starfishes of the West Indies, Florida, and Brazil.
SHIMEK, Bonumiu. The plant geography of the Lake Okoboji region.
WickHaAM, H. F. New fossil Coleoptera from the Floris- sant beds.
Collected studies and reports.
Cuark, H. L. Report on the Crinoidea and Echinoidea collected by the Bahama expedition from the Univer- sity of Iowa in 1893.
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NEW SERIES NO. 92 es He ae MARCH 20, 1915
BULLETIN OF THE STATE |
_ UNIVERSITY OF IOWA
BULLETINS FROM THE LABORATORIES OF NATURAL HISTORY |
“VOLUME VII ite NUMBER 1
Report on the Starfishes of the
West Indies, Florida, and Brazil
INCLUDING THOSE OBTAINED BY BAHAMA EXPEDITION FROM THE / UNIVERSITY OF IOWA IN 1893 \
BY ADDISON EMERY VERRILL
PROFESSOR EMERITUS, YALE UNIVERSITY NEW HAVEN. CONNECTICUT
PUBLISHED BY THE UNIVERSITY, IOWA CITY
ISSUED TWENTY-ONE TIMES DURING THE: ACADEMIC YEAR; MONTHLY FROM OCTOBER TO JANUARY, WEEKLY FROM FEBRUARY .TO JUNE. ENTERED : AT THE POST OFFICE AT IOWA CITY AS SECOND CLASS MAIL MATTER —
ee * aaa
IN THE SERIES OF RESEARCH BULLETINS OF THE UNIVERSITY
BULLETINS FROM THE LABORATORIES OF NATURAL Plo PORY
EDITORIAL CHARLES C. NUTTING
GEORGE F. KAY - BOHUMIL SHIMEK -
VOLUME VII
STAFF
- - - = Zoology - - - + Geology =) sue) Botany
NUMBER |
CONTENTS
Report on the Starfishes of the West Indies, Florida, and Brazil, including those obtained by the Bahama expedition from the University of Iowa in 1893.
ADDISON EMERY VERRILL, Professor Emeritus, Yale University
PUBLISHED BY THE UNIVERSITY IOWA CITY, IOWA
Since several years have elapsed since the publication of the last re- port on material secured by the Bahama Expedition of 1893 from the State University of Iowa, it has been deemed advisable at this time to give a list of papers in which the results of the expedition are discussed, together with the full tities, authors and dates of issue.
Contribution to the anatomy of the Gorgonidae, together with observa- tions on living specimens. Pages 97 -160, and 10 plates. ©. C. Nur- TING. Bull. Lab. Nat. Hist., S. U. I., Vol. I, No. 2. Nov., 1889
(This paper was based on material secured during a preliminary Expedition to the Bahamas by the author in 1888, and therefore should find a place here.)
Narrative and preliminary report of the Bahama Expedition. 251 pages, and 20 full-page illustrations. C. OC. NurTtnc. Bull. Lab. Nat. Hist., S. U. I., Vol. III, Nos. 1 and 2. Jan., 1895
The mollusks and brachyopods of the Bahama Expedition of the State Unwersity of Iowa. 20 pages, and 1 plate. Wm. H. Datu. Bull. Lab. Nat. Hist., S. U. I., Vol. IV, No. 1. Dec., 1896
Notes on the Hymenoptera collected by the Bahama Expedition from the State University of Iowa. 4 pages. WILLIAM H. ASHMEApD. Bull. Lab. Nat. Hist., S. U. P Vol. Ve.sNiow de Dec., 1896
Report on the fishes collected by the Bahama Expedition, of the State University of Iowa, under Professor C. C. Nutting. 20 pages. 4 plates. SaMUEL GARMAN. Bull. Lab. Nat. Hist. iS]. ie) Vol. Tv, No. 1. Dee., 1896
Report on the Actinaria collected by the Bahama Expedition of the State University of Iowa. 25 pages, 3 plates. J. PLAYFAIR Mc- Mourricu. Bull. Lab. Nat. Hist., 8. U. 1, Viol: (iveuNo. 3 June, 1898
The Brachyura of the expedition to the Florida Keys and the Bahamas im 1893. 65 pages. 9 plates. Mary J. RATHBUN. Bull. Lab. Nat. Hist. S. U. 0, Vol. TV, No. 3) Sune asos
Report on the Ophiuroidea collected by the Bahama Expedition in 1893. 88 pages. 8 plates. A. E. VERRILL. Bull. Lab. Nat. Hist., SUS Vol ING. in Sept., 1899
American Hydroids. ©. C. Nurtie. Smithsonian Institution, U. S. Nat. Museum, Special Bulletin No. 4, Part 1, issued 1900; part 2, issued 1904; part 3, issued 1915 (The writer, in undertaking to prepare this monograph, agreed to publish in
it all descriptions of hydroids secured by the Bahama Expedition instead of issulng a separate Report on that group).
West Indian Starfishes found at a depth of 40 fathoms or less, includ- mg a report on material secured by the Bahama Expedition from the State University of Iowa. 230 pages, and 29 plates. A. E. VERRILL. Bull. Lab. Nat. Hist., S. U. I. Vol. , No. » 1915
Descriptions of about 70 new species of marine invertebrates from the Bahama expedition collections are found in the papers listed above. Some important groups, such as the Echinoidea, Vermes, and Corals are still to be reported on; but it is hoped that the series will be com- plete at a reasonably early date.
The material upon which these papers are based is in the museum of natural history of the University. C. C. NuTtTine
INTRODUCTION
That portion of this memoir relating to the special collection of the Bahama Expedition was nearly completed and the plates were made more than ten years ago, but its publication was de- layed for several reasons, but largely because of the desirability of publishing, in advance of it, a much more extensive work, by the writer, on the starfishes of the North Pacific coast of Amer- ica (Harriman Series) then in type, and in which the classifica- tion and many of the families and genera are revised.
Unavoidable and unexpected delay of the larger work had like- wise delayed this. In the meantime, owing to the lack of any general work on the shallow water starfishes of the West Indies, and to the very scattered and unsatisfactory condition of the literature relating to them, it has been thought desirable to en- large the report so as to include descriptions of all the families, genera, and species known to occur in the West Indian fauna, in waters less than 150 fathoms deep.
The West Indian faunal region is here extended so as to in- clude the entire region from northern Florida to Rio de Janeiro, Brazil, and to include, also, the Bermudas and the whole of the Gulf of Mexico.
As compared with other tropical seas, the West Indian starfish fauna is very poor in littoral and shallow water species. It is rarely that more than twelve to fifteen species can be found by careful shore and reef-collecting at any one place.
Professor Clark (1898), after two seasons of shore and reef collecting of the echinoderms at Jamaica, enumerated only four- teen species of starfishes, although he visited both sides of the island, as well as outlying reefs. The entire number of littoral and very shallow water species probably does not exceed twenty to twenty-two over large areas.
This is not more than one-half or one-third the number to be found in many other similar regions.
+ NATURAL HISTORY BULLETIN
At depths of 100 to 1,000 fathoms, however, the starfishes are very numerous, both in species and individuals.
The most prolific zone for starfishes, as well as for most other groups, is on the continental border in 100 to 500 fathoms.
The Bahama Expedition dredged at a number of localities be- tween 100 and 200 fathoms, especially at the famous erinoid ground, off Havana, which isa locality very favorable for marine invertebrates of many kinds. Therefore, the collection obtained includes an unusual number of species, considering the small number of dredgings made, and the smallness of the equipment, as compared with that of the ‘‘Blake’’ or the ‘‘ Albatross.”’
A few of the more common West Indian and conspicuous shallow water starfishes were mentioned by the early writers, such as Maregraft (1648) ; Sloane (1725) ; Linck (1733) ; Seba (1761), and others.
This was especially the case with the very large and conspic- uous species, the ‘‘large warted starfish’’ of Brown, 1756, now known as Oreaster reticulatus, which is very common in many places in very shallow water, and therefore sure to attract the at- tention of the most casual observers.
Yet the number of species actually known to occur in the West Indies was very small up to the period of Gray (1840) and of Miller and Troschel (1842).
Linné referred only two or three of his species to the West Indies and some of those doubtfully. Lamarck (1816) recog- nized only two or three as West Indian. Thomas Say (1825) recorded six species from the southern coast of the United States, five of which he described as new. All of these, except two, are also West Indian species.
Gray, 1840, gave the West Indian region as the source of eight of his species. Miiller and Troschel (1842) recorded but five as West Indian, though they described others from this fauna whose origin was unknown.
The first attempt to give a critical synopsis of the West Indian species was made by Prof. Christian Liitken,! a most careful
1 Dr. Christian Frederik Liitken, professor of the University of Copen-
hagen; lieutenant in the war of 1849-50; died February 6, 1901, aged 73 years.
His works on the starfishes and ophiurans of northern Europe, Green-
WEST INDIAN STARFISHES 4)
writer, in 1859. He then described with care thirteen species, in the Museum of Copenhagen, mostly from the Lesser Antilles. Six were described as new, mostly with doubt. At least four of these proved to be synonymous with those of Gray and other pre- vious writers, who had poorly described them. His careful de- scriptions are, however, invaluable for the determination of most of the early known species, and mark an era in work of this kind.
In 1867 the present writer published a faunal list of West Indian species, for comparison with that of the Panama region. This list included twenty-four nominal species, but at least five of these have since become synonymous. On the other hand, two or three previously described species were omitted. Thus the number of species of this fauna actually known at that time was about twenty.
The beginning of dredging in somewhat deep water was made by Pourtales in 1867 and 1868, but only one additional species of starfish was then added by him.? Subsequently, these investi- gations were carried on by the ‘‘ Bibb,’’ in 1869, and the ‘‘ Hass- ler’’ in 1872, under the direction of Pourtales, and in 1872, un- der Dr. Wm. Stimpson by the ‘‘Bibb’’ and ‘‘Bache.’’ But no separate lists of the starfishes taken in those years has been pub- lished. Some are included in the ‘‘Revision’’ by Perrier, 1875 and 1876 (see Bibliography) ; others in his 1884 report on the Blake starfishes.
Mr. Alexander Agassiz carried out his extensive dredgings in the Caribbean Sea and Gulf of Mexico, etc., on the ‘‘Blake’’ in 1877, 1878, 1879, and 1880. About 350 stations were occupied by the Blake.
These explorations* added numerous species of starfishes to the fauna, not only from great depths, but also many from com- paratively shallow water.
The starfishes thus obtained by Agassiz were described by Per- land, and both coasts of tropical America are classical and indispensable. He also wrote on fishes, ascidians, actinians, and other subjects.
For many years he was a most esteemed correspondent of the writer.
2 Catalogued by Agassiz as Pteraster militaris, now called P. caribbeus Perrier.
3 A very complete and interesting account of these explorations is given by Mr. A. Agassiz in ‘‘ Three Cruises of the Blake.’’ Vols. I, II. 1888.
6 NATURAL HISTORY BULLETIN
rier in several articles from 1880 to 1894, but most were brought together in his memoir of 1884.
Dredging operations of great extent were carried on by the U. S. Fish Comm. steamer ‘‘ Albatross,’’ in the West Indian re- gion during the winters of 1884, 1885, and 1886.* Many species of starfishes were then obtained, including a considerable num- ber of new species. But those collections have not yet been sys- tematically reported upon, though the writer described and fig- ured a number of the species in 1899.
According to Mr. A. Agassiz (1888, p. 4), the Blake took 54 species of starfishes, of which 46 were considered new species by Perrier. The actual number has since proved to be slightly larger.
The Bahama Expedition, from the University of Iowa, dredged mostly in waters less than 200 fathoms deep. The most interest- ing were off Havana, on the crinoid grounds, where a number of additions to the starfish fauna were taken. Others of interest were from off E. Florida, on the ‘‘Pourtales Plateau.’’ The total number of species taken was about thirty.
In consequence of these more modern dredging expeditions the number of species of West Indian shallow-water starfishes, living in less than 150 fathoms, has been very much increased, while the deep-water species have become even more numerous.
I have had opportunity to examine most of the types of the ‘‘Blake Expedition’’ starfishes, described by Perrier, and also many of those taken in the West Indies by the Albatross. But I have not had opportunities to make any thorough studies of the latter as a whole, and, therefore, can not give any useful state- ment as to the number of additions to the fauna contained in them. Some of the species from the Albatross explorations were revised, and a number of new species were described and figured by me in 1899. (Revision of Genera and Species of Starfishes, see Bibliography. ) ;
At present about 125 species are known to me from that gen- eral faunal region, or from Florida to Brazil. Of these about 42
4Lists of all the dredging stations occupied by the Albatross, Blake, and other U. S. vessels up to 1887, with physical data, were published by
Sanderson Smith, in the Annual Report of the U. S. Fish Commission, for 1886, pp. 873-1014.
WEST INDIAN STARFISHES i
species have not been taken in less than 150 fathoms, and several are still undescribed.
Some of the West Indian shallow-water species have a wide range. A few range from Florida to the Abrolhos reefs, Brazil, or to Bahia. Two or three are said to be found on the West Coast of Africa, or at the Cape Verde Islands. Four reach Ber- muda. None are identified with those of Panama, though some are closely related. Among the deep-water species a few have been taken also off the eastern coast of the United States, north of Cape Hatteras, and three or four are thought to be identical with East Atlantic species from similar depths. Yet most of the species are peculiar to this fauna.
Very little is known concerning the starfishes to be found on the extended coasts of the Gulf of Mexico, beyond the coasts of western Florida and Alabama. Mr. Ives has listed several species from Vera Cruz and Yucatan. Two are peculiar to Vera Cruz, so far as known. Yet there are more than two thousand miles of that coast line about the marine fauna of which we know almost nothing.
That coast is almost everywhere sandy, and therefore not fa- vorable to many shallow-water invertebrates, except in sheltered localities.
It is known, however, that several of the Florida species reach Vera Cruz and Yucatan; others reach Colon. I can now add that one species (Astropecten americanus), common at moderate depths north of Cape Hatteras, occurs at similar depths on the northern coast of Colombia.
The Brazilian species are here included, because several, or the majority, are identical with West Indian and Florida species. The Brazilian starfishes have been listed by Mr. Richard Rath- bun, 13 species (1879), and later by Ludwig, 1882. At least nine or ten of the Brazilian species occur also in the West Indies.
In this report all those species which are known to have been obtained in less than 150 fathoms are included, though a num- ber of them really belong to the deeper water series, but find their upward limits in 100 to 150 fathoms. Doubtless other deep- water species will hereafter be found at such depths, especially the young. A few other deep-water species have been included for the sake of comparisons with their shallow-water allies.
8 NATURAL HISTORY BULLETIN
It is proposed to include, at the end, a special list of the other deep-water species (about 40) not reported upon, for the sake of completeness.
Thus the systematic part of this report will include about 82 species and subspecies, nearly all of which are redescribed in de- tail, with their known distribution, ete.
Comparatively few, even of the shallow-water species, have ever been figured. Therefore some of them, though not new, have been figured in this report.
The drawings of structural details are by A. Hyatt Verrill, and also most of the photographs.
Class ASTERIOIDEA MORPHOLOGICAL FEATURES
The Asterioidea® have a polygonal or star-shaped body, in which the rays are direct prolongations of the body itself, and contain extensions of the body-cavity and more or less of the viscera, especially one or more pairs of gonads and a pair of di- gestive glands; generally, also, a lobe of the saccular stomach.
The actinal or ventral side of the disk and rays has deep, radial, ambulacral grooves, extending to the tips of the rays. The roof of the groove is supported by two rows of ambulacral os- sicles, arranged like rafters, or in close, inverted, V-shaped pairs of plates or bars, between which are rows of pores for the pas- sage of the tubular ambulacral feet or ‘‘podia,’’ which are usual- ly terminated by a muscular sucker, but are pointed in the sub- order Paxillosa.
In the middle line of each ray and external to the ambulacral plates are situated the radial nerve and blood-vessel; and deeper within, the median radial ‘‘water-tube’’ or ambulacral tube. The radial water-tube supplies water to the podia through the medium of muscular ampulle situated internally, above the ambulacral plates. Usually there are two of these to each podium, but in Echinaster, Ctenodiscus, and some other genera there is only one.
The grooves are bordered on each side by a row of plates called adambulacral, which always bear spines.
5This spelling of the name is preferred because it is derived from As- terias (not from Aster.)
WEST INDIAN STARFISHES 9
The mouth is central, dilatable, and surrounded by soft mem- brane. The so-called jaws are merely the adoral ambulacral and adambulacral plates, more or less modified and coalesced. The ‘‘teeth’’ are only slightly modified adambulacral spines.
The stomach is very saccular and usually evertible. It usually has a lobe and a pair of digestive glands extending into the cavity of each ray, but in the case of some multiple-rayed species (Heliaster), it has lobes corresponding only to the primary five rays. In this case the five stomach-lobes do not enter the rays, but the five pairs of digestive glands do. In a few slender-rayed genera, also, the stomach is confined to the disk. The intestine is rudimentary or abortive, and not functional in most cases; usually there is a median dorsal nephridial or ‘‘anal’’ pore, main- ly for the discharge of the secretions from a lobulated gland.
The respiration is partly dermal, but especially by tubular dermal outgrowths, called papule. The madreporic plate is dor- sal, usually single, but sometimes two or more are present in autotomous species. The rays terminate in an enlarged apical or ocular plate, carrying an ocellus.
The skeleton is made up of large numbers of ossicles and plates of various kinds, mostly articulated so as to be more or less mov- able, giving flexibility, both to the rays and to the disk, though in some species (e. g. certain Goniasteride) the flexibility is slight, except at the tips of the rays.
The external skeletal plates are classed as dorsal or abactinal; marginal; interactinal; adambulacral; ambulacral; oral; and ocular or apical. The dorsal plates are very diverse in form and arrangement. They may be irregularly reticulate, tesselated, or imbricated. They may form regular radial rows; sometimes they are abortive, or nearly so. They are concealed by a thick dermis in certain genera.
The dorsal plates, like the marginals and interactinals, com- monly bear spines or small spinules, but they may be covered with granules, or with a smooth soft integument, or even appear quite naked, being then covered only with a thin membrane.
These plates and their armatures of spinules take several special names, according to their forms and structure, and are often characteristic of special genera and families and higher groups.
10 NATURAL HISTORY BULLETIN
When they become columnar or of hour-glass shape, and have the summit covered with a radiating cluster of small slender spinules, they are called paxille. These are the most typically formed in Astropecten, India, and allied genera, and are char- acteristic of the suborder Paxillosa.®
In nearly all the families there are two rows of specialized plates on each side of each ray, known as superomarginal and inferomarginal plates. These are often the largest and most con- spicuous of all the plates, especially in the Phanerozona, such as Astropecten, Gonaster, ete. In other cases both rows are much reduced in size and are not easily distinguished, as in some of the Asteriide. Or only the upper row may be reduced and in- distinct, as in Lwidia. Rarely one or both rows are abortive in certain deep-sea genera. Secondary intermarginal rows of plates are often interpolated between the marginal rows in some fam- ilies.
Between the inferomarginals and the adambulacral plates there may be one or more rows of plates called interactinals. The first and most important of these is next the inferomarginal row, and is called the peractinal row in this article. It may extend nearly or quite to the tip of the Ray. The other rows are called subactinal rows, and may be much shorter and later formed. All these interactinal rows may be lacking in many genera, or they may be rudimentary and spineless.
They appear later in development than the marginal plates, and the new ones are not formed next the ocular plates, as are the marginals.
Pedicellariz of one or more kinds are generally present, but are rare or entirely lacking in certain families, such as the Solas- teride, Echinasteride, Pterasteride, and in some genera of other families. Their uses are imperfectly known. One use it to keep the body free from dirt and parasites.
These curious organs are not found in other classes of Echino- derms, except in the Echinoidea. In the latter they are com- monly mounted on long stalks and generally have three valves — sometimes four or more or only two.
In the Asterioidea they are usually sessile, but in the order
6 For various other special forms and their names, see below, under Phanerozona and Valvulata.
WEST INDIAN STARFISHES ata
Forcipulosa they have very short pedicels, and they are most fre- quently bivalved.
They are of several different types, which are commonly char- acteristic of the orders and families, and often of the genera.
Sometimes there may be bivalved, trivalved, four-valved, and five-valved ones on a single specimen, as in Dermasterias im- bricata. In some eases they are not really valvular, but consist of several movable spinules, arranged in convergent groups or opposed rows. ‘The sessile valvular forms are generally situated over a pore, penetrating a plate, for the passage of nerves, etc.
Some of the special forms are described below, under the sev- eral orders. ;
In nearly all species there are numerous small, soft, tubular outgrowths from the body-wall, called papule, serving for respir- atory purposes. They may be in large groups, especially be- tween the dorsal and lateral ossicles, or stand singly, or they may occur only in special areas. Rarely they are branched, as in Tuadia.
Most shallow-water starfishes that have been studied in this respect have a free-swimming, bilateral larval form, known as a brachiolaria. A considerable number carry their eggs and young till they develop the starfish form. These have an abbreviated metamorphosis. The two methods may occur in different genera of the same family (e. g., Asterude).
The embryology of the deep-sea species is unknown in most cases, and may be diverse, or even more abbreviated.
CLASSIFICATION OF ASTERIOIDEA.
In this report I have adopted the division of the class into three large orders, viz.: J. ForcrpuLosa or FoRcIPULATA. II. SprnuLosa, with suborders, AVELATA and VELATA. III. PHANEROZONA. The last has three suborders, considered orders by some writers, Viz. : I. VALVULOSA or VALVATA. Il. Noromyora Ludwig (as an order). JIT. Paxm.osa.
12 NATURAL HISTORY BULLETIN
Order ForciepuLosa Verrill or Forcipuuata Perrier.
Stelleride forcipulate Perrier, Mem. Etoiles de Mer, pp. 166, 188, 1876.
Forcipulata Perrier, Exped. Sci. Trav. et Talisman, p. 27, 1894.
Asteries Ambulacraires ViIGUIER, Squellette des Stellerides, Arch. Zool. Exper., Vil, p. 93, 1878.
Cryptozonia (pars) Sladen, Voy. Chall., xxx, pp. xxxiv, 397, 1889.
Adetopneusia (pars) Sladen and Leptostroteria (pars), op. cit., p. XXXiv.
Forcipulosa Verrill, op. cit., p. 24, 1914a.
In this order the form is always stellate, often with long rays, commonly five, but often multiple. Ambulacral plates, except the orals, are usually short and closely crowded (leptostroterial), but not in Brisingide. The proximal pair is elongated and, ex- cept in Pedicellaster and a few others, forms the inner end of the jaw. Ambulacral feet are generally arranged in four rows; but in two rows in Pedicellasteride, Brisingide, and Zoroaster- ide. In certain large species of Pisaster there may be more than four rows.
Aside from the ambulacral and adambulacral plates, the skel- eton of the rays consists of five fundamental or primary rows of ossicles, viz., the median dorsal or carinal; the two superomar- ginals; and the two inferomarginals. The latter may or may not be confined to the ventral side.
The ossicles of adjacent rows may be articulated directly by their lobes, or small, simple connecting ossicles may intervene.
This simple or primitive type of skeleton is found in Cor- onaster briareus (see pl. 1x, fig. 4) and some other genera, espe- elally when young.
In most cases more or less numerous ossicles or rows of ossicles are interpolated during growth between the five primary rows, either above or below, or between the marginals, thus giving rise to many different styles of skeleton in the more complex genera.
The carinal or median dorsal plates usually remain distinct, but are often small and not easily recognizable. The two mar- ginal rows are nearly always easily recognizable, though small, and they generally bear spines larger than the dorsals. They never form a stout margin, as they do in the Paxillosa.
Interactinal plates are various. Usually there is at least one row, the peractinals, but these are often small and spineless and sometimes are lacking. In other genera there may be several rows of spiniferous interactinals.
WEST INDIAN STARFISHES 13
The Forcipulosa, as the name implies, are specially character- ized by the possession of forcipate pedicellariz, not found in other orders. Two forms of pedicellariz are nearly always pres- ent, except in the young.
The minor or forcipulate pedicellariz, peculiar to this order, generally form wreaths or large or small clusters on the spines, but they may also occur, either singly or in clusters, on the in- tegument of the dorsal or lateral plates, on the papular areas, and in other situations. Their two blades are prolonged proximally and crossed, with muscles attached to the ends, so that they can open and shut like forceps or scissors. Their valves are concave and are usually armed with denticles.
The major or forficulate pedicellarie are generally ovate or lanceolate, sometimes wedge-shaped, stone-hammer-shaped, lyrate, or spatulate. The tips may be acute, plain, flat or denticulate.
They are mostly dermal, but may also occur on the ventral and oral spines, or along the inner edges of the grooves, or attached to pedicels in clusters within the grooves, or even on the pedicels of the ambulacral feet. They may be of several sizes and forms on one specimen.
Certain species of Asteriidew, Coronaster, ete., have a peculiar very large form of dermal pedicellarie, often as thick as the spines, in which the broad valves terminate in a series of curved denticles, so that when closed the valves resemble a pair of clasped hands, or better still, a pair of clasped feet of a cat, with the claws protruded. These may be called felipedal or wngwecu- late. (See pl. ix, fig. 4c.)
FAMILIES AND SUB-FAMILIES OF FORCIPULOSA.
This order now includes the following families and sub-families : J. Family AsTermp2.
Subfamilies ASTERINZ ; STICHASTERINZ ; PYCNOPODINZ ;
HELIASTERINZ. II. Family ZoROASTERID. If. Family PEpIcELLASTERID2. IV. Family Brisincip2. Subfamilies BrisincINzZ; LABIDIASTERINE (Type, Lab- idiaster Lutk.).
14 NATURAL HISTORY BULLETIN
The first three families are represented in depths less than 150 fathoms, though all except the first family are essentially deep- sea groups.
Family ASTERIDZ Gray (emended).
Asteriide Gray, Aun. & Mag. Nat. Hist., vi, p. 178, 1840; Synopsis, p. i, 1866. Perrier, Revis. Stell., Arch. Zodl. Exper. et Gen., iv, p. 302, 1875; Mem. Etoiles de Mer, pp. 167, 198, 1876. Viguier, Squellette des Stellerides, pp. 93, 99, pl. v, figs. 1-10, 11-12, 1878.
Asteriide (emended) + Stichasteride Sladen, Voyage Chall., xxx, pp. 430, 560, 1889. Perrier, Exp. Trav. et Talisman, pp. 43, 105, 107, 128, 1894; Contrib. 1’étude des Stellerides Atlan. Nords, pp. 25, 31, 1896. Verrill, Monog. Starfishes, North Pacific, pp. 27-44, 1914a (emended.)
Stellate starfishes, most often with five or six elongated rays; sometimes with a variable number of unequal rays in autotomous species; sometimes with numerous rays, increasing with age by the outgrowth of interpolated rays, as in Heliaster and in Pycno- podia. Madreporic plate generally single in five-rayed and six- rayed forms, but often two or more in those with a variable num- ber of rays, due to autotomy or budding. Dorsal and actinal plates various in form and arrangement, most often reticulated. Odontophore usually formed of a single piece. Oral ambulacral plates elongated and forming the jaws. Those along the grooves become very short and compressed. Papule occur both on the actinal and dorsal sides. Ambulacral feet usually crowded in four rows, sometimes more. Pedicellariz of two kinds are pres- ent. Dorsal plates generally bear spines or spinules, various in size and kind, and sometimes granules. They are never true paxille. 3
The minor pedicellariz may be attached directly to the in- tegument, either singly or in clusters, or to the spines, to the pedicels of the larger forms, to saccular dermal growths around the spines, to the inner edge of the ambulacral grooves, or rarely even to the ambulacral feet. They are most commonly attached by slender and sometimes long pedicels, and aggregated into wreaths or clusters attached to a contractile fold of dermis on the spines. They are often so abundant, in the wreaths around the dorsal spines, that they nearly or quite conceal the spines and integument in living specimens. In some cases they are at- tached in large numbers to large muscular dermal sheaths or
WEST INDIAN STARFISHES 15
sacks loosely surrounding the spines nearly or quite to the tips when extended in life. They often have a formidable array of minute sharp denticles, and are frequently very characteristic of species.
Subfamily ASTERIIN. Asteriide (restr.) Sladen, op. cit., 480, 560, 1889. Perrier, 1894, pp. 105, 128; 1896, pp. 25-31. Astertine Verrill, op. cit., p. 42, 1914a.
The Asterune are chiefly characterized by the more or less openly reticulated arrangement of the dorsal and lateral ossicles, especially of the dorsal ossicles, which are usually more stellate or lobulate and more slender than in the Stichaster group, and generally united either by the ends of the lobes or apophyses, or else by means of interpolated ossicles, thus allowing more flexi- bility in the rays, and larger papular areas. The dorsal plates may be in definite rows or irregularly reticulated.
The Stichasterinew, on the contrary, are chiefly characterized by the more or less regular subtesselated or imbricated arrange- ment of the ossicles, both on the sides and dorsal surface of the rays, the ossicles themselves being larger, broader, and more rhombic, and usually united in more definite, longitudinal and transverse rows, with smaller and more regular intervening pap- ular areas. In either group, as at present understood, the inter- actinal plates may have the regular serial and subtesselated ar- rangement.
The discovery of many new generic and specific types inter- mediate between typical Stichaster* and Asterias, renders it difficult to define the limits of the two so-called families, typified by these genera. ;
The only West Indian genus that has been referred to the Stichasterinz is Stephanasterias. To me this seems to be more nearly related to Leptasterias and therefore one of the Asteriine.
Genus ORTHASTERIAS Verrill. Orthasterias Verrill, Starfishes North Pacific, pp. 168-185, 1914a. Long-rayed diplacanthid starfishes with a small disk. Dorsal
7 The type of Stichaster is S. striatus M. and Tr. = S. aurantiacus of later writers. It is a Chilian species.
16 NATURAL HISTORY BULLETIN
skeleton consists of lobed, wide, subimbricated or contingent plates, arranged in three, five, or rarely more regular radial rows, and sometimes with extra interpolated ossicles; the rows are con- nected by transverse ossicles. The principal plates bear, on a central boss, large, isolated spines, usually making three or five longitudinal rows above the marginal rows.
Superomarginals are large, usually monacanthid, four-lobed ; the descending lobe is large and usually joined directly to the inferomarginals.
The inferomarginals are strong, convex ossicles, each bearing usually two large spines.
Uusually there is one row of interactinal plates, with or with- out spines, closely united to the adambulacrals and inferomar- ginals; sometimes they are rudimentary and without spines. (Stylastervas. )
Papule are numerous and clustered.
The lateral and dorsal major pedicellariz are large, and are usually of two or more forms. The larger are erect, wedge- shaped, stone-hammer-shaped, or ovate, with the tips of the valves wide and usually denticulate. Others are more slender, with the valves spatulate or narrowed in the middle and the tips unguicu- late, with interlocking teeth. Others similar, or smaller, with acute tips, often occur on the adambulacral spines and margin. Type, O. columbiana V.
This genus includes several large species on the North Pacific coast. Off the eastern coast of the United States one species (O. tanner Ver.) is abundant at moderate depths, from south of Cape Hatteras to southern New England.
The subgenus Stylasterias Ver. (op. cit., p. 48, 1914a), was sep- arated mainly because it has only rudimentary and spineless interradial plates, which are not visible without preparation.
The following West Indian species belong to this group or sub- genus. The young of O. tanneri, when two or three inches across, have the same features.
ORTHASTERIAS SUBANGULOSA Verrill.
Asterias angulosa Perrier, Bull. Mus. Comp. Zodlogy, vol. ix, p. 3, 1881; Mem. Etoiles de Mer, p. 202, 1884 (non Miiller.)
Orthasterias (Stylasterias) subangulosa Verrill, Starfishes N. Pacific Coast, pp. 168, 370, 1914a.
WEST INDIAN STARFISHES 17
Plate ii; figures 1, 2. Plate ix; figures 1, la.
The largest specimen has the radii 9™™ and 66™; ratio, 1:7.33. This was from the Bahama Expedition, station 43, in 15 fathoms. The disk is small and has a very evident central pentagon of plates and spines. The rays are slender, somewhat angular, reg- ularly tapered to subacute tips.
The dorsal plates form a conspicuous raised median carina, and a smaller dorso-lateral row on each side, well separated from the prominent superomarginal row by a wide lane, containing large papular areas and crossed by stout transverse ossicles uniting the plates of the two rows, but without spines.
The spines generally stand singly on all the dorsal, radial, and superomarginal plates, and all have the same form. They are rather long, terete, regularly tapered, and acute. They are sur- rounded, near the base, with a dense wreath of minor pedicel- lariz of small size.
The inferomarginal plates form a conspicuous row well sep- arated from the upper ones by a wide lane, which contains a row of large papular areas and bears many rather large, erect, lanceo- late or subacute major pedicellariz.
Each plate usually bears two spines, one nearly above the other. The upper oneisthe larger. It is larger than the supero- marginals and dorsals, and is somewhat flattened, often a little expanded distally or sublanceolate, and subacute; the other is distinctly smaller and more acute. Sometimes there is prox- imally a much smaller third spine below the second, apparently arising from the same plate, but perhaps, in some cases, arising from a rudimentary interactinal ossicle not visible at the surface. The third spine is lacking in specimens a little smaller.
The upper inferomarginal spines bear, at about midheight, a dense cluster of numerous minor pedicellarize on the upper side. There are also some of the rather large lanceolate major pedicel-
lariz between their bases. Similar major pedicellarie occur on and between the adambulacral spines, on the inner margins of the ambulacral furrows, and on the interradial areas.
The adambulacral plates bear each two rather long, slender, flattened, subequal spines, which form two regular rows; the and more flattened or subspatulate. The peroral spines are- outer ones are the larger. The adoral spines are longer, larger,
18 NATURAL HISTORY BULLETIN
rather stout and blunt. It appears to be the undetermined star- fish described by Nutting (Narrative Bahama Exped., bottom of page 168). This species was dredged by the Bahama Expedition at station 43, off Sand Key, Florida, in 15 fathoms; at station 47, off Key West, in 80 fathoms; and at station 61, off Key West, in 75 to 80 fathoms. Several specimens were obtained. Perrier recorded it from the Blake stations, off Florida, in 85 and 110 fathoms, one from each place. The type had the radii 6™ and 43mm, ORTHASTERIAS CONTORTA (Per.) Verrill.
Asterias contorta Perrier, Bull. M. C. Z., vol. ix, p. 1, 1881; Mem. Htoiles de Mer, p. 199, 1884. O. contorta Verrill, op. cit., p. 48, 1914a.
This species was not figured by Perrier and only imperfectly described. The size was not given. I did not examine his cotypes, in the Museum of Comp. Zoology, except in the case of a single lot of small specimens (sta. 155, 88 fathoms). These appeared to me to be the young of O. subangulosa.
According to Perrier’s description it has no dorsal rows of plates except the carinals, which might be due to immaturity. He also mentions that the submarginal papule are large and placed singly. In O. subangulosa they are rather small and stand two or three together.
Without farther study of the types it is not possible to say whether they are distinct species.
Perrier recorded this from ten Blake stations, in 11 to 188 fathoms, from Florida reefs to Cariacou and Barbadoes. |
Genus COoSCcINASTERIAS Verrill.
Cosinasterias Verrill, Trans. Conn. Acad. Sci., i, p. 248, 1867. (Type, C. muricata V. = (?) C. calamaria (Lam.).)
Stolasterias (pars) Sladen, Voyage Chall., vol. xxx, pp. 563, 583, 1889. (Type, C. tenuispina.) Non Perrier.
Coscinasterias and Polyasterias Perrier, Exped. Trav. et Talisman, p. 108, 1894.
Coscinasterias Verrill, op. cit., pp. 45, 46, 1814a.
Disk small, rays elongated, somewhat angular, usually carinate. Dorsal plates of the rays stout, four-lobed, usually arranged in three or five regular rows, besides the upper marginals, which form regular lateral rows. Interactinal plates form one primary
WEST INDIAN STARFISHES 19
(peractinal) row, like the lower marginals, but they may be rudi- mentary and spineless in half-grown specimens, and only bear spines in large, mature individuals, as in C. tenwispina.
Minor pedicellariz are abundant. Large forficulate or major pedicellariz of the usual forms are present, often in considerable numbers, on the dorsal and lateral plates, and smaller ones occur on the marginal areas. Large unguiculate pedicellariz are lack- ing. Adambulacral plates are monacanthid or nearly so.
Stolasterias, which was proposed as a subgenus by Sladen in 1889, was nearly identical with Coscinasterias, characterized by me in 1867. He gave no reason for changing the name. It should be regarded as a synonym of the latter and be eliminated. Sladen’s type was A. tenwispina, which is a Coscinastervas with the interactinal plates rudimentary and without spines, except in the larger specimens.
As C. tenutspina often divides spontaneously, it belongs to Polyasterias in Perrier’s arrangement. This would make the lat- ter strictly synonymous with Sladen’s typical Stolasterias.
M. Perrier (1894) correctly retained Coscinasterias for the typical forms, like calamaria, but separated those species that are known to undergo spontaneous fission under the name of Polyastervas = typical Stolasterias Sla., though they do not ap- pear to differ much in structure from the preceding. He re- tained Stolasterias Sla., in a restricted sense, for those allied to glacualis, although C. tenwispina was named by Sladen as the type.
A. glacialis had, long before, been made the type of Marthas- terias by Julien.
COSCINASTERIAS TENUISPINA (Lam.) Ver.
Asterias tenuispina Lamarck, op. cit., 1816, vol. ii, p. 561. Perrier, Re- vision, op. cit., p. 306, 1875.
Asteracanthion tenwispinus Mull. and Trosch., Syst. Asteriden, p. 16, pl. i, fig. 1, a. b., 1842. Perrier, Pédicellaires, p. 32, pl. i, figs. 3-3c, 1859. Lutken, op. cit., 1859, p. 95.
Asterias (Stolasterias) tenwispina Sladen, pp. 565, 583, 1889. Verrill, The Bermuda Islands, vol. ii, part v, p. 324, in Trans. Conn. Acad. Sci., vol. xii, p. 280, plates xxxiv, fig. 2; xxxiv c, fig. 2, 1907.
Coscinasterias tenuispina Verrill, op. cit., 1814a, p. 45.
Asterias atlantica (pars) Verrill, Transactions Conn. Acad. Sci., vol. i, p. 368, 1868. R. Rathbun, Brazilian Echinoderms, in Trans. Conn. Acad., vol. v, p. 144, 1879.
20 NATURAL HISTORY BULLETIN
Plate xxvi; figure 2. Plate xxvii; figure 4.
Disk small. Rays long, rather slender, angular, variable in number. Dorsal and marginal spines are elongated and rather slender. Adambulacral spines slender, one to a plate. Color in life usually pale purplish or violaceous above, yellowish below.
This species is autotomous and therefore is commonly found with the rays unequal and variable in number, from two or three to nine or more, but most frequently with six to eight, three or four being often much shorter than the others. There are usual- ly two or more madreporic plates. Sometimes it is regularly six- rayed or five-rayed. It becomes six to eight inches in diameter at the Bermudas.
The dorsal plates are rather openly united and form a definite median or carinal row and one main dorsolateral row on each side, often with one imperfect second row proximally in large specimens. Each dorsal plate usually has a rather strong, some- what elongated spine.
The superomarginal row of plates is well marked and forms the lateral margin of the ray. Each plate bears one, or some- times two, elongated spines, longer than the dorsals. The supero- marginals are also strongly developed and have a row of large spines close to the adambulacrals.
The interactinal plates are small and usually without spines, but in very large specimens some of them may bear a small spine.
The adambulaeral plates normally bear one long slender spine, but in very large specimens a few bear two spines. In life the color varies. It is usually some light shade of violet or pale purple; yellow below.
This is a common Mediterranean species. It has been reported from Madeira, the Canary Islands, Cape Verde Islands, Brazil, Abrolhos Reefs, Brazil (coll. C. F. Hartt, Yale Mus.), Cuba, the Bermudas, and West Indies. It is the only common starfish at the Bermudas.
It is well adapted, by its habits, to be carried from one country to another, adherent to the bottoms of vessels.
CoSCINASTERIAS TENUISPINA, var. ATLANTICA Verrill.
Asterias atlantica (pars) Verrill, Corals and Echinod. of Brazil, Trans. Conn, Acad. Science, vol. i, p. 368, 1868.
WEST INDIAN STARFISHES 21
One of the types of this supposed species was a regularly five- rayed specimen from Cuba. It differs sufficiently from the ord- inary form of the species to receive a varietal name, and there- fore I propose to designate it as above.
The rays of the type of this ferm are decidedly angular, rel- atively shorter and stouter than in the ordinary form. The median radial and two marginal rows of spines and plates are strongly developed, and one dorso-lateral row is less so. The adambulacral spines are monacanthid, as in the typical form. The dorsal and marginal spines are shorter and stouter than in the latter.
The type was from Cuba. I am not prepared to say that the other American forms are fully identical with those of Europe, for I have never been able to study a good series of both in any museum. Valid specific characters may have been overlooked by authors who unite them.
CosCINASTERIAS LINEARIS (Perrier) Verrill.
Asterias linearis Perrier, op. cit., p. 2, 1881; Etoiles de Mer, p. 201, pl. iii, fig. 5, 1884.
The type of this species is small and evidently young. It is monacanthid and evidently a Coscinasterias. It may, perhaps, prove to be the young of C. tenwispina when a good series can be studied.
The type had the radii 5™” and 50™"; ratio, 1:10. The rays are five, slender, angular. The dorsal side of the rays has three rows of plates, each of which bears a single elongated spine, thus forming three longitudinal rows of spines. There are three rows of ventral spines. The outer one forms the lateral, larger (in- feromarginal) row of spines. Between these and the upper lat- eral row are large isolated papule; on the dorsal side the papule are smaller and clustered. There appear to be two rows of in- ‘ teractinal spines proximally. Major pedicellariz, elongated in form, were found on the disk.
Adambulacral spines are one to a plate. They are straight and not tapered. (Condensed from Perrier’s description.) The type was from Blake station 45, in 101 fathoms, off West Florida.
22 NATURAL HISTORY BULLETIN
Genus LEPTASTERIAS Verrill.
Leptasterias Verrill, Proc. Boston Soc. Nat. Hist., x, p. 350, 1866. Type, A. miilleri Sars. Sladen, Voy. Chall., xxx, p. 563, 1889. Perrier, Exp. Trav. et Talism., p. 108, 1894. Verrill, op. cit., p. 116, 1814a.
An extensive group of small, more or less diplacanthid star- fishes, closely related to typical Asterias, usually with a single row of peractinal plates and spines. Rays five or six. The more typical forms have slender rays and a small disk, with the dorsal ossicles irregularly arranged, or showing several imbricated ra- dial rows, with numerous spines. It differs from typical Asterias in the diminished number and larger size of the dorsal and lat- eral papule, and the small number of actinal plates, of which there is generally only a single row, but more profoundly in hav- ing the genital pores on the actinal side, near the mouth, and in the ovaries, which are different in form and produce larger eggs, which have an abbreviated development, passing through no free-swimming larval stages.
The eggs and young are carried in clusters, adhering in front of or around the mouth in all the species studied in the breeding season.
LEPTASTERIAS FASCICULARIS (Per.) Verrill.
Asterias fascicularis Perrier, op. cit., 1881, p. 1; Mem. Etoiles de Mer, p- 200, pl. ii, fig. 3, 1884.
The type had the radii 9™™ and 57™™; ratio, 1—6.3. The five slender rays are slightly contracted proximally. The dorsal and dorso-lateral surfaces of the rays have about seven alternating rows of plates, separated by papular areas smaller than the plates, and containing usually two papule, and holding about two or three dermal minor pedicellariz.
The plates of the median row have a curved row of three or four short, blunt spines; those of the other rows have one or two similar spines; some single spines are also situated on the inter- polated ossicles.
The disk is small and covered with spines like those of the rays; between them are some oval major pedicellariz with curved valves, touching only at tips.
On the under side of the rays there are two rows of plates, near the adambulacrals; each plate bears a single spine and often a dermal minor pedicellaria.
WEST INDIAN STARFISHES 23
The adambulacral spines are rather long and slender, two to a plate, divergent, forming two regular rows.
This species seems to be a true Leptasterias, allied to LD. compta of New England. Its mode of reproduction and position of genital pores are unknown, and therefore its generic position cannot be positively determined.
The type was from off Guadaloupe, in 309 fathoms.
LEPTASTERIAS HARTI R. Rathbun. Leptasterias hartiti R. Rathbun, Echinoderms of Brazil, Trans. Conn. Acad. Science, vol. v, p. 145, 1879.
This is a small, six-rayed species. Radii of the type were about 3.2" and 19™™; ratio, 1:6.
Rays are well rounded above, tapering regularly to slender tips. The row of median radial plates is distinct; each plate bears a V-shaped row of about five small spines, becoming re- duced to two or three distally. Spines on the disk are similar to those on the rays, closely and irregularly arranged.
Above the first lateral radial row there are, on each side, about two longitudinal series of small spines, with many interpolated on the connective ossicles. The rows are not very regular. The first lateral row, probably superomarginals, has two small spines to a plate, distinct to tip of rays; the series above this has, on each plate, transverse, irregular groups of four or five spines, de- creasing to a single one distally.
On the under side there are about three rows of plates and spines. The inferomarginal row has usually one spine, seldom two, to a plate.
The two marginal rows are well separated.
The spines of the interactinal plates form oblique series of two or three. They are larger than the adambulacrals, but not much longer, and have compressed clavate tips.
The adambulacral spines are regularly two to a plate, slender and clavate proximally; distal ones tapering; outer ones a little ’ larger.
Major pedicellariz are large and numerous, variable in size, generally scattered, above and below. They are triangular or ovate in form, with a strongly curved basal piece; about two- thirds as broad as long. The larger ones equal or exceed in length the lateral and dorsal spines, and are much stouter. They
24 NATURAL HISTORY BULLETIN
are most numerous between the lateral and ventral spines. The papule are mostly isolated.
Taken from a telegraph cable, about 30 miles east of Cape Negro, Brazil, in 62 fathoms.
The deseription above is condensed from that of Mr. Rathbun.
LEPTASTERIAS MEXICANA (Liitk.) Verrill. Asteracanthion mexicanum Liitken, Vidensk. Meddel., p. 94, 1859. Asterias mexicana Verrill, op. cit., p. 344, 1867.
This was described by Liitken from two small specimens from Vera Cruz. They were evidently young. The larger was 1.5 inches in diameter. I am not aware that anyone has obtained additional specimens.
According to Liitken the rays are five; the upper side has about seven rows of short, obtuse, rough spines surrounded by a dense basal wreath of minor pedicellarie. The papule stand two or three together.
The adambulacral plates are one and two to a plate, in pretty regular alternation. They bear pedicellariz at about midheight. They are slightly flattened, slender, and obtuse. The ventral side has two rows of small spines. (Condensed from Liitken.)
Litken compared it to L. miiller1, the type of Leptastervas. Although it appears to have the external characteristics of Lep- tasterias, it may be the young of a larger species of Asterias, like A. forbest.
There is no reason why A. forbes: should not be carried to Vera Cruz adherent to the bottoms of vessels from our southern ports, even if it does not habitually live on the Mexican coast. - Many other creatures are thus carried to distant places and no doubt, in many cases, become naturalized in new homes by this means. I have not seen A. forbes: from the Gulf of Mexico, but should expect to find it on the coast of West Florida and Ala- bama.
Genus STEPHANASTERIAS Verrill, 1871. Type, S. albula.
Stephanasterias Verrill, Bull. Essex Inst., iii, p. 5, 1871; Expl. of Casco Bay, Proc. Am. Assoc. Adv. Sci. for 1873, pp. 356, 359, 364, 1874; Check List Invert., 1879; Expl. Albatross, 1883, p. 540, 1885.
Nanaster Perrier, Exp. Trav. et Talism., pp. 129, 131, 133, 1894; Camp. Scientif. 1’Hirondelle, p. 27, 1896. (Type, S. alGula.)
Stichaster (pars) Verrill, 1866, p. 351. Perrier, p. 347, 1875. Sladen, p. 432, 1889.
WEST INDIAN STARFISHES 25
Stephanasterias Verrill, Revision Genera and Species of Starfishes, p. 222, 1899; North Pacific Starfishes, op. cit., p. 146, 1814a.
Small autotomous starfishes, commonly found with five to nine unequal rays, due to incomplete replacement and regulation of rays after division. Two or three longer rays, and three or four shorter ones are common forms. When fully mature, often reg- ularly six-rayed.
The whole surface is thickly covered with minute thorny spinules, placed in clusters on the plates, and longer on the under side and margin.
The median-radial and marginal rows of plates are usually distinet, with wide plates, closely imbricated radially. The sev- eral dorsolateral rows have the plates small, short, and relatively wide, irregularly arranged, but with longitudinal rows of papular pores between them.
The inferomarginal and interactinal plates form regular, close, longitudinal rows.
The adambulacral spines are slender, two to a plate, or alter- nately one and two. Papule are isolated or in small groups. Both kinds of pedicellariz are present.
Perrier changed the name of this genus to Nanaster, because he erroneously assumed that Stephanasterias had been preoccu- pied by Stephanaster Ayres. The two names are distinct and differently derived.
STEPHANASTERIAS GRACILIS (Per.) Verrill.
Asterias gracilis Perrier, op. cit., p. 4, 1881; Nouv. Arch. du Mus., vi, p. 204, 1884.
Stephanasterias gracilis Verrill, Trans. Conn. Acad. Sci., vol. x, p. 223, 1879; Starfishes N. Pacific, p. 146, 1914a.
Plate ix; figures 2-2c.
Disk small; rays slender, variable in number, commonly six. The species is autotomous, and the rays are often unequal.
The dorsal surface is closely covered with numerous nearly uniform minute, slender, subdiaphanous spinules, several on each plate. The marginal spines are larger, flattened and enlarged distally, mostiy with a blunt and thorny tip, and more or less hyaline,
The adambulacral spines are two to a plate, shorter and much
26 NATURAL HISTORY BULLETIN
more slender than the marginals, slightly clavate, subhyaline; oral and adoral spines similar.
This closely resembles S. albula, of the northern coast and arctic regions. It seems less variable in number of rays and is perhaps less often divided by autotomy. Its marginal spines are longer, more expanded and flatter.
I have examined several of the types of this species in the Museum of Comp. Zodlogy, from the Blake Expedition, described by Perrier.
It was taken by the Blake at nine stations in 56 to 183 fathoms, among the Lesser Antilles, and at two stations in the Gulf of Mexico, off West Florida, in 95 and 101 fathoms. The Albatross also took it at many stations.
It was obtained by the Bahama Expedition, off Havana, in 110 and 200 fathoms (stations 4 and 13), and off East Florida, in 110 fathoms (station 64, 6 specimens).
STEPHANASTERIAS HEBES Verrill, sp. nov. Plate ix; figure 3.
General appearance and size much as in S. albula, but stouter with less numerous and less delicate spines.
The adambulacral spines form two perfectly regular divergent rows. They are relatively short and not very slender, terete or slightly clavate and blunt. The ambulacral furrow is relatively very wide and the sucker feet large. Owing to the shortness of the spines and width of the groove, the adambulacral spines - cover only a small part of its margins.
The interactinal and inferomarginal spines are much larger and stouter than the adambulacrals and much more so that in S. albula. They are about twice longer and more than twice thicker than the adambulacrals adjacent. They are clavate, with rounded or obtuse tips, not so thorny as in S. albula, nor so trans- lucent. The dorsal spines are similar, but shorter, arranged in close clusters on the plates.
Taken by the Albatross at station 2766, in — fathoms. (No. 18,426, Nat. Mus.)
This is very much stouter than S. gracilis, with much larger and more clavate spines.
WEST INDIAN STARFISHES 20
Family ZoROASTERIDZ Sladen.
Zoroasteride Sladen, op. cit., pp. xxxvi, 416, 1889. Perrier, Exped. Trav. et Talisman, p. 112, 1894. Ludwig, op. cit., 1905, p. 159.
Disk small; rays long and slender, angular or rounded.
The disk is covered largely by the ten enlarged basal radials and interradials, and the large central plate.
A row of large overlapping carinal plates stiffens the rays, each usually bearing a spine. Marginal plates scarcely differentiated from the dorso-laterals and interactinals.
All these form close, regular, imbricated, longitudinal rows, often six to eight proximally. They are also imbricated in trans- verse rows.
All the plates are granulated or finely spinulose and the lateral and interactinal rows usually bear, also, a single elongated spine, and usually a dermal major pedicellaria. Papular pores are single and placed between all the rows of plates.
Adambulacral plates are alternately unlike. The larger are compressed, carinate, and project far into the groove. They bear a divergent row of several spines transverse to the groove, and have one or two small spines in the groove. The alternate small plates do not project into the groove, are not carinate, and bear fewer spines. Spines of both kinds of plates may bear large major pedicellariz.
Minor or crossed pedicellariz appear to be lacking. Sucker- feet are in four rows proximally, becoming reduced to two rows distally. Their suckers are small.
This is essentially a deep-sea family. Only a few species have occurred in less than 150 fathoms.
Genus ZOROASTER Thomson.
Zoroaster W. Thomson, Depths of the Sea, p. 154, 1873. Sladen, op. cit., p. 416, 1889. Perrier, op. cit., 1894, p. 115. Fisher, op. cit., 19110, p. The central plate of the disk has few smaller plates around it. The basal radials are prominent, but not much larger than the next radials and of the same character. The basal interradials are smaller and flatter, without a central spine. The central plate and basal and median radials have a central short articulated spine. The whole surface is covered with a rather thick dermis
and with small articulated spinules, attached to basal papille.
28 NATURAL HISTORY BULLETIN
In addition to these the lateral and interactinal plates have one slender, articulated spine.
The dorso-lateral row of plates next the carinal row is differ- entiated, the plates being larger than the others and similar to the carinals. The rest of the lateral and interactinal plates are all similar, forming six to eight regular rows, imbricated both longitudinally and transversely, leaving only small papular pores between them. Marginals are not much different from the rest.
Adambulacral plates of two sorts are very unlike. The larger carry four to six divergent slender spines, along the thin carinate edge; the larger one at the margin of the groove may bear one or several major pedicellariw, often large; other spines may bear smaller ones. The much smaller alternate plates are not carinate and bear usually two slender outer spines and a small one in the furrow. Major pedicellarie are scattered on the back and sides.
ZOROASTER ACKLEYI Perrier.
Zoroaster ackleyi Perrier, Comptes rendus, 1880, p. 436; op. cit., p. 6, 1881; Etoiles de Mer, p. 197, pl. iii, fig. 1, 1884; Exped. Trav. et Talisman, p- 117, 1894. A. Agassiz, Three Cruises of the Blake, vol. ii, p. 105, fig. 382.
The largest specimen examined by me has the radii 10™™ and 105™; ratio, 1:10.5; diameter of rays at base 10™™; height dD ieee
The small disk is convex or a little swollen. The long slender rays are well rounded at the base, but become somewhat carinate on the attenuated distal half. .
The disk is occupied largely by the ten basal radial and inter- radial plates, and a large prominent central plate, with a central mamilla supporting a short obtuse spine.
The dorsal radials are larger and more elevated than the inter- radials and have a central boss, which bears a short obtuse spine. They are rounded with small notches or emarginations around the margin, where there are isolated papular pores. Similar pores surround the central plate.
The basal interradials are flatter and more angular and have no central spine. They are in contact with the radials laterally. One of them is partly covered by the madreporie plate, which is rather large and convex, with radiating gyri.
WEST INDIAN STARFISHES 29
The central plate of the disk is surrounded by several small plates, and by papular pores.
All the plates of the disk and dorsal side of the rays are thickly covered with minute rough spinules that leave little round scars when rubbed off.
Pedicellarie of the forficulate kind, much larger than the spinules, are scattered over the whole surface, usually but one to a plate. When closed they are ovate-lanceolate, and some- what compressed, subacute.
The median radial row of plates is conspicuous. They are rather large, convex, with a small central boss, carrying a small short, thick spine, which is somewhat enlarged or obtuse at tip; the distal ones are smaller and acorn-shaped. They are easily detached and are often rubbed off. They are often absent from the distal part of the rays. These radial plates are shield- shaped, with rounded sides and overlap each other serially.
The sides and lower surfaces of the rays are proximally made up of seven to nine regular rows of small shield-shaped plates, all much alike, except the first row, so that it is not easy to dis- tinguish the marginal rows. The row next the median radials has larger, angular plates, similar to the median, though smaller. The others are imbricated transversely, in regular vertical rows, decreasing in size downward, and overlapping with their angular lower end, like slates on a roof.
In five or six lower rows each plate bears on a boss, a long, slender, very acute spine, decreasing upward in length, where they nearly intergrade with the spinules of the upper plates. These spines are also accompanied by small, acute spinules, and each plate usually has a rather large lanceolate or acute-ovate pedicellaria. Distally a similar spine appears also on the row next to the carinal row.
The adambulacral plates are alternately large and small. The larger ones are carinated and project far into the groove and ' bear, on the curved edge, five or six slender acute spines, of which two or three are within the groove, one far within, and the others form a divergent row, transverse to the furrow, the middle or marginal ones longest. Many of these spines, both within and outside the furrow, bear a single large, forficulate or major pedicellaria, some of them much thicker and about as long
30 NATURAL HISTORY BULLETIN
as the spine, and much larger than those of the dorsal surface. Some spines have also two or more smaller ones. The larger ones, on the marginal spine, are compressed, long ovate, with acuminate or contracted tips; the smaller ones are acute-ovate, lke the dorsal ones. The largest occur on the outer dorsal spines.
The peroral and adoral spines are considerably elongated and acute. The oral area is deeply concave, so as to conceal most of the spines. The smaller alternate adambulacral plates are not carinate and do not project into the grooves. They usually bear one very small inner spine and two longer very unequal outer ones.
The ambulacral sucker-feet are large. Proximally they form four rows; farther out they are in two zigzag rows on each side; toward the end of the rays, they are reduced to two nearly straight rows.
This was taken by the Blake, at station 134, off Santa Cruz, in 248 fathoms, 10 specimens; and at station 157, off Montserrat, in 120 fathoms, one specimen.
The specimens studied by me were mostly taken by the Al- batross. The larger, described above, was from station 2396, off Pensacola, Fla., in 335 fathoms, gray mud, bottom temperature 41.80° F. (No. 18,450, Nat. Mus.) The other was from station 2394, also off Pensacola, in 525 fathoms, light gray mud (No. 10,422). It was also taken at several other localities.
Family PEDICELLASTERID Perrier.
Pedicellasteride Perrier, Etoiles de Mer., pp. 167, 194, 1884; Exped. Trav. et Talism., p. 92, 1894. Sladen, op. cit., p. 556, 1889. Verrill, op. cit., p. 202, 1914a,
Pedicellasteride (pars) Perrier, op. cit., 1885, p. 15.
Disk small, rays usually five or six, sometimes ten to twelve or more, as in Coronaster. Dorsal skeleton usually reticulated, or with few rows of plates.
Ambulacral feet are in two rows, at least distally. The jaws are formed by modified adambulacral plates. Pedicellarie are of two kinds, much like those of Asteriide.
WEST INDIAN STARFISHES 31
Genus CORONASTER Perrier. Coronaster Perrier, Ann. Sci. Nat. Zool., vol. xix, No. 8, pp. 5, 9, 1885; Exped. Trav. et Talism., p. 92, 1894. (Type, C. parfaiti.) Sladen, op. cit., pp. xxxix, 592, 1889.
Delicate starfishes with a small disk and numerous slender rays, covered with long, slender spines in radial rows. The dorsal skeleton is weak, made up of the rows of median and superomarginal plates, connected together by slender transverse ossicles. Each plate of the longitudinal rows usually bears one slender spine.
Both rows of marginal plates well developed and spiniferous. No interactinal plates. Adambulacrals diplacanthid. Large, felipedal, dermal major pedicellariz occur above and _ below. Minor pedicellarie form large cireumspinal wreaths, borne on contractile sheaths. Tube-feet are relatively large, in two rows, not crowded. A pair of rather large peroral spines on the mar- gins of the oblong jaws, with groups of oral marginal pedicel- lariz.
CORONASTER BRIAREUS Verrill, 1914a, p. 49.
Asterias briareus Verrill, Brief Cont. to Zoodl., No. 50, Amer. Journal Sei., vol. xxiii, p. 220, 1882; Notice of Rem. Maine Fauna, in Annual Rep. Com’r. of Fish and Fisheries for 1882, p. 659, 1884, ditto for 1883, p. 540; Amer. Jour. Science, ser. 3, vol. xlix, p. 209, 1895.
Starfish ‘‘near Asterias volsellata,’’?’ Nutting, Narrative Bahama Exped., p- 168, fig. 3, 1895.
Plate i; figures 1, 2. Plate ix, figures 4—4c.
Disk small,rays slender, elongated. Rays variable in number, ten to twelve in the larger specimens, one of the larger has the radu 8™™ and 76™":; ratio 1 :9.5.
The dorsal skeleton is openly reticulated. There is a marked medial radial carina, with larger plates and longer spines. The marginal plates also form ridges. These five rows of plates are connected by slender transverse ossicles, leaving large rectangu- _lar papular areas, with numerous papule in clusters.
The dorsal and marginal spines are long, slender, acute, not very near together. A few arise, also, from the transverse ossi- cles.
The spines are all much alike in form and size. They bear large dense wreaths of small minor pedicellarix, usually on a
32 NATURAL HISTORY BULLETIN
sheath which may be contracted to near the base or extended to near the tip in preserved specimens.
Solitary felipedal or unguiculate major pedicellariz of large size and remarkable in form are scattered on the dermis, above and below; others are on the inner furrow margins. They are hand-shaped, with curved, acute terminal denticles, interlocking like fingers or claws. The valves are wider distally and there are generally about five or six interlocking denticles.
The adambulacral spines are two to each plate, slender, elong- ated. There are no interactinal plates nor spines; the inferomar- ginals join closely to the adambulacrals.
It was taken by the Bahama Expedition at Station 61, off Key West, in 75 to 80 fathoms, and at Station 64, in 110 fathoms, off American Shoal, Florida.
It was dredged by the U. S. Fish Commission, at several sta- tions in 31 to 373 fathoms, between N. lat. 37° 18’ 11” and 36° 41’ 05”.
This is the species described by Professor Nutting, as the most beautiful starfish obtained. See Narrative of Bahama Ex- pedition, p. 168, and figure on opposite plate.
Genus PEDICELLASTER Sars.
Pedicellaster Sars, Oversigt Norges Ech., p. 77, 1861. Perrier, op. cit., 1884, p. 194; Exped. Trav. et Talism., p. 99, 1894. Sladen, op. cit., p. 557, 1889. Verrill, op. cit., p. 202, 1914a.
Delicate starfishes with a small disk and usually five or six slender rays, sometimes more. Dorsal plates form a reticulation. - Ambulacral feet in two rows. Ambulacral groove relatively wide and open. Dorsal and marginal plates are covered with small spinules.
Pedicellariz of both kinds are formed on most, if not all, species when adult.
PEDICELLASTER POURTALESI Perrier. Pedicellaster pourtalesi Perrier, Bull. Mus, Comp. Zool., vol. ix. p. 7, 1881; Etoiles de Mer, p. 194, pl. iii, fig. 4.
Rays very slender, fragile at base, elongated, terete, obtuse. The type had the radi 4™™ and 21™™; ratio, about 1:5. Dorsal plates are delicate mostly in three rows, a carinal and
WEST INDIAN STARFISHES 33
one dorsolateral on each side, united transversely by small ossi- eles. Each principal plate bears a single small spine, marginal plates and spines are similar.
The adambulacral plates are relatively large; each bears two or three small, slender spines in a transverse row.
The madreporic plate is small, with few gyri. Minor pedicel- lariz are dermal, scattered between the dorsal spines.
Recorded by Perrier from four Blake stations, among the Lesser Antilles, in 127 to 250 fathoms. It was also taken by the Albatross in the same region.
Order SpPInvuLosa Perrier.
Stelleride Spinulose Perrier, Nouv. Arch. Mus. Hist. Nat., vi, p. 154, 1884.
Spinulosa Perrier, Exped. Trav. et Talism., pp. 27, 138, 1894.
Phanerozonia and Cryptozonia (pars) Sladen, Voy. Chall., xxx, p. xxiii, 1889.
Spinulosa Fisher, op. cit., 1911b, p. 251. (Table of known families.) Ver- rill, op. cit., pp. 24, 304, 1914a.
This order, established by Perrier, includes eryptozonate (rare- ly subphanerozonate) starfishes, which usually have ;but two rows of ambulacral feet (four in Diplopteraster). The am- bulaeral plates are not crowded. The dorsal plates, generally reticulated or imbricated, spinulose, granulose or naked, often form pseudopaxille or parapaxille, but they rarely, if ever, form true paxille. The dorsal plates are sometimes isolated, rudimentary, or absent in deep-sea genera. The pedicellarie are commonly lacking; when present they are neither forcipulate nor forficulate. They may be pincer-like, with two or more simple valves, and are always dermal.
Perhaps the structure of the pedicellarie is the most positive character for separating certain genera of this group from some of the aberrant Forcipulosa.
The marginal plates are nearly always small and sometimes indistinct; the lower ones are usually the larger; they never form a thick rigid margin; the margin may be acute or rounded. Papulz may occur both on the dorsal and on the actinal sur- faces, but they are restricted to the dorsal surface in most fam- ilies. The odontophores are adambulacral and of various forms. The ambulacral grooves are usually narrow. The ambulacral feet always have a terminal sucker. The ambulacral ampulle may be single (Echinasteride) or double.
34 NATURAL HISTORY BULLETIN
A number of the genera do not have free-swimming larve,* but are known to carry their eggs and larve attached about the mouth, or else in a marsupial pouch (gonocodium) until they become true starfishes, large enough to care for themselves. This habit is usually associated with the lack of pedicellarie. It is conducive to the formation of local varieties. The order may be naturally divided into two suborders:
Suborder [— Avenata Ver., op. cit., 1913; 19144, p. 204.*
This includes the more typical forms in which there is no dorsal, tent-like marsupial chamber for the protection of the eggs and young. The spines are either all free, or partially or wholly webbed together into groups, as in Solasteridw, in which the groups of adambulacral spines form transverse combs complete- ly webbed together, and the dorsal spinules are partially so.
It includes the following families represented in this collection:
Echinasteride ; Solasteridw; Korethrasterida, deep sea; Aster- mde; Poranide. Also: the following extralimital families, be- sides other smaller groups: Acanthasteride; Mithrodide; Py-. thonasteridew, deep sea; Myxasteridw, deep sea.
Il — Suborder VeEuatTa Perrier (as an order).
This group includes only the family Pterasteride. It is re- markable for having most of the spines webbed together in clus- ters and for the remarkable development of a superdorsal mem- brane, more or less completely uniting the paxille together, and usually forming a dorsal marsupial pouch or gonocodium in which the eggs and young are carried. :
Suborder AvELATA Verrill (See above).
Family EcHINASTERIDa Verrill (restricted).
Echinasteride Verrill (pars), Trans. Conn. Acad. Sci., i, p. 343, 1867. Per- rier (pars), Revis. Stell., Arch. Zo6l., iv, pp. 299, 358, 1875.
Echinasterine Viguier, Arch. Zoél. Exper. et Gen., vii, p. 123, 1878 (struc- ture.)
Echinasteride Sladen (pars), Voy. Chall, xxx, p. 535, 1889. Bell, Catal. Echinod., pp. 23, 95, 1890. Perrier, Etoiles de Mer, Nouv. Arch. Mus. Nat. Hist., vi, p. 164, 1884; Exp. Trav. et Talism., pp. 28, 141.,.
* This is the case with the genera Henricia, Pteraster, Hymenaster, and others. * There once misspelled as Alvelata.
WEST INDIAN STARFISHES 30
1894. Ludwig, Fauna Arctica, i, p. 472, 1900. Fisher, op. cit., 1911b, p. 258, analytical table known genera, p. 259. Verrill, op. cit., p. 205, 1814a.
Dorsal skeleton usually formed of small, sometimes over- lapping, reticulated or areolated plates; sometimes they form longitudinal radial rows. Median dorsal row is often indistinct. Upper marginals small, often not specialized. Inferomarginals usually distinct. One or more interactinal rows are usually present. Dorsal and marginal spines may be large and isolated or minute and clustered, but not truly paxilliform. They are often covered with a distinct, thick, dermal membrane, which may contain calcareous granules.
The ambulacral ampullez are usually single, one to each tube- foot.
Ambulacral grooves narrow. Adambulacral spines small, two or several to a plate, usually in a transverse group or a single transverse row. No pedicellarie are known.
The most prominent genera are Echinaster and Henricia, both of which occur in the West Indies.
Genus Ecurinaster Miiller and Troschel, emended.
Echinaster (pars) Miller and Troschel, Monatsber., Berlin. Wis. Akad., April, 1840, p. 102; Wieg. Arch., 1840 (non Gray); Syst. Aster., p. 22, 1842 (pars).
Othilia Gray, Ann. Mag. Nat. Hist., vi, p. 281, Dec., 1840; Synopsis Star- fishes, p. 12, 1866. Fisher, 1911b, p. 260.
Echinaster Perrier, Revis. Stell. du Mus., Arch. Zoél. Exper., iv, p. 364, 1875. A, Agassiz, North Amer. Starfishes, p. 97, pl. x, 1877 (structure of skeleton.) Viguier, Squelette Stell., Arch. Zodl. Exper., vii, p. 123, pl. vii, figs. 1-7, 1878 (structure of skeleton.) Sladen, Voy. Chall., Zool., Xxx, p. 553, 1889. Perrier, Exp. Trav. et Talism., p. 146, 1894. Ver- rill, op. cit., p. 206, 1914a.
Stellate, usually with the disk rather small. Rays normally five, of moderate length, rounded. Larger dorsal and marginal plates strong, lobate, convex in the middle, and having a central mammilla and pit for the attachment of the usually solitary spine. The larger ossicles are united into reticulations by small, rounded, connective ossicles, leaving large papular areas; soli- tary papule may occur in the marginal and actinal regions.
The whole surface is covered in life with a rather thick, soft integument, containing small granules on the spines. Dorsal
36 NATURAL HISTORY BULLETIN
spines are conical, acute, mostly isolated, sometimes in small groups.
Upper and lower marginal plates are not very distinct, with spines like the dorsals. Adambulacral spines form a small oblique or transverse row of two to four; the inner ones are smaller, inserted in the groove, just above the margin.
The genus Othalia Gray, December, 1840, appears to be a syno- nym of Echinaster Miller and Troschel, April, 1840. The type of the latter was E. spinosus* = A. echinophora Lam., both by virtual tautology and by designation. (See foot-note in Miller and Troschel, Syst. Ast., p. 22, 1842.) Hchinaster of Gray, 1866, is Acanthaster of later writers.
This genus is found in all tropical and subtropical seas, chief- ly in shallow water.
ECHINASTER SENTUS (Say) Liitken.
Asterias sentus Say, Journ. Philad. Acad., v, p. 143, 1825.
Othilia aculeata Gray, Annals Nat. Hist., vi, p. 281, 1840; Gray, Synopsis Starfish, p. 12, 1866.
Echinaster spinosus (pars) Miller and Trosch., Syst. Ast., p. 22, 1842. Verrill, Notes on Radiata, p. 343, 1867. Ives, Proc. Acad. Nat. Sci. Philad., for 1890, p. 325. Clark, Echinoids and Asteroids of Jamaica, p- 6, 1898.
Othilia spinosa Agassiz, Bull. Mus. Comp. Zo6l., p. 308, 1869.
Echinaster sentus (Verrill, MSS., 1866). Liitken, Vid. Meddel., pp. 60, [284], 1871; 9 Perrier, Arch. Zool. Exper., iv, p. 366, 1875; A. Agassiz.
North American Starfishes, p. 97, pl. x, figs. 1-6, 1877 (figs. of living and structure.) KR. Rathbun, Echinoderms of Brazil, p. 147, 1879.
Plate xxix; figure 2. Young.
The two marginal rows of plates are pretty regular but not much differentiated from the others in size or form. The upper row usually has only one stout conical spine, with a mammiform base, on each plate. It turns upward proximally, reaching the dorsal side, and leaving a wide interradial intermarginal area, in which there are a number of spineless plates, and usually one short intermarginal row of plates bearing one spine each in a typical specimen from Florida.
8 The name spinosus (Retz.) is not tenable for any species of the genus for Asterias spinosa R. was antedated by A. spinosa Pennant, 1777.
9Liitken here refers to Verrill (correspondence) and to his determina- tion in 1866 (op. cit., p. 348), of sentus.
WEST INDIAN STARFISHES 37
The intermarginal plates are like the upper ones, but extend regularly to near the small adoral area. Some of them some- times bear two, or even three, conical spines, close together, but ordinarily only one. The peractinal plates form a single row of small, oblong, transverse connective ossicles between the infero- marginals and the adambulacrals. Between them are large pap- ular pores, either isolated or in small groups; many of these ossicles are without spines, but some of them usually have a small conical spine, near the outer adambulacrals.
The adambulacral spines are small and subacute, in some- what oblique transverse series of three. They are much smaller and more numerous than the adjacent spines, about four of their plates corresponding to one marginal plate. The inner furrow- spine is much the smaller; the next is nearly as large as the outer one. These last are webbed together basally and thus form a continuous regular border to the furrow. Most of the dorsal spines can be referred to five irregular rows, above the supero- marginals. These are one pretty distinct median radial row, and two less distinct dorso-lateral rows on each side. There are also some irregular interpolated spines on the connective ossi- cles. Many of the larger plates bear two and some three clus- tered spines.
The papular areas are large and the papule are very numer- ous, not only on the dorsal side, but also between the marginal rows, and are present also next to the adambulacrals, where they become less numerous.
The madreporic plate is raised, fiat, wart-like, covered with rough projections, like stout spinules.
The Young. Plate xxix; figure 2.
A young specimen, apparently of this species, was taken by the expedition in shallow water on the Great Bahama Bank. Its radi are 7"™ and 26™™. It has the two rows of marginal spines well developed and pretty regular. There are about eight or nine in the upper series and about twelve in the lower. They are relatively large, conical, acute.
The dorsal spines are similar in size and form. They form, on some of the rays, too pretty definite dorso-lateral rows of five or six, with a few in the median row. In other cases the median
38 NATURAL HISTORY BULLETIN
row has five or six spines and the doso-lateral ones fewer. The papular pores are small and numerous dorsally; few and mostly isolated below the inferomarginal spines. <A row of small inter- actinal plates separates the inferomarginal plates from the adambulacrals and some of these bear rudimentary spines, and many have small solitary papule between them.
The adambulacral plates have three small spines in an oblique row; the inner one is very small and deep in the furrow; the outer ones are webbed together longitudinally. Color, when dry, pale yellowish brown. The madreporite is small, round, rough with minute spinules.
TERATOLOGY.
A regular four-rayed specimen was obtained by the Bahama Expedition at the Great Bahama Bank, in shallow water, May 17, 1893.
The radii are 13™™ and 53™"; ratio, 1:4.1. The dorsal spines are rather numerous, subequal, of moderate size and acute. They do not form definite or obvious rows, but appear as if in about Six or seven zigzag and irregular rows proximally. They are not crowded. :
Papular areas, between them, are large, with many rather large pores; these areas form about six irregular rows. Additional ones exist between the rows of marginal spines and scattered papule occur between the inferomarginals and the adambula- crals, and between the interactinals.
The rows of superomarginal spines are not clearly distinct from the dorsals, being of the same size and form and the rows are irregular. The inferomarginal row is more regular, but the spines are of the same size. This row is near the adambulacral plates and about four of the adambulacral plates correspond to each of these spines in the amount of space occupied. There are about twenty to twenty-four of these spines in each row. Proximally there is a short irregular intermarginal row, and also a row of small interactinals, the latter mostly spineless.
Ihe adambulacral plates mostly bear three partially webbed spines. Two on the outer surface are subequal, one nearly
WEST INDIAN STARFISHES 3g
above the other; the other is much smaller and deeper in the groove; distally there is often another smaller furrow-spine still deeper in the groove. The madreporite is round, elevated, wart- like, and covered with. small rough spinules. The color of the dry specimens is yellowish brown, to dark purplish brown.
Mr. A. Agassiz (op. cit., 1877, pl. x) gave excellent figures of this species from living specimens as well as many structural details of the skeletal parts.
It is the common shallow water species found on our coast, from North Carolina to Florida. In the number and size of the spines and the number of the rows it exceeds E. echinophora, but has many less than E. spinulosa.
Most of the earlier writers, including Lamarck, Miiller and Troschel, and many others combined this with ,E. echinophora, under the specific name ‘“‘spinosus,’’ a name that was often at- tributed to Retzius, 1805 (as Asterias spinosa), which was ante- dated, however, by A. spinosa Pennant. It is doubtful, as shown by Liitken, whether the brief diagnosis of Retzius refers to any Echinaster. In any case it was preoccupied and cannot be used for any species of this genus.
9)
The Asterias spinosa of Say, 1825, was primarily the common Asterias forbesi of our coast, but also included A. vulgaris, for he had specimens from the coast of Maine.
This determination, and also the fact that Say’s A. sentus, was an Echinaster like spinosus of authors, was published by me in 1866. (Proc. Boston Soe. Nat. Hist., vol. x, p. 348) and was earlier communicated to Liitken. I did not publish the combined names at that time.
This species occurs in shallow water on the southern coasts of the United States, from North Carolina to the Florida Reefs, and to Yucatan. It is also common in the West Indies, from the Bahamas to St. Thomas. I have seen it from Cuba, Ba- hamas, and Hayti. Yucatan (Ives), St. Thomas (Liitken), Ba- hia and Pernambuco, Brazil (R. Rathbun), Jamaica (Clark).
The Bahama Expedition took specimens on the Great Bahama Bank, including one four-rayed specimen, described above.
40 NATURAL HISTORY BULLETIN
ECHINASTER SPINULOSUS Verrill.
Echinaster spinulosus Verrill, Proc. Boston Soc. Nat. History, vol. xii, p. 386, 1869. Liitken, op. cit., p. 285 [61], 1871. Ives, op. cit., p. 326, 1890.
Plate iv; figures 1, 2.
This species is characterized by having long, slender, terete, tapered rays, covered with numerous small, conical, subacute or often blunt spinules, in many more or less evident rows, and often standing two or three on a plate. The number in each row may be forty or more.
The number of rows, exclusive of inferomarginals, may be from fifteen to twenty-one in the larger examples. A medium- sized specimen (type) has the radii 12™™ and 72™; ratio, 1:6. The largest specimens have the radii about 14™ and 80™.
A typical specimen of medium size from West Florida has these characters; the two rows of marginal plates can readily be distinguished in this and in most other dried specimens, not only by their larger size and more rounded form, but also by a finely granulated and well defined patch on the convex part of each plate and mostly below the single spine, but often including most of the surface of the plate. Over this granular surface the in- vesting dermal membrane is so thin that the granules can usually be seen without preparation, and sometimes they appear as if naked. They are not found on the intermarginal and peractinal plates.
The two rows of marginals are pretty regular and can be traced to the tips of the arms, each bearing one small conical spine. On the distal third of the ray the two rows are contigu- ous; on the proximal part they are separated by two or three short, irregular rows of intermarginals, each bearing a conical spine. In the interradial area additional spineless plates are interpolated, broadening the area, for here the upper ane row turns upward to the dorsal side.
Many papular pores are found between the marginals and intermarginals, but none below the inferomarginals in most speci- mens.
Granulated patches also occur on the plates of the principal dorsal rows.
The interactinal or peractinal plates and spines form a pretty regular row, extending nearly to the tips of the rays,
WEST INDIAN STARFISHES © 41
where they become very small. Most of them bear one small conical spine, close to the outer adambulacral. The outlines and sutures of these plates are usually concealed by dermis; some- times proximally, a number of them are spineless and visible, when they appear as narrow, flat, oblong, transverse ossicles, ex- tending between the inferomarginal and adambulacral plates. The adambulacral plates have two small furrow-spines, one above the other. The most interior one is the smaller; on the outer margin, and not quite in line with the others, there is a somewhat larger and stouter but small, conical spine. Those on consecutive plates are webbed together for about half their height and also webbed to the upper one of the furrow-series, forming a continuous row.
The madreporie plate is prominent, flattish, wart-like, covered with thin, rough, irregular ridges and points.
Aleoholic and well dried specimens of this species are usually purplish brown, but many, when dried, become yellowish brown. In life it is reddish brown.
This species is abundant on the west coast of Florida, at Tam- pa Bay, Egmont Key, and many other localities.
In the Yale Museum, besides the types from Egmont Key, there are specimens received from the U. S. Nat. Mus. from the following places: Pine Key, Cedar Key, Goodland Point, and Marco, W. Florida; Gulf of Mexico, S. of Alabama (stations 2370, 2373, 2387, steamer Albatross), in 23 to 32 fathoms. I have seen no West Indian specimens nor any from East Florida.
This is more nearly allied to EL. braziliensis than to either of the other species.
It has many more rows of spines than E. sentus, and many more spines in each row, while the spines are also much smaller and more slender. The rays are also longer and more slender.
ECHINASTER BRAZILIENSIS M. and Tr.
Echinaster braziliensis M. and Trosch. (pars), Syst. Aster., p. 22 (not the figure, pl. i, fig. 4). Liitken, op. cit., p. 9 [67], 1859; p-. 284 [60], 1871. Verrill, Notes on Radiata, pp. 343, 368, 1868. Perrier, Archiv. Zool. Erp., vol. iv, p. 367, 1875. RK. RAGhbon: Trans. Conn. Acad Sci., vol. v, p. 148, 1879. Ludwig, Mem. Cour. Acad. Roy. Belg., xliv, p. 7, 1882. Ives Proc. Acad. Nat. Sci. Philad., for 1890, pp. 324, 326, pl. vili, figs. 16-18, 1890.
Othilia braziliensis A. Agassiz, Bull. Mus. Comp. Zo6l., vol. i, p. 308, 1869 (Florida, shore to six fathoms.)
“- <2 Ve
42 NATURAL HISTORY BULLETIN
Plate xxvi; figure 1.
The ratios of the radii are about 1:5. The rays are terete and rather slender. The spines are numerous, small, acute-conieal, not much larger than the outer adambulacrals. On the rays they form about nine to eleven rows in the adult, with thirty or forty in the median row, in specimens 112™™ in diameter. The adambulacral plates have three spines.
The spines are larger and fewer than in E. spinulosus, but much more numerous and smaller than in E. sentus.
Color in life rust-red, paler below. (Ives.)
The figure of a portion of a ray given by Miiller and Troschel does not agree with their description. It probably represents E. sentus. }
This Brazilian species has been recorded from shallow water on the Florida coast by Mr. A. Agassiz; from Jamaica, by Liit- ken and Verrill; and from Yucatan by Ives. On the Brazilian coast it has been recorded from Pernambuco (Rathbun) to Rio de Janeiro (Perrier). The Yale Museum has it from Bahia and Pernambuco (coll. C. F. Hartt, Nos. 5206, 5208, 4559). Also a specimen, perhaps of this species, from Cuba (No. 263). Mr. Rathbun doubts the Rio de Janeiro record. Some of the other records are also doubtful, owing to confusion with allied species. That of Florida is doubtful. I have seen no specimens from Florida nor from the Bahamas.
ECHINASTER ECHINOPHORUS (Lam.) Perrier.
Asterias spinosa (pars ?) Retzius, Dissert. sist. spec. cog. Asteriarum, p. 18, 1805 (non Asterias spinosa Pennant 1777).
Asterias echinophora Lam., Anim. sans vert., ed. I, vol. ii, p. 560, 1816.
Othilia spinosa Gray, Ann. and Mag. Nat. Hist., vi, p. 281, Dec., 1840; Synopsis Starfishes, p. 12, 1866.
Echinaster spinus Mill. and Trosch. (pars), Syst. Ast., p. 22, 1842; Liitken, Vidensk. Meddel., p. 90, 1859. Perrier, Pédicell., p. 57, 1869.
Echinaster (Othilia) crassipina Verrill, Trans. Conn, Acad., i, p. 368, pl. iv, fig. 7, 1868.
Echinaster crassispinus Liitken, op. cit., p. 283 [61], 1871. Ives, op. cit., 1890, p. 326.
Echinaster echinophorus Perrier, Arch. Zéol. Exper., iv, p. 364, 1875. R. Rathbun, Echinod. Brazil, p. 147, 1879. Ives, Proc. Acad. Nat. Sci. Philad. for 1889, p. 171.
WEST INDIAN STARFISHES 43
The rays are angulated and not very long; not slender. Radii of a medium sized specimen 12™ and 47™™; ratio about as 1:4. On the dorsal and lateral surface of the rays there are about five irregular rows of rather distant, relatively large, stout, acute, conical spines, on mammilary bosses of the principal plates, and partially covered by a thick dermis. The two outer of these rows may be reckoned as superomarginals, but they are like the others.
The inferomarginals, which are on the actinal side near the adambulacrals, but separated from them by a naked lane, form a more regular row of about fifteen, somewhat larger than the dorsals, but similar in form.
The disk bears a few spines, like the radials. Beneath, the disk is covered by a smooth, naked dermis, without spines. Pap- ular pores are numerous in large groups on the dorsal side; fewer on the sides and beneath.
The adambulacral spines are two to a plate; the outer one is much the larger and they form a close, regular, longitudinal row, partly webbed; the inner one is much smaller, not half as long, and situated deep in the furrow. Above description is from a Brazilian specimen.
In life this species is usually bright red or crimson; it be- comes dull reddish brown in alcohol, and usually dark purplish brown when carefully dried.
Not uncommon in shallow water and on reefs throughout the West Indies. It extends to the Abrolhos Reefs, Brazil, and to Yucatan. Abundant in Bay of Bahia, Pernambuco, on rocky bottoms, ete. (R. Rathbun). Yucatan (Ives).
T have not seen it from Florida, nor from the Bahamas. The Yale Museum has it from the Abrohos Reefs, Brazil (coll. C. F. Hartt, No. 1527, type of FH. crassispina), and young specimens, ap- parently of this species, from Parahyba de Norte (coll. Hartt, Nos. 5209, 5210, 5211).
ECHINASTER MODESTUS Perrier.
Echinaster modestus Perrier, op. cit., p. 7, 1881. Etoiles de Mer, p. 206, pl. iii, fig. 7, 1884.
The type was evidently a young specimen. Its radii were 6=™™ and 36™: ratio, 1:6.
ot NATURAL HISTORY BULLETIN
The dorsal surface is formed of openly reticulated plates, leaving large sunken papular areas, containing groups of three or four papule. The investing dermis is thick.
The dorsal, dorsolateral, and lateral spines are very small and form about nine irregular rows, with scattered spines between them. The marginals are not distinct.
Besides the nine rows there is a regular row of interactinal spines, which ceases at about midlength of the ray. The adam- bulacral plates have a short, thick, obtuse furrow-spine, and ex- terior to it, an equally larger but rather shorter spine; outside of this there may be, in the same transverse line, one or more small spines.
Taken by the Blake at two stations among the Lesser Antilles, in 120 to 123 fathoms, and in the Gulf of Mexico, off West Flor- ida, in 101 fathoms.
This has much smaller spines than specimens of E. spinulosus of the same size, and the surface appears more areolated.
THYRASTER SERPENTARIUS (M. and Tr.) Ives.
Echinaster serpentarius M. and Trosch., Syst. Ast., p. 24, 1842. Perrier, op. cit., vol. iv, p. 370, 1875. Liitken, op. cit., p. 285 [61], 1871. Thyraster serpentarius Ives, op. cit., p. 329, pl. viii, figs. 19-21, 1890.
The genus Thyraster, established for this species by Ives (1890), is separated from Hchinaster especially by the character of the dorsal and marginal plates, which are quadrangular, or rhombic, overlapping by their angles, and placed in regular longitudinal rows. They bear three or four small, thorny spines, in a row, on the distal edge. The adambulacral plates mostly bear a row of three or four small spines on the distal margin, with a small furrow-spinelet. The marginal plates usually have three spines on the distal edge, rather larger than those of the dorsal plates, but still quite small. Diameter of the type, 4 inches.
It is a Mexican species, recorded by Miiller and Troschel, and later by Ives and others, from Vera Cruz.
Genus HEnrIcIA Gray.
Henricia Gray, Ann. Mag. Nat. Hist., vi, p. 184, November, 1840; Synopsis Starfishes, p. 5, 1866. Bell, Ann. and Mag. Nat. Hist., vi, p. 473, 1890;
WEST INDIAN STARFISHES 45
Catal. British Echinod., p. 95, 1892. Fisher, 19116, p. 266. Verrill, op. cit., p. 209, 1814a.
Cribella Forbes, Brit. Starfishes, p. 100, 1841, or December, 1840.
Echinaster (pars) Miller and Troschel, April, 1840; Syst. Aster., p. 22, 1842.
Cribrella Liitken, Gronl. Echinod., p. 30, 1857. Norman, Ann, and Mag. Nat. Hist., xv, p. 124, 1865. Verrill, Proc. Boston Soc, Nat. Hist., x, p. 345, 1866. Perrier, Revis. Stell. Mus., Arch. Zool. Exper., iv, p. 373, 1875. A. Agassiz, North Amer. Starfishes, p. 113, pl. xviii, 1877 (structure of skeleton). Viguier, Squellette des Stell., Arch. Zool. Exper., vii, p. 126, pl. vii, figs. 8-15, 1878 (odontophore). Sladen, Voy. Chall., xxx, p. 540, 1889. Perrier, Exp. Trav. et Talism., p. 143, 1894. Ludwig, Fauna Arctica, p. 473.
Stellate starfishes usually with five terete, slender rays; some- times with short rays and wide disk. Sometimes with six or more rays in autotomous species.
Dorsal plates are numerous and small, united into a rather closely reticulated structure, bearing numerous papular pores, either isolated or in small groups. Dorsal spinules minute and nearly uniform, crowded in divergent clusters on the convex ossicles of the dorsal, lateral, and actinal surfaces.
The madreporic plate is inconspicuous and usually spinulose ; sometimes two are present in autotomous species. There is no thick dermis over the plates.
Supermarginals are small, often indistinct, and like the dor- sals. Inferomarginal plates distinct, often larger than the su- peromarginals, contingent with the latter distally, but often sep- arated proximally by one, two, or more interpolated rows of marginal ossicles, varying in number and extent, and thus caus- ing great variations in the thickness and taper of the rays. In- teractinals and inferomarginals similar and closely joined, con- vex, and covered with small spinules; the rows are separated by papular pores, which may also occur between the interactinal and adambulacral plates.
The two rows of marginal plates, the interactinal and the adambulacrals agree nearly or quite in radial length, at least proximally, so that they form regular transverse series.
The adambulacral plates are transversely oblong. Their actinal face is covered with numerous spinules in two or more transverse rows, or in a compact group; those near the furrow margin are longer, and one to three may stand on the margin; there is one
46 NATURAL HISTORY BULLETIN
or sometimes two furrow-spines within the groove. Pedicellarie have not been found in any species.
The eggs and young are carried in clusters around and below the oral region until the young take on the stellate form pro- vided with sucker-feet.
This genus differs from Echinaster in the absence of a thick external integument; in the covering of minute spinules; and in the nearly uniform small ossicles of the dorsal and lateral skel- eton, the connective ossicles being almost indistinguishable from the primary series; and in the quadrangular form and regular arrangement of the marginal and interactinal plates.
According to the accepted rules of priority it is necessary to adopt Henricia instead of Cribrella as the name of this genus, for it has at least a month of priority. Cribrella of Agassiz, 1835, was a different group.
HENRICIA ANTILLARUM (Perrier). Verrill, 1914a, p. 210.
Cribrella antillarum Perrier, Bull. Mus. Comp. Zool., vol. ix, p. 8, 1881; EKtoiles de Mer, p. 207, pl. ui, fig. 3, 1884.
Rays, in the type, five, rather long, slender terete, regularly tapered. Radi 7™™ and 42™™; ratio, 1:6.
Dorsal ossicles small, reticulated, leaving numerous papular areas mostly smaller than the ossicles and containing a single papula. They are covered with numerous minute, short spinules. Madreporic plate single, raised, about midway between the cen- ter and margin of the disk, and covered with minute spinules similar to those of the dorsal ossicles. :
Two marginal rows of plates are well developed, rectangular, closely covered with minute spinules.
There are two or three rows of interactinal plates proximally, but only one reaches the middle of the ray. Their larger plates are quadrangular and of the same length as the marginals and adambulacrals.
The latter are transversely oblong; their actinal surface is covered with small spinules; five or six of them become abruptly larger near the furrow margin and stand in two rows, two or three in each row.
Dredged by the Albatross at station 2671, off Georgia, in 280 fathoms. (No. 18,392, N. Mus.)
WEST INDIAN STARFISHES 47
Perrier recorded it from five Blake Exped. stations, in 127 to 734 fathoms, among the Lesser Antilles.
The specimen from off southern Georgia (No. 18,392) has the proximal interactinal plates transversely oblong or rectangular, and similar to the marginals in size and form.
The adambulacral spines are strongly differentiated ; the three nearest the groove show a decided approach to the formation of a true marginal comb. Those next to these are about twice as long as those farther back. There is a small furrow-spine deep in the groove, and distally there may be two.
Small isolated papular pores form a row between the inter- actinals and marginals, and also between the former plates; a few are located proximally between the interactinals and mar- ginals.
HENRICIA SEXRADIATA (Perrier) Ver.
Cribrella sexradiata Perrier, Bull. Mus. Comp. Zool., vol. ix, p. 8, 1881; Etoiles de Mer, p. 209, pl. iv, fig. 6, 1884. Henricia sexradiata Verrill, N. Pacific Starfishes, p. 371, 1914a.
Plate xi; figure 7. Details.
Rays slender, variable in number, often unequal in size; most frequently there are six of which about three are shorter than the others, due to autotomy.
The type described by Perrier had the radii 6™™ and 138™™; ratio, 1:2.16; others have relatively longer rays.
The dorsal surface is covered with numerous small reticulated ossicles, carrying very small spinules; the papular areas are smaller than the ossicles and mostly have a single papula. Two madreporic plates are usually present, small, granulated, one each side of a single ray.
The two rows of marginal plates are very distinct; the plates are quadrangular and covered with nearly uniform minute spin- ules, a little larger than those on the dorsals. There is a short row of peractinal plates of similar size and form, proximally.
The adambulacral plates are transversely oblong; they bear, near the inner margin, about five small obtuse spinules, larger than the others, and forming two longitudinal rows; two belong to the inner or marginal row; three to the outer one. The rest
48 NATURAL HISTORY BULLETIN
of their actinal surface is covered with smaller, short, obtuse spinules, resembling granules.
Perrier recorded this species from the Blake Expedition from off West Florida,’?® in 101 fathoms, and off Barbados in 150 fathoms. I have seen it, also, from Blake station 297, in 123 fathoms, off Barbados. The Bahama Expedition dredged it off Havana, in about 200 fathoms.
This may be only a variety or form of H. antillarum. Such differences as have been noticed may be due to the variations caused by autotomous division. The possession of six rays can- not be regarded as a specific character, especially in autotomous species.
HENRICIA MICROSPINA Verrill, sp. nov.
A small species with a small disk and five very slender terete rays. Radi of the type 4.5™ and 23™; ratio, 1:5.1.
The dorsal surface is evenly covered with very small irregular plates many of them roundish, forming a finely but rather evenly riticulated structure, in which the plates and ossicles are narrower than the sunken papular areas between them. The papule are mostly solitary. The madreporic plate is small, somewhat raised, nearly destitute of spinules.
The spinules are remarkable small and thin, almost micro- scopic in size, very slender and short. They are rather nu- merous on the larger plates, but are not crowded, partly owing to their thinness. :
Plates of both marginal rows are very distinct, transversely
oblong, nearly equal, and covered with numerous very minute, spaced spinules like those of the dorsals but a trifle larger. The two rows are separated only by a narrow naked groove distally, but at the base of the rays a few intermarginal plates lhe between them. : The interactinal plates form a single regular row. They are squarish, about half as large as the marginals, and spinulated in the same way. There is a simple row of minute papular pores each side of the interactinal row of plates.
10 Perrier gives this station erroneously as No. 47. It should be No. 40.
He also errs in giving the longitude as 33° 21’ East. It should read 83° 21’ East. It is in N. lat. 25° 33’, Northwest of Dry Tortugas.
WEST INDIAN STARFISHES 49
The adambulacral spines are very characteristic. There is a strictly marginal comb, directed horizontally across the groove, and composed proximally usually of three slender spines, webbed at base. Distally these spines are reduced to two and near the tip of the ray to one. Deep in the groove there is sometimes a minute furrow spine, especially distally, but prox- imally it is usually lacking or rudimentary.
On the actinal face of the plate there are numerous minute spinules, of which four to six, near the inner margin, become de- cidedly longer than the rest.
The adambulacral plates and spines project strongly into the groove, but are so much separated serially that they leave large rounded interspaces for the passage of the sucker-feet, which appear to be larger than usual.
Taken by the Albatross at station 2655, in 338 fathoms, north of Bahamas, N. lat. 27° 22’; W. long. 78° 7’ 30”. (No. 18,390, Nat. Mus.)
Family SoLasTEeripa Perrier.
Solasterine (sub-family of Echinasteride) Viguier, Squellette des Steller- ides, Nouv. Arch. Zool. Exper., vii, p. 133, pl. viii, figs. 1-7, 1878, (structure).
Solasteride Perrier, Etoiles de mer, p. 210, 1884. Sladen, Rep. Voy. Chall., Zool., xxx, p. 442, 1889. Perrier, Exped. Trav. et Talism., p. 151, 1894. Fisher, op. cit., 1911b, p. 305. Verrill, op. cit., 1914a, p. 242.
Echinasteride (pars) Perrier, Revis. Stell., Nouv. Arch. du Mus., iv, p. 358, 1875.
Rays varying in number from five to thirteen or more. Dorsal skeleton generally formed either by openly reticulated or loosely imbricated (rarely detached) ossicles, which have a convex or elevated central boss, bearing a cluster of slender paxilliform movable spinules, webbed together into stellate or pencillate groups (pseudopaxille). Both series of marginal plates are usually distinguishable, at least distally, alternate or opposite, or nearly in one line. The upper ones are often the smaller and essentially like the dorsal pseudopaxille. Both rows bear paxilliform spinelets.
Actinal interradial pseudopaxille are generally present and similar to the dorsals. Adambulacral plates are transversely elongated and bear, each, one to five or more spines, usually
20 NATURAL HISTORY BULLETIN
webbed, in a longitudinal row on the inner margin of the groove and one or more clusters or a transverse webbed series or comb of movable spines on the outer surface. True pedicellarie are not known in most species, but large specimens sometimes have a few small bifid spines in the furrow series, and in S. stimpsoni a few minute simple bivalve pedicellariz sometimes occur on the dorsal paxillar areas. Sladen (1888 and 1889) divided this family into two subfamilies: Solasterinw and Korethrasterine. In this he was followed by Perrier (1894, pp. 154, 158). Later these have been considered separate families.
Genus LoPHASTER Verrill.
Lophaster Verrill, Amer. Journ. Sci. ser. III, xvi, p. 214 (type, L. furcifer), 1878. Expl. Albatross in 1883. Ann. Report U. S. Fish Comm., pp. 531, 541 [29, 39], pl. xvi, figs. 49, 49a, 1885. (L. furcifer); Amer. Journ. Sci., xlix, p. 201, 1895. Sladen, Voy. Chall., xxx, pp. 458-461, 1889. Fisher, op. cit., 1911b, p. 334.
Lophaster Verrill, op. cit., 1914a, p. 366.
Solaster (pars) Danielssen and Koren, Asteroidea, Norske Nordhavs-Expd. Zool., xi, p. 47, pl. viii, fig. 12; pl. ix, figs. 9-11, 1884 (structure). Sarkaster Ludwig, Mem. Mus. Comp. Zool., vol. xxxii, p. 185, 1905 (t.
Fisher).
Rays nominally five, sometimes six. Disk not very broad. Dorsal surface covered with nearly uniform paxilliform plates (parapaxille) which have a columar or raised boss, and a lobed polygonal or cruciform base; their lobes are articulated by over- lapping, without intermediate ossicles, and thus form a closely reticulated structure with polygonal or subquadrate meshes, and usually solitary papule.
Marginal plates of both series well developed, in regular series, parapaxilliform, like the dorsals, but larger.
Interactinal plates closely united in interradial angles, with paxilliform spinules; a single row usually extends far along the rays in adult specimens, with or without spinules. Adambulac- ral plates have a furrow series of webbed spines, and a trans- verse comb on the actinal surface.
Species of this genus have been found in deep water, in all the oceans.
WEST INDIAN STARFISHES d1
LOPHASTER RADIANS Perrier.
Korethraster radians Perrier, Bull. Mus. Comp. Zool., vol. ix, p. 12, 1881. Korethraster hispidus Perrier, Etoiles de Mer, p. 212, pl. vi, figs. 9-11, 1884 (non W. Thompson). Solaster radians Perrier, Etoiles de Mer, p. 275 (in explanation of plates),
pl. vi, figs. 9, 10, 11, 1884. Lophaster radians Perrier, op. cit., pp. 167-170 (in lists of species), 1884. Korethraster? radians Sladen, op. cit., p. 459 (footnote), 796, 1889.
Plate v; figure 2. Plate vii; figure 3. Plate xi; figures 1—1c.
Form stellate with a rather broad convex disk and short sub- acute rays. Radii of the largest (sta. 64), 17™™ and 40™"; ;ratio, 1:2.4. It occurs with four to six rays, usually five. The dorsal skeleton is rather openly reticulated with large quadrangular papular areas and numerous large grouped papule.
The pseudopaxille are rather large and high, with an enlarged convex summit, which is covered with numerous (often forty to fifty) very slender, divergent, thorny spinules; those of the mar- ginal fringe are longer and interlock across the spaces interven- ing between the plates. The spinules are hyaline and often forked at the tip, and diverge in all directions.
On the rays there is no evident median series, nor do the plates form many regular transversely oblique rows on the sides. The bases of the plates are mostly deeply four-lobed or substellate. Their arrangement is evidently in radial rows.
The two marginal rows of plates are very evident and pretty regular, and nearly opposite. The upper ones are rounded and very similar to the larger dorsals, and higher than the adjacent lateral ones. The inferomarginals are high, elliptical and nearly twice as large with a correspondingly larger number of terminal rough spinules on the convex summit.
There is a row of small peractinal plates, extending to about the distal fourth of the rays, and a few other interactinal ones on the small interradial areas. These have very slender, thorny spinules.
The adambulacral plates have a prominent convex inner mar- gin, which bears a regular compact row of four or five long, slender, thorny, subhyaline furrow-spines, the middle ones long- est. Their actinal surface bears a row usually of three or four similar rough spinules, but rather stouter. In this dry specimen there is no very evident web between these spines, except near
o2 NATURAL HISTORY BULLETIN
their bases. It is more evident in others, but it is thin and delicate.
The oral plates are large and have a broadly and evenly rounded margin, that of the pair combined forming a semi- circle, and bearing along the edge a very regular compact row of eighteen slender subequal spines, nine to each plate. Epioral spines are also numerous and slender.
Ambulacral sucker-feet and their pores are very large, reg- ularly biserial. (
The type deseribed by Perrier was smaller, radii 4™™ and 10™™.
Perrier, in his report on the starfishes of the Blake Expedition (1884, pp. 167, 212, and 275), placed this species in three dis- tinct genera, and under two different specific names (without ex- planations or cross references), as shown in the synonymy given above. His description was given under ‘‘ Korethraster hispidus, sp. nov.’’; but it was a reprint of his earlier description (1881), which was given under Korethraster radians.
The name, Lophaster radians, occurs in his lists of species dredged; and Solaster radians occurs only in the explanation of plate vi. There is no reference to these figures in connection with the description. I am not aware that he corrected these references in any other later works.
The type of this speciés, described by Perrier, was quite young (radii 4™ and 10™™), only about one-fourth as large as the one now described. This will account for various differences.
Perrier recorded this species from off Barbados, in 56 fathoms, and off Havana, in 80 fathoms. It was also dredged at several stations in the West Indies, by the ‘‘ Albatross.’’
It was taken by the Bahama Expedition at several stations. The largest, described above, was from station 64, in 110 fath- oms, off Florida. Smaller ones occurred at sta. 28, off Sand Key, in 116 fathoms; and at sta. 56, Pourtales Plateau, in 200 fathoms. Another (radii 10™™ and 33™™) was from station 62, off Florida, in 80 fathoms.
Genus SoLASTErR Forbes.
Solaster (pars) Forbes, Mem. Wern. Soc., viii, p. 120, 1839; British Star- fishes, p. 109, 1841. Gray, Ann. and Mag. Nat. Hist., vi, p. 183, No- vember, 1840; Synopsis, p. 4, 1866. Verrill, Proc. Boston Soe. Nat.
WEST INDIAN STARFISHES 3)
Hist., x, p. 345, 1866 (Solaster and Crossaster first separated). Agassiz, North American Starfishes, pp. 111, 112, 1887 (structure). Perrier, Ex- ped. Trav. et Talism., p. 154, 1894. Sladen, op. cit., p. 450, 1889. Verrill, North Pacific Starfishes, p. 242, 1914a.
Solaster (pars) Viguier, Nouv. Arch. Zool, Exper., vii, p. 138, 1878 (struc- ture).
Solaster (pars) Danielssen and Koren, Norw. N. Atlantic Exp., Asterioidea, pp. 42, 52, 53, 1884 (structure). Fisher, op. cit., 1911b, p. 306. Crossaster (pars) Muller and Troschel, Monatsb. Preuss. Akad. Wiss., Ber-
lin, April, 1840, p. 103.
This genus includes large, handsome, multirayed species, usually with a broad disk and slender rays.
The rays are seven to fifteen, variable in each species, but most frequently ten to twelve. Dorsal ossicles small, mostly slightly four-lobed, sometimes stellate, usually reticulated, or in quin- cunx, on the disk and proximal part of the rays, but closely im- bricated near ends of the rays, and often arranged in regular quineunx order on the sides of the rays. They are convex or mammillate, with a central boss, and bear a stellate or fasciculate cluster of slender spinules, webbed together, and usually enclos- ing one or several central spinules, connected by the web, form- ing pseudopaxille. The outer circle of webbed spinules often forms a funnel-shaped structure in life, or when well preserved ; but the spinules are movable, and in dry specimens they are usually mostly collapsed or tipped over, thus forming irregular groups or pencils of small spinules, from five to twenty or more in a group.
The superomarginal plates are small and close to the larger inferomarginals, usually alternating. They are frequently hard- ly distinguishable from the dorsal pseudopaxille, except near the ends of the rays, but they extend regularly to the apical plate.
Inferomarginals are much larger and more elevated, usually transversely oblong, forming a rather conspicuous row. They bear a large number of paxilliform spinules in two or more transverse rows. Adambulacral plates usually have three to six shorter groove-spines, webbed together, and an actinal transverse row of four to nine or more longer spines, also webbed.
The interradial actinal areas are small, but distinct, and bear pseudopaxille, much like the dorsal ones.
Papule are numerous on the dorsal surface, but stand singly
o4 NATURAL HISTORY BULLETIN
or in small groups. A single row of peractinal pseudopaxille usually extends along the proximal part of the rays, to about the middle.
SOLASTER CARIBBZUS Verrill, sp. nov. Plate xxviii; figures 1—la. Type.
Disk rather large; rays seven or eight; most often eight, rather short. Radu 14™ and 36™:; ratio, 1:2.6.
The rays are rounded above and taper regularly. They are closely covered with very small, elevated pseudopaxillx, having very small spinules. Papule are numerous, but mostly solitary. Both rows of marginals are distinct and alternate. The upper ones are much the smaller, compressed, and not much larger than the dorsals.
The inferomarginals are more than twice larger, transversely oblong, and bear two or more rows of short spinules.
The adambulacral plates have an inner or furrow-comb of three or sometimes four graded, webbed spines, the middle ones longest; and a transverse actinal comb of about four larger ta- pered spines, of which the second and third are the larger.
The jaws have a peroral row of about eight relatively long, tapered, subacute, graded spines (four on each side), of which the two middle ones are considerably longer and larger than the next, while the latter are about twice as long as the third.
All the specimens are quite young. Taken by the Albatross at several stations in the West Indies.
Family KoRETHRASTERIDZ Dan. and Kor.
Korethrasteride Danielssen and Koren, Asteroidea, Norske Nordhavs-Expd. Zool., xi, p. 99, pl. xii, figs. 1-14, 1884 (structure). Fisher, op. cit., 1911b, pp. 252, 340.
Korethrasterine (subfamily of Solasteridz) Sladen, op. cit., p. 462, 1889. Perrier, Exped. Tray. et Talism., pp. 151-154, 158, 1894 (deser. and analytical table).
Stellate starfishes with short rays and convex disk, fiat below. Dorsal surface covered with thin, flat, roundish plates, overlap- ping by their edges, with a boss in the middle, bearing a fascicle of long, divergent spinules. Inferomarginal plates are trans-
WEST INDIAN STARFISHES 19)
versely elongated, their outer ends forming the margin, and bear- ing spines.
Adambulacral plates correspond with the inferomarginals and join them. They bear a transverse row of slender spines, usual- ly without webs. Interactinal plates are usually lacking.
Jaws carinate, with a row of several adoral spines and a larger terminal pair; actinal surface with a pair of larger epioral spines.
Dorsal papule are said to be lacking in the type genus; they are present in Remaster.
Genus REMASTER Perrier. Type R. palmatus Per.
Korethraster (pars) Perrier, op. cit., 1881, p. 12; Etoiles de Mer, p. 211, 1884. Remaster 12 Perrier, op. cit., pp. 150, 161, 1894 (as subgenus).
Dorsal plates have a four-lobed base, the lobes overlapping, leaving large papular pores between them; from the central boss arises a divergent circle of slender webbed spinules.
Inferomarginal plates form the border and bear marginal spines. Two or three rows of imbricated interactinal plates on the rays, corresponding to the adambulacrals, and bearing about two webbed spines. The jaws have large epioral spines.
REMASTER PALMATUS Perrier.
Korethraster palmatus Per., Bull. Mus. Comp. Zool., ix, p. 12, 1881; Nouv. Arch. du Mus., vi, pp. 311, 376, pl. vii, figs. 5, 6, 1884. Sladen, op. cit., pp. 463, 796, 1889.
Korethraster Nutting, Narrative, p. 167, 1895.
Remaster (subgenus) palmatus Per., Exp. Trav. et Talism., pp. 158, 161 (redescribed), 1894.
Plate xi; figure 2.
The type of Perrier had the radii 4.7™™ and 14™™; ratio, 1:3.
The form is stellate, with a swollen disk and short rays.
On the dorsal surface a deep interradial groove runs up to the dorsal pore, dividing the disk into five parts. The dorsal pore is surrounded by granules, but has no visible plates.
Five primary radial plates are distinguishable. The rest of the radial areas are covered by squarish plates, with emarginate
12 The name refers to the large flat oral spines (t. Perrier.)
56 NATURAL HISTORY BULLETIN
sides and rounded angles, and imbricated by the two distal lobes of each, overlapping the edge of the two plates beyond it. Thus each plate joins four others.
Each dorsal plate has a round central boss, carrying a fascicle of ten to twelve long (about 1™™), slender, movable, webbed spinules capable of becoming either divergent or fasciculate. Two large isolated papular pores are situated on the adoral side of each plate in the notches.
The adambulacral plates are short, but elongated transversely. Each has a single small furrow-spine, and on its actinal side three large flat, striated spines, truncated and fringed at the tip, -each of which seems to be composed of three united slender spines. These form three longitudinal rows.
Outside of the adambulacrals is a range of imbricated plates, corresponding to them in length, but a little oblique. Each of these bears a fiat fascicle of coalesced spinules like those of the adambulaeral plates. These are succeeded by an outer row of similar, but less regular, plates which form the margin of the under side.
Kach jaw-plate bears a large, flat, lanceolate, hyaline spine, turned outward and appressed. Other oral spines are small and webbed together.
Ambulacral feet are rather large, in two regular rows, with well developed suckers.
The type was taken by the Blake off Barbados, in 163 or 200 fathoms, with three others; a young one was taken near the same place in 76 fathoms. It was also dredged in the West Indies by the Albatross.
Taken by the Bahama Expedition at station 56, on the Pour- tales Plateau, N. lat. 20° 16’, in 200 fathoms (3 specimens).
Family ASTERINIDA.
Asterinide (pars) Gray, Ann. and Mag. N. Hist., vi, p. 238, 1840; Synopsis, p. 15, 1866. Perrier, Revis. Stell., iv, p. 291, 1875; v, p. 209, 1876. Viguier, Squellette des Stell., Arch. Zool. Exper., vii, p. 205, pl. xiv, figs. 1-13, 1878 (structure). Sladen, Voy. Challenger, xxx, p. 374, 1889. Perrier, Expl. Trav. et Talism., pp. 141, 163, 1894. Fisher, op. cit., 1911, p. 253 (Table of genera). Verrill, Revision Asterinine, Amer. Journal Science, vol. xxxv, p. 477, 1913 (Table of genera); Starfishes North Pacific Coast, p. 262, 1914a.
WEST INDIAN STARFISHES oT
Body usually rather flat, often thin, sometimes stellate with long rounded rays; usually with five or six short rays, rarely up to eight. Margins usually thin and formed mainly by the inferomarginals; superomarginal plates small, usually scarcely larger than the adjacent dorsals, generally with a comb or cluster of spinules. Dorsal plates usually lobed or cross-shaped, flat and more or less imbricated, sometimes not imbricated ; generally covered with minute spinules, often in tufts or combs; some- times covered with a soft, naked, or granular dermis. Ventral side flat; interactinal plates angular, flatish, imbricated or close- ly united, usually covered with small combs or tufts of spinules, sometimes with only one; these plates form regular oblique rows. Adambulacral spines simple, divergent; generally in two sets; the furrow-spines form small, usually webbed combs of two to eight. Those on the outer surface are in groups or fans, like the interactinals, or single. Pedicellariz usually lacking; when present they are two-bladed, erect, forficiform.
There is generally no single, dorsal, median, radial row of plates; its place is taken by two, three or more alternating rows; the larger of these plates have a papular pore or several pores under the proximal edge. In some eases, especially while young, there is a symmetrical median row of plates. These larger plates usually show only one of the edges and part of one side; seen from the inside they appear much larger, often four-lobed, and obliquely imbricated. The dorsal plates of the outer inter- radial areas, in many species, have conical or pillar-like processes extending downward and meeting similar uprising processes from the outer interactinal plates, as stalactites meet stalagmites; or they may not join directly, but be united by an intermediate ossicle. In some genera they are differently joined. In typical Asterina, and doubtless in many other genera, if not in all, the genital pores are on the ventral side, in a pair, just outside the jaw plates, but they are seldom visible in preserved specimens. The madreporic plate is usually rather large and placed close to the central area of the disk; sometimes there are two or more, rarely five.
The papular pores are dorsal and form several rows on the median and lateral parts of the rays, and sometimes on adjacent parts of the disk, but are absent from the more or less extended
08 NATURAL HISTORY BULLETIN
interradial areas, where the plates are more closely imbricated and different in shape.
ASTERINIDES Verrill. Type, A. foliwm (Ltk.)
Asterina (pars) Litken; Sladen; Perrier, ete. Asterinides Verrill, Revision Asterine, pp. 477, 482, 1913. N. Pacific Starfishes, p. 263, 1914a.
Margins of disk and rays thin, subacute; rays short, depressed.
Interactinal or ventral plates in regular oblique rows, each with a fan-shaped group of two to eight small spines, usually webbed.
Dorsal plates of papular areas thin, nearly all of one kind, the exposed part usually roundish, elliptical, or shield-shaped, wholly or partly spinulose. Principal dorsal plates are all closely imbricated. Small interpolated plates few and mostly solitary. Adambulacral spines form a fan or comb within the furrow-edge, and another fan on the outer surface; ventral plates and interspaces are not covered by a granulated dermis. No pedicellariz occur on the dorsal plates nor on the intervening dermis.
This genus differs from typical Asterina, type, A. guibbosa,"* chiefly in lacking the characteristic pedicellarie of that genus. Its dorsal plates are also thinner and more scale-like, more close- ly imbricated, and often partly naked and areolated.
ASTERINIDES FOLIUM (Liitk.) Verrill.
Asteriscus folium Liitken, Vidensk. Nat. Foren., Kjobenhavn, p. 60, 1859. Asterina folium A. Agassiz, North Am. Starfishes, p. 106, pl. xiv, figs. 7-9, 1877. Sladen, op. cit., 1889, p. 393.
13 Asterina gibbosa (Pennant). This, which is the type of the genus and family, is from Southern Europe and the Mediterranean Sea.
It has the following characters: Dorsal plates, in the papular areas are thick, subequal, imbricated; their exposed proximal ends are rounded, con- vex, with the margin obtuse and bearing a cluster, usually of four to six small, rather stout, divergent, blunt spinules, and more or less numerous two-bladed pedicellariz, both on the plates and between them, nearly as thick as the spinules; naked portion of the plates is finely areolated. The median plates of the rays are scarcely larger than others; usually there is a band of four alternating rows on the median area, with no special median row, unless in the young. The plates of the median radial areas are shield- shaped, with a pair of papular pores on the proximal edge, one either side
WEST INDIAN STARFISHES og
Asterina minuta Gray, (non Gmel., nec Brug.14) Ann. Mag. Nat. Hist., p. 289, 1840; Synopsis Starfish, p. 16, 1866. Perrier, Arch. Zod]. Exper. et Gen., v, p. 229, 1876.
Asteriscus foltum Verrill, Trans. Conn. Acad. Sci., i. p. 74, 1867.
Asterina folium Verrill, Trans. Conn. Acad. Sciences, vol. xii, p. 281, pl. xxxiv, ¢, figs. 3, a, b, 1907.
Asterinides foltum Verrill, Revision Asterinine, p. 479, figs. 1913a; Star fishes N. Pacific, p. 263, 1914a.
Plate i; figure 5. Plate xi; figure 4. Plate xxviii; figure 2.
The rays are usually five; sometimes six or four. The dorsal plates are usually nearly naked, often bearing only a single or double range of small and short spinules on the most elevated portion. Dorsal radial plates are shield-shaped, notched for the isolated papular pores. Papular pores form six simple radial rows. They are absent from the interradial areas.
Plates of the interradial areas are closely imbricated, not lobed ; their exposed part is more or less rhombic.
The interactinal plates are closely united; each bears a fan of usually three or four, sometimes five on the larger ones, of slen- der webbed spinules.
Adambulacral plates have, in the furrow-comb, about four slender spines, and three or four in the actinal cluster. The madreporite is near the central area of the disk. There are no pedicellariz.
A specimen before me, from Curacoa, has the radii 6™ and of the median angle, in an emargination of the edge. Small intermediate plates are few and isolated.
The papular pores form about fourteen to eighteen short rows, extending nearly to the margins and leaving only small interradial areas without pores.
Adambulacral furrow-spines slender, in regular combs of three to five; those on the outer surface are much stouter, mostly two to a plate. In- teractinal plates closely united, mostly with two acute spinules to a plate.
Marginals very small, those of the two rows subequal, convex, finely spinulose.
The pedicellarie have the basal piece swollen, ovate, larger than the tapered acute blades. They stand at all angles on dry specimens, but were probably erect in life.
14The Asterias minuta of Gmelin, 1788, and of Brugiéres, 1792, is without much doubt the same as A. exigua Lam., but certainly not the present species.
60 NATURAL HISTORY BULLETIN
11™™"; ratio, 1:1.8. It is the same one from which figures of the details of structure have been made.
The rays are convex; ends obtuse. All the plates of the upper side were naked, as received, dried. Probably they had some spinules originally. The plates are minutely punctulate, and closely imbricated.
Two alternating rows of plates, slightly larger than those ad- jacent, take the place of a median radial row. They are angu- larly shield-shaped. The adcentral edge is slightly raised and emarginate to accommodate the single papula standing there. No small intermediate plates are now left.
The plates in the rows of the sides of the rays, where there are papular pores, are similar to the median ones, but slightly smaller and mostly notched for the papula. They are somewhat oblique or one-sided.
The papule are in six rows on the rays; others are on the disk. The central dorsal pore is easily seen. The madreporic plate has a small convex, round, grooved top, but the basal part is considerably larger, with lobed margin, and has no gyri.
The adambulacral plates have four very slender graded spines in the marginal comb, and a close group of two, or sometimes three larger ones, on the actinal surface. The interactinal plates have mostly two or three small spinules. The apical peroral spines are large and acute. Curacoa, No. 7469 (Nat. Mus. eoll.).
The color, in life, seems to be quite variable. At Bermuda, where I found it common, adhering to the under side of large flat masses of limestone, it was always of a pale, but distinct, blue color.
This small species is the most common shallow-water member of the family, from the Florida Keys and reefs to the Lesser Antilles and to the Bahamas and Bermudas. Curacoa (Yale Mus. specimen described). Perrier (1876) recorded ten speci- mens from Guadeloupe, among which one had four and another six rays. The Bahama Expedition took it at the Tortugas (one specimen was six-rayed), Jamaica (Clark).
Bermuda, common under large stones. Old Providence (Yale Mus.).
The only closely related species is A. modesta Verrill, from
WEST INDIAN STARFISHES 61
Panama. For the sake of comparison a revised description of the latter, from the type, is given here.
Asterinides modesta Verrill.
Asterina modesta Verrill, Trans, Conn. Acad. Sci., vol. i, p. 277, 1867. Asterinides modesta Verrill, op. cit., p. 482, 1813.
Plate xxvii; figures 1—2.
Rays five, short. Dorsal radial plates are thin and closely imbricated, with a convex outer surface; beneath concave, over the isolated papular pore. A solitary, small, rounded plate stands adcentrally to the papular pore; it bears a tuft of small, slender spinules like those of the larger plates. The latter have, near their higher margin, a row, usually single, of small, slender, acute, divergent, thorny spinules. Pedicellariz are lacking.
The interactinal plates are few, convex, minutely punctulated. Most of them bear a comb or fan of two or three slender spinules.
Adambulacral piates have a furrow-comb of about three slen- der, tapered spinules.
Panama and Pearl Islands at low tide. (Yale Museum.)
Genus ENopLopatiria Verrill. Type, E marginata (Hupé).
Enoplopatiria Verrill, Revision Asterinine, p. 480, 1913a. Starfishes N. Pacific, p. 263, 1914a.
Dorsal plates are imbricated; a median radial row is distinct in the young. The principal plates of the papular areas have the exposed portion usually elliptical, curved, or crescent-shaped, prominent and spinulose; small interpolated plates are present in small groups or solitary. Ventral plates have a fan of two to five spinules; adambulacrals have a furrow-comb of three to five or more spines. Pedicellarize are present on the dorsal plates, especially on the small interpolated plates.
In this group the papular area is plainly differentiated on the rays; it consists, in the adults, of about four or five median and submedian rows of larger and broader, four-lobed or five-lobed plates, curved and notched proximally for the papular pores, and of two or three rows of narrower, obliquely placed, lobed plates, having an angular proximal lobe, and notched proximally over the papular. pore.
62 NATURAL HISTORY BULLETIN
Between all the larger plates are a few small rounded plates, either solitary or in small groups, and often bearing a single bivalved pedicellaria.
On the dorsal interradial areas, beyond the papule, the plates are smaller, closely imbricated, subtriangular or pelecoidal, thick- ened, usually with a solitary small ossicle between their angles.
ENOPLOPATIRIA MARGINATA (Hupé) Verrill.
Asteriscus marginatus Hupé, Voyag. Castelneau, Zool., iii, p. 100, 1857. Perrier, Pedicell., p. 97, 1869.
Asteriscus braziliensis Liitken, op. cit., 1859, p. 57.
Asteriscus stellifer Mobius, Hamburgur Abhand. Geb. Natur., iv, 1859. Verrill, Trans. Conn. Acad., i, p. 343, 1867.
Asteriscus minutus (pars) M. and Tr., System Aster., p. 41, 1842. (non L., non Gmel., nec Gray, etc.)
Enoplopatiria marginata Verrill, Revis, Asterinine, p. 480, 1813; Star- fishes N. Pacific, p. 263, 1914a.
Plate vii; figure 2. Brazil.
Depressed, stellate, with short, rather obtuse rays, and rather evenly incurved interradial margins. Becomes 65™™ or more in diameter. Larger dorsal plates rather conspicuous with thick- ened exposed portions, mostly broad crescent shaped. The larger ones bear a curved single or double row of about six to twelve spinules, not covering the whole surface, and usually several forked pedicellariz, with a large rounded basal piece, nearly as thick as the spinules. A similar pedicellaria is usually borne on each of the small interpolated plates.
Small specimens, up to 70™™ in diameter, usually have a dis- tinet median dorsal row of subrhombic or shield-shaped plates, flanked on each side by a row of obliquely placed plates of sim- ilar size, but less symmetrical. Larger specimens usually show the median row of plates proximally, but on the distal third or half of the ray the dorsal rows of plates become confused and crowded irregularly.
The plates of the dorsal interradial areas are regularly im- bricated in alternating rows, becoming thicker proximally, and often have a small central boss. They bear a group of small divergent spinules.
The ventral or interactinal plates are regularly arranged in
WEST INDIAN STARFISHES 63
oblique rows, thickened or convex, with a finely areolated sur- face, and bearing a central fan of two to five or more spinules. The smaller specimens usually have only two or three spinules; larger ones may have four or five on the proximal ones and two on the distal ones.
The proximal plates are so arranged that in the dry specimens a distinct pore, similar to the dorsal papular pores in size, is situated between their angles, but whether occupied by a papula cannot be ascertained from the dry specimens. The furrow- series of spines form a regular graded, webbed comb of about four or five on each plate, the central spines longer. On the actinal end of the plate there are usually two to four webbed spines, in a fan, the number increasing with the age.
West Indies; Colon; south to the Abrolhos Reefs, and Rio de Janeiro, Brazil. West Africa; Senegal (Perrier); Canary Islands.
Perrier records a six-rayed example from Brazil.
The Yale Museum has specimens from Rio de Janeiro (coll. C. F. Hartt, No. 5205) and other localities.
Perrier and several others regard the West African species (EL. stellufera {Mob.]) as identical with that of the American coast. I have seen no African specimens and cannot express any personal opinion as to their identity.
The following species, from Panama, is very closely related to the above.
Enoplopatiria siderea Verrill.
Enoplopatiria siderea Verrill, Revision Asterininz, p. 480, 1913; Starfishes N. Pacific, p. 365, pl. cix, fig 3, 1914a.
Plate xxvii; figures 3—3a. Type.
This is a rather large species. The type (dry) has the radii 24™ and 45™™; ratio, 1:1.83.
It differs from the preceding in having the principal papular plates larger, thicker, more deeply lobed, and more finely areo- lated ; in having the distal, dorsal interradial plates more round- ed, and provided with very regular stellate rosettes of five to seven slender, divergent, interlocking spinules, and often one
64 NATURAL HISTORY BULLETIN
central spine. Combs of ventral plates have two to four slender, tapered, subequal, webbed spines, but mostly three.
Furrow-spines are in convex combs of four or five slender, webbed spines. Ventral plates are covered with a tough dermis. Pedicellariz are abundant, two-valved. Marginal plates small, convex, regular, about equal, regularly paired; lower ones are the more prominent.
Panama. One dry specimen, presented by Capt. Dow (Yale Univ. Mus.).
Genus ASTERINOPSIS Verrill. Type, A. penicillaris (Lam.).
Asterinopsis Verrill, Revision Asterinine, p. 480, 1913.
This genus is peculiar in having a fascicle of slender spinules on the ventral plates, instead of a fan-shaped group, which is usual in the family. Pedicellariz are lacking. The dorsal and marginal plates also bear fascicles of slender spinules. Adam- bulacral furrow-spines are in regular combs. The principal dorsal plates are lobed or cross-shaped.
In addition to the type, which is Indo-Pacific, this genus in- cludes the following two species.
ASTERINOPSIS PILOSA (Perrier) Verrill.
Asterina pilosa Perrier, Bull. Mus. Comp. Zool., vol. ix, p. 16, 1881; Nouv. Arch. Mus., vi, p. 219, pl. iii, fig. 8, 1884. Asterinopsis pilosa Verrill, Revis. Asterinine, p. 480, 1913. This species was described from a single six-rayed specimen ; radi (67> and 102". ; It is evidently very closely related to the A. lymani and may not be distinct from it. Asin the latter the ventral and dorsal plates bear fascicles of elongated, slender spinules, appearing silky. They agree in having six rows of papule; four slender spines in the furrow-combs; and in most other respects. Off Dominica I., 118 fathoms, Blake Expedition.
ASTERINOPSIS LYMANI (Perrier) Verrill.
Asterinopsis lymant Verrill, op. cit., 1913, p. 480. Asterina lymani Perrier, Bull. Mus. Comp. Zo6l., vol. ix, p. 15, 1881; Nouv. Arch. du Mus., vol vi, p. 219, 1884.
WEST INDIAN STARFISHES 65
The following notes were made by me several years ago, on the type specimen preserved in the Museum of Comparative Zoology.
Rays unusually long for this group. Principal dorsal plates are small, lobate or cross-shaped, separated by numerous rather large papular pores; they bear close pencils of rather long, slen- der spinules, ten or twelve or more in each pencil. The spinules of the plates in the interradial areas are more slender and di- vergent.
On the actinal side the outlines of the interactinal plates are not distinguishable; each plate bears a group or pencil of four to six very slender, long, acute spinules, longer than those of the dorsal plates. Marginal plates very prominent, but not very large; not flat. Each bears a terminal close cluster of long, slender spinules, like those of the interactinal plates. The ad- ambulacral plates bear a comb of about four or five slender, acute, regularly placed spinules in the furrow-series, and a group of three to six similar spinules on the outer surface.
The type was dredged by the Blake Expedition, off Barbados.
Genus STEGNASTER Sladen. Type, S. WESSELI Per.
Stegnaster Sladen, Voy. Chall., xxx, p. xxxiv, 376, 1889. Fisher, op. cit., p. 254, 1911b. Verrill, Revision of Asterinine, p. 481, May, 1913.
Form depressed, pentagonal or stellate, with very short rays; margin thin.
Adambulacral spines form a continuous webbed series, the in- dividual combs of three to five being united together; no spin- ules on the outer surface. Dorsal and ventral plates imbricated and, like their interstices, covered with a finely granulated dermis, which also forms a web between the minute spinules forming a feeble fan, on the interactinal plates. No dorsal spin- ules. Papular pores form six or more rows on the rays. In- ternal dorso-ventral columns are present near the margins. Form depressed, pentagonal, with thin margins. A few small ossicles occur between the dorsal plates; usually solitary. No pedicel- larie have been observed.
Besides the typical species, this genus seems to include S. i- flatus Hutton, of New Zealand.
66 NATURAL HISTORY BULLETIN
STEGNASTER WESSELI (Perrier) Sladen.
Asterina wesseli Perrier, Arch. Zool. Exper. et Gen., vol. v, p. 231, 1876; Etoiles de Mer., p. 220, 1884.
Stegnaster wesseli Sladen, Voy. Chall., xxx, p. 778, 1889. Verrill, op. cit., p. 481, 1913. ;
Plate ii; figures 3, 3a.
The form is pentagonal with emarginate sides, or substellate, with very short, wide rays. Radii of one of the larger speci- mens, 1677 and 20™™; ratio, 1:1.5. No. 758, Yale. 3
Central part of dorsal surface is convex. The dorsal surface is everywhere covered with a granulated dermis, both on and be- tween the plates. The granules are minute, but sharp and rough. The larger plates of the papular areas are convex, curved and deeply notched for the papular pores; in front of the notch is a small ossicle, bordering the pore proximally.
The interactinal plates are rather thick, closely imbricated, and covered, together with their interspaces, by a sharply and densely granulated dermis, which also invests the bases of the row of minute spinules; it also forms a continuous web nearly to the tips of the marginal adambulacral spines, uniting them in an uninterrupted series to the ends of the rays; it also covers the jaws and peroral spines, the granules becoming there higher, like small acute spinules. Many of the interactinal plates bear a small transverse group or rudimentary fan of two to four very small, sharp spinelets, which are much obscured by the sur- rounding granulated skin. The granules are pointed, rough, and crowded.
The furrow-spines are sub-equal, slender, obtuse, mostly two or three to a plate, standing in a nearly straight row.
Dorso-ventral pillars or trabecule are well developed, rather stout, often curved. Jaw-spines form a continuous series with about six or seven on each side, grading into and united with the adambulacral row; apical ones stouter. No epioral spines seen. A pair of small pores, probably genital, are situated in each interactinal area, one on either side of the convex outer end of the jaw-plates.
The central part of the dorsal side of the disk is surrounded by five larger and more prominent lunate plates, which are the
WEST INDIAN STARFISHES 67
basal radials or carinals. The madreporite, which is triangular, small, and depressed, is bordered externally by two smaller lunate plates, just outside the row of basal carinals.
The interradial areas, destitute of papule, are relatively ex- tensive, running up in a narrow area, nearly to the central pentagon. The plates on the interradial areas are rhombic, closely united, and a little convex.
The median line of each ray is occupied by a row of small elliptical plates, elongated radially, without papule. The pap- ular pores are large, round, and form six radial rows, where most developed; a few are situated on the central area of the disk. Superomarginal plates are transversely oblong, sometimes con- stricted across the middle, or double, and about twice as large as the roundish infero-marginals; the latter, however, project a little more and thus form the edge of the margin. Both series are covered with sharp granules, like the dorsals. No pedicel- lariz were found.
Variations.
A somewhat smaller alcoholic specimen (diameter 28™™) ap- pears somewhat different. The principal dorsal plates are more distinct and appear more convex and more regularly arranged. The outlines of many of the plates can be seen under the gran- ulated dermis. The upper marginal plates show their outlines beneath the granules. When the granules are cleaned off, they are oblong with rounded angles, with the longer axis transverse to the disk-margins. They are of the same width and number as the lower plates, and about twice as long, vertically. Their width is about the same as that of the adjacent abactinal plates, and they are covered with the same form of granules. Ventral interradial plates bear somewhat prominent flattened fans of minute spinules, webbed together at their base and showing three to seven small points at the outer edge. Jaw-plates are raised in the middle in the form of low verruce, covered with uniform and very small spinules. The peroral spines are slender, rather longer than the adambulacral spines, and closely webbed to- gether. The adambulacral spines are free only at the tips, the web being continuous along the whole length of the groove.
Off Florida; Bahamas; Jamica; West Indies generally; Colon (coll. W. F. Bradley, Yale Mus., No. 758).
68 NATURAL HISTORY BULLETIN
Family Poranup& Perrier.
Gymnasteriade (pars) Perrier, op. cit., 1884, pp. 165, 229, Sladen, 1889, . 3005. aoe ane Perrier, op. cit., 1894, pp. 163, 227. Asteropide (pars) Fisher, Smithsonian Mise. Coll., vol. 52, p. 90, 1908; op. cit. 1911b, p. 247. Verrill (pars), op. cit., 1914a, p. 304.
Disk usually large; rays short and broad. Dorsal ossicles sometimes in regular radial rows, on subimbricated, more often irregular, sometimes partly abortive, either covered with a thick, smooth or granulated dermis, or spinulose.
Marginal plates sometimes prominent; often small; usually oblique or overlapping; the lower ones projecting, and often with a small group of spinules on the outer edge, or with a single spine. :
Interactinal plates often numerous; usually in regular oblique rows or chevrons or else in rows running to marginal plates. Papule in dorsal radial areas; sometimes interomarginal. Pedicellariz lacking.
Adambulacral spines may form a simple row, transverse to the groove, or there may be also an inner comb of two, three or more, often partly rubbed.
The structure of some of the genera is very much like that of the Asterinide, with which the group might, perhaps, be united as a subfamily, without doing violence to their natural affinities.
The genus Poraniella, recently described, is more nearly inter- mediate than any other.
It is intended here to separate those genera of Asteropide closely related to the type, and especially those having bivalve or trivalved pedicellariz, as a distinct family of the Valvulosa. Por- aniide appear to belong rather with the Spinulosa, next to As- terinide.
Genus PoRANIELLA Verrill.
Poraniella Verrill, Annals and Mag. Nat. History, vol. xiv, p. 19, July, 1914c.
Small starfishes with short rays. Dorsal surface covered with flat imbricated plates, in regular radial rows on the rays, not concealed by dermis, and bearing few spaced marginal spin- ules; medial radial row is distinct. Papule are isolated and placed under the proximal edge of the radial plates.
WEST INDIAN STARFISHES 69
Superomarginal plates well developed, larger than dorsals, naked, or with few spaced submarginal spinules.
Inferomarginal plates are about as large as the upper ones, more prominent and form the thin margin; their outer edge has a comb of slender spines; their upper surface has a second par- allel row of small spines, no papule pores between the rows.
Interactinal plates are imbricated in regular chevrous, each of which, except the first, has an impaired median plate. Most of these plates have two or three spinules standing side by side, otherwise the plates are naked.
Adambulacral plates have a horizontal regular inner comb of about three small spines discontinuous with the spines on the actinal face, which stand in a transverse line, usually of two or three. This genus resembles Asterina and Astervndes pretty closely, not only in appearance, but also in the arrangement of the imbricated dorsal plates and their spinules, and the position of the papule. The plates are larger, however, and do not be- come so numerous and small toward the margins. The mar- ginal plates are much larger, and thicker, and therefore the margin is not so thin and acute.
The interactinal plates and their spinules are much as in As- terina, though larger and much fewer. The presence of a regular furrow-comb of adambulacral spines is another similar feature, though the spines on the actinal face stand in a transverse row, not as they do in Asterina.
This genus, therefore, is a sort of connecting link between Asterinide and Poraniide. Its affinities seem, on the whole, to be more with the latter, though the thick dermis found in most genera of the latter is lacking. The genus most nearly related is, perhaps, Lasiaster™ of Sladen, if this be distinct from the earlier genus Rhegaster Sladen, which is denied by some.
15 Lasiaster Sladen, op. cit., p. 371, 1889. All the plates, above and below, and both marginals covered with thick dermis and all have a compact cluster of numerous small spinules, but they are absent from the sutural lines. Superomarginal plates are rather large and their outlines are visible through the dermis. Inferomarginals not larger, but more prominent, finely spinulated on both sides and with a row of spinules on the outer end.
Adambulacrals have two furrow spines, nearly or quite side by side prox- imally and becoming oblique distally; actinal face has three spines in a
70 NATURAL HISTORY BULLETIN
Poramella also closely resembles, in general appearance, Mar- ginaster of Perrier. The type of the latter (so designated by Sladen, 1889) is M. pectinatus Per. (1881.) This has a different structure of the dorsal skeleton and very different adambulacral spines, which do not form a furrow-comb. In fact, it is covered with a thick dermis and is, without much doubt, the young of a large species of Porania or some closely similar genus.**
The Marginaster echinulatus Perrier, is nearer Poramella in structure, for it has furrow-spines in a comb of two or three and’ a transverse row on the actinal face as in the latter.
PORANIELLA REGULARIS Verrill.
Poraniella regularis Verrill, Annals and Mag. Nat. History, vol. xiv, p. 19, July, 1914c.
Plate vii; figures 1—la. Plate xv; figures 5—5b. Details.
A small depressed stellate starfish, with a wide and somewhat convex disk and short, rapidly tapered, subacute rays, with rath- er thin fringed margins. Radii of the type, 6™™ and 12™"; ratio,
transverse row. Interactinal plates are in regular chevrons, each with a com- pact cluster of small spinules.
Type, L. villosus Sladen (1889), said by Sussbach and Breckner (op. cit., 1910, p. 219), to be the same species as L. hispidus (Sars, 1871); P. rosea Dan. and Kor. (1884) ;Rhegaster murrayi Sladen (1883); Poraniomorpha spinulosa Verrill (1879); R. borealis Verrill (1878). See Plate x, figures 3, 3d.
If this be correct, the name should be Rhegaster hispidus (Sars), for Rhegaster is six years earlier than Lasiaster. Our two New England Species appear to me to be distinct and should be called Rhegaster spin- ulosus and R&. borealis. See Verrill, 1914c, pp. 17, 18, pl. i, figs. la, 10, 2.
The latter is very closely related to R. murrayi, the type of Rhegaster. It differs in having the interactinal spinules longer and in special clusters, but the whole dorsal and marginal surfaces are densely clothed with uni- form erect miliary spinules, entirely concealing the plates and sutures, as in Rhegaster. Whether either of-our species is the same as Rhegaster hispidus (Sars) I am not able to say. That is the earliest name of any species; borealis is next in order (1878.) In Lasiaster the spinules do not cover the sutures, and therefore the outlines of the plates are more or less visible, and the superomarginal plates are larger and thicker. See also 1914c¢, in Annals and Magazines Nat. History, vol. xiv, pp. 17, 18.
16 I have before me equally small specimens of Porania wmsignis V. and Rhegaster spinulosus V., which agree very closely with Marginaster in de- tails of structure. |
WEST INDIAN STARFISHES fo
1:2. Marginal plates are eight in each row. The two series are not quite opposite.
The dorsal surface is covered with rather thin and relatively large imbricated plates, overlapping adcentrally, and adradially, forming rows subparallel with the median radial row. Two rows each side of the median radial row have plates larger than the median, flat, thin, imbricated toward the median, with the free edge curvilinear and slightly notched or crenulate, and bearing three to five minute, acute, marginal, appressed spinules. Rest of the surface without spinules and not obscured by dermis. Isolated small papule are situated adcentrally to these plates, one or two under the free edge of each, except distally. These are followed toward the interradial margin by two short curved rows of similar, but smaller, plates without papule between them.
The median radial row of plates is narrow but distinct. Close to the tip of the ray the plates are small and regular, angular, and he flat, with no pores between. Soon they become notched and turned up with two papular pores under the free edge. Proximally they stand up nearly edgewise and become smaller and are depressed below the level of the adjacent rows of plates. They bear two or three minute acute marginal spinules. The basal radial plates are much larger, but stand up obliquely, showing only one edge and part of the side. Each bears five or Six minute spaced spinules on the proximal margin, and three or four scattered on the surface.
At the base of the interradial areas are two similar closely joined or united plates, of larger size, with a notch between them, from which a narrow naked line runs down to the interradial margin. The small madreporite lies in, or partly above, the notch between two of these plates, which with the basal radials, form a raised rim, surrounding a central group of smaller plates, mostly rounded, and having few minute marginal spinules.
The superomarginal plates, which in the larger specimens number nine or ten, are rather larger and thicker than the radial plates, and a little convex, with curved outer and inner margins. Each one bears two (rarely one) small submarginal spinules on the outer side.
The inferomarginal plates are similar in size and number, but more prominent, with the edge rounded, forming a rather thin
12 NATURAL HISTORY BULLETIN
scalloped margin to the rays. Each bears two rows of spinules; a short row of about three on the upper surface, which are acute and divergent; and a marginal row, usually of four slender half- webbed spines, the middle ones largest, so that the outline of the group is curved. On the under side they are naked, trans- versely oblong, and the surface is covered with microscopic, raised dots.
The interactinal plates form three regular chevrons, with a few additional small plates in the interradial angle. (In smaller examples there are but two chevrons). In the first series there may be twelve or more plates, all paired. In the second and third chevrons there is an impaired interradial plate. All these plates have two or three small acute spinules, near the aboral edge, standing side by side, and subappressed. The plates are somewhat imbricated, more or less ovate, or subangular with rounded lateral edges, variable in size. The surface is covered with raised microscopic dots and covered only with a thin dermis that does not conceal them.
The adambulacral plates are small; each has a regular furrow eomb of three slender webbed spines, directed horizontally. On the actinal face, and separated from the inner ones by a naked groove, are two or three, usually two, larger divergent spines, standing transversely to the groove.
The adoral and epioral spines are similar to the ordinary adam- bulacrals, but rather larger and longer; the perorals are some- what stouter and blunt.
This was dredged by the Albatross, off West Florida, at sta- tions 2334 (No. 10,190, Nat. Mus., type) 67 fathoms; 2324, No. 10,129, 23 fathoms; 2338, No. 10,179, 169 fathoms; 2323, No. 18,484, in 163 fathoms. Also Bahamas, station 2649, No. 18,474, in 36 fathoms.
Young. The smallest specimen seen (No. 18,474) has the radii 4mm and 6.5™™. Marginal plates seven in each row.
The dorsal plates are arranged nearly as in the type, but the specimen is preserved in better shape and all parts are neatly regular.
The plates are not only much smaller, but lie flatter and their outlines are less distinct because the dermis is a little thicker, most of the plates bear one small sharp spinule, situated on the
WEST INDIAN STARFISHES 73
inner edge, just over the papular pore, or between them when there are two pores.
The median radial row of plates is less compressed proximally than in the type, and less depressed, being nearly on the same level as those adjacent, but the plates have the same form, and distally near the ends of the rays they become flat, polygonal, and have no pores between them, as in the type; most of them bear a single median sharp spinule on the inner edge. There are three radial rows of plates on each side of a ray, the outer one of about three plates. Papule form two rows on each side of a ray, with a few in a third row. The five primary radial and five interradial plates are larger, thicker and more distinct than the rest, but are not turned up, as in the type, and they bear very few minute spinules. They surround a central area of small plates covered with dermis and bearing isolated spinules. A minute dorsal pore is present, surrounded by microscopic spinules; marginal plates are essentially as in the type, but all parts are much smaller, and the upper ones mostly bear but one minute spinule. The comb of marginal spinules and the in- feromarginal plates are largely webbed and the spines are very slender.
Interactinal plates form two chevrons with a few of a third, most of them have two small webbed spines on the aboral end.
Adambulacral spines are like those of the type, but much smaller,
Other specimens are intermediate in size between the above and the type, and have intermediate conditions, as to plates and spinules, but all agree so closely that their identification is easy.
PORANIELLA ECHINULATA (Perrier) Verrill, 1914c.
Marginaster echinulatus Perrier, Bull. Mus. Comp. Zool., ix p. 17, 1881; Etoiles de Mer, p. 230, pl. i, figs. 6, 7; Exped. Trav. et Talism., p. 169, 1894, (redescribed). Sladen, Voy. Chall., xxx, 366, 1889.
Poraniella echinulata Verrill, Annals and Mag. Nat. History, vol. xiv, p. 20,
_ duly, 1914e.
Radii of the type are 3™™ and 5™; ratio 1:1.66. The form is somewhat stellate. According to Perrier (1894) the type has the following characters:
On the dorsal side the central plate is surrounded by ten plates, five larger basals (interradials), and five radials, the last two
74 NATURAL HISTORY BULLETIN
sets are in contact. The radials are pentagonal. In this cen- tral area are no papular pores, but a large one stands next to the suture between the radials and basals.
The carinal row, beyond the basal, has five pentagonal plates. The proximal lobe overlaps the edge of the preceding plate, and has a papular pore each side of it. The other dorsal plates, sim- ilar in form, are in oblique short rows that run from the ecarinal to the marginal plates; the first row has three plates; the second and third two plates each.
There are isolated papular pores each side of the plates of he first row, forming four radial rows of pores. There are also two rows of small interradial plates.
The dorsal plates mostly bear from one to three small spines. No pedicellariz were found.
The superomarginal plates are four to each side of a ray; they are longer than wide, destitute of a marginal row of spinules, but bear small scattered spinules. The ocular plate is rather large, ovate, unarmed. The inferomarginals are five to each side of aray. They are longer than wide and each bears a row of five to seven marginal spinules, which are short and divergent, forming a marginal fringe. The distal plates are very small.
The few interactinal plates form two chevrons. The first has no impaired interradial, each of its halves has five plates ad- joining the adambulacrals and larger than the latter, so that one corresponds to two adambulacrals.
The second chevron has an impaired interradial plate, and each half has two plates; a second odd interradial lies opposite the median suture of the inferomarginals. These plates are not imbricated and each usually bears two small central spinules.
The adambulacral plates project into the groove and the inner edge bears a divergent row of three to five small spinules; on the actinal side they bear a somewhat oblique transverse row of two or three somewhat larger spinules.
The jaws are elongated, each half bears a large terminal or peroral spine and a row of four smaller graded adorals; also one enlarged epioral, inclined toward the mouth.
Blake Station 278, in 69 fathoms, off Barbados (3 specimens. )
This species seems to be nearest Poraniella, but the cotypes, which I have examined, are all evidently very young. Thus the
- EIN, TD ST i a
WEST INDIAN STARFISHES (fs)
adult might be a Porania or some other related genus. I have seen only one type. It differs from the other similar small or young forms, in having more spines in the furrow-comb, as in Poramnella.
From P. regularis it differs especially in the form and ar- rangement of the dorsal plates. In the latter they are more numerous and show no tendency to form oblique rows running from the radials to the marginals, nor are the radial plates pent- agonal, for their exposed edges are meniscoid or pelecoid. For the present, or until larger specimens can be found, I prefer to place this species under Poraniella, on account of its having the same arrangement of adambulacral spines and interactinal plates and spinules.
Genus MarcGInaster Perrier. (Type. M. pectinatus).
Marginaster Perrier, Bull. Mus. Comp. Zoél., ix, p. 16, 1881; Etoiles de Mer, p. 229, 1884. Sladen, Voy. Chall., xxx, p. 364, 1889. Perrier, Exp. Trav. et Talism., p. 164, 1874.
Form pentagonal or stellate with short rays. Disk and rays
a little convex, edges thin, bordered by a fringe of marginal
spinules. Dorsal plates well developed, flatish, imbricated, cov-
ered with a thin integument, which often partially conceals the plates in alcoholic specimens; median radial row distinct. The larger plates usually bear few small spinules, often only one or two to a plate. Papule few isolated between the plates. Upper and lower marginal plates distinct, about equal in number ; the lower ones are larger and more prominent, and bear
a terminal row of several spinules, forming a marginal fringe.
Upper ones have fewer and smaller spinules, in a row in the type.
Papular pores do not occur between the two rows of plates, in
the type, but are present in some of the related species.
Interactinal plates few, rather flatish, imbricated or tesselated, arranged in short rows or chevrons, usually bearing one to three small clustered spinules. One or two plates are impaired be- tween the jaws and the median interradial suture.
The adambulacral spines are small and in typical species form
a short, transverse, usually oblique row of two to five spines;
the two inner ones often stand nearly side by side. Jaw-plates
relatively large, convex, separated by a median groove, and bor-
76 NATURAL HISTORY BULLETIN
dered by a regular lateral row of several spines, and a pair of larger peroral spines.
Papular pores occur mostly singly over most of the dorsal surface of the rays and disk, and sometimes between the mar- ginal plates. In M. pectinatus, the type, they are few and confined to the rays.
When first established by Perrier this genus included two very small unlike species (M. pectinatus and M. echinulatus) from the West Indies. The former should be the type. These were both incorrectly figured. Later (op cit., 1894) Perrier rede- scribed them in detail, and added a third. They are quite unlike, as shown by the original types, which I have examined, and may represent two diverse genera, owing to the different character of the adambulacral armature, for in the second species there is an inner longitudinal fan of three webbed spinules on each plate.
As in Rhegaster they may both be the young of other genera, such as Porana, ete. I have studied a considerable series of these and allied species, but they are all small and probably mostly quite immature. Therefore, the status of the type of the genus seems to me rather uncertain.
At present, it seems best to restrict the genus to those species, which, like the type, have a simple transverse row of adambula- eral spines. Those that have an inner longtitudinal comb should be referred to Poraniella, or to Rhegaster.
MARGINASTER PECTINATUS Perrier.
Marginaster pectinatus Perrier, Bull. Mus. Comp. Zodél., ix, p. 16, 1881. Nouv. Arch. du Mus., vi, p. 229, pl. i, figs. 4, 5, 1884. Exp. Sci. Trav. et Talism., p. 167, 1894, redescribed. Sladen, Voy. Chall., xxx, p. 366, 1889. Verrill, 1914c, p. 18.
The type of this species, which is preserved in alcohol in the Mus. Comp. Zodlogy, has been examined by me.
The adambulacral plates are small and crowded; each bears an inner or furrow-spine and one or two outer ones, there being but one of these distally, while on the middle part of the groove there may be two, or sometimes three spines, forming a small simple transverse row. These spines are all very small, slender, and crowded, most so near the middle of the rays.
The interactinal areas contain about ten to twelve plates,
WEST INDIAN STARFISHES 17
which, when moist, are visible through the thin dermis that cov- ers them. A few of them bear one, two, or sometimes three small clustered spinules.
The carinal radial row of plates is rather conspicuous; the plates are larger than those adjacent, irregularly four-lobed, overlapping by their lobes. Small, isolated, short, acute spin- ules are borne on most of the dorsal plates. The superomar- ginal plates of which there are six on a side, do not reach the margin; they bear only a few granule-like small spinules, as now preserved.
The inferomarginal plates project strongly beyond the margin; those in the middle of the interradial margin are decidedly larg- er than the others. The marginal fringe of spines, that they originally bore on each series, was mostly rubbed off before I first saw the specimen, in 1896, but Perrier (1894) states that there were four or five flat, lanceolate spinules on each plate of the lower series. His figure, however (1884), represents them as slender and acute. The figure is incorrect in various other respects.
According to the later description by Perrier (1894, p. 167), the type of this species has the following characters:
Radii 4"™ and 5™". The adambulacral plates bear one inner spine and another, or more often two, on the actinal side, a little larger than the inner one, divergent and placed one behind the other in a transverse row.
On the dorsal side the central, five radial, and five interradial plates are distinct. The basal radials are triangular with round- ed corners, with the wider end turned proximally. The five car- inals are imbricated and slightly cruciform, decreasing regularly distally. The ocular or terminal plate is semicircular and with- out spines.
The central plate has three spinules: the interradials six or seven; the basal radials three; the carinals two or one.
Transverse rows of smaller elliptical dorsolateral plates run from the carinals to the marginals. The first and second series have apparently, three plates; the third two plates; the fourth and fifth each have one unarmed plate. The larger of these plates bear two or sometimes three, small spines; the smaller only one spine. On the disk there are a few other plates irreg-
78 NATURAL HISTORY BULLETIN
ularly placed. The papule are large, all dorsal, and arranged in rows.
The superomarginal plates correspond with the lower ones; each bears a marginal series of three or four flat, lanceolate spinules.
The inferomarginals are five on each side of a ray. They are rectangular, broader than long; each bears a marginal row of four or five flat, lanceolate, acute, hyaline spinules, forming a marginal fringe of spines.
The jaws are triangular, of moderate size, not prominent. Each half bears an angular peroral spine, four lateral spines, and two or three epiorals.
The interactinal plates form a triangular area; three larger ones stand in an impaired median row, running from the jaws to the median interradial suture; one small plate intervenes be- tween the second interradial marginal plate and the third adambulacral, a larger plate lies between the third marginal and the fifth adambulacral. A series of three plates, the middle one alone well developed, runs from the suture between the first and second, and that of the sixth and seventh adambulacrals; the last plate lies between the sutures of the second and third marginals and that of the eighth and ninth adambulacrals. Some of these plates bear a single spinule; most are without spinules.
Perrier refers to the type specimen as ‘‘unique’’ (1884, p. 230, foot note). He states that it was accidentally irregular, which was the case in the type seen by me. The only locality given was the Blake Station 32, in 95 fathoms, N. lat. 23° 52’.
PoRANIA (?) AUSTERA Verrill.
Marginaster austerus Verrill, Revision Genera Starfishes, Trans. Conn. Acad. Sci., vol. x, p. 221, 1899. Porania ? austera Verrill, Ann. and Mag. Nat. History, vol. xiv, p. 20, 1914c.
Plate iii; figures 1—la. Plate xi; figures 6—6a. Details.
Body fiat below, swollen above, with short, thick, rapidly tap- ered acute rays. Radii of the type 11™™ and 17™;; ratio, 1:1.6; marginal spines eleven. Whole surface covered with a rather thick dermis which conceals the outlines of all dorsal plates and most of the ventral ones, except the interactinals.
WEST INDIAN STARFISHES 79
The dorsal surface is covered irregularly with small, short, blunt or clavate rough spines. These stand partly isolated, part- ly in little irregular rows of two to six. These rows run in various directions, but on the rays mostly obliquely or trans- versely to the ray, but without order. The concealed plates are irregularly lobed and reticulated, so as to leave rather large isolated papular pores, which are scattered over the whole dorsal surface, and also form a regular row between the two rows of marginal plates, where they are either single or two together. Madreporic plate, which is nearer the center than the margin, is small, oval, with fine gyri, and surrounded by small obtuse spines. Dorsal pore is small, surrounded by very small spinules.
The superomarginal plates are irregularly three-lobed or four- lobed, emarginate on both edges for papular pores. Hach one bears about three or four small erect spinules like those around them, and irregularly placed.
Inferomarginal plates are prominent, beyond the upper ones, and form the margin. They have marginal notches between them. Each bears a horizontally projecting, single, outer mar- ginal row of about five or six closely placed, shghtly divergent, and only a little unequal, oblong, blunt spines. Above these, on the upper side, there is another row, corresponding in number, but not half so long, of small, acute spinules.
The under side of these plates, as seen through the thinner dermis, is transversely oblong, with the lateral margins slightly incurved, and with a small groove and marginal ridge at the inner end; the ridge may end in a slight tubercle at one or both ends. The rest of the surface is covered with minute rounded elevations.
The interactinal plates are easily visible, and mostly without spinules. They form oblique forked rows running from the marginals to the adambulacrals. The rows are separated by spaces covered only by the dermis, but the plates in each row are in close contact or slightly imbricated.
These rows or series of plates start from an angular plate op- posite the suture between two inferomarginals. This is joined adorally by two (sometimes one) long-ovate, narrower plates, with the larger end centered; each of these is followed by two
80 NATURAL HISTORY BULLETIN
similar but smaller plates, each of which abuts against two adam- bulaerals.
The two primary interradial rows start from an impaired plate, in line with the suture between the first marginals. Its two halves diverge rapidly, in a curve, leaving a small, nearly smooth and naked, triangular area outside the jaw-plates. A few irregular small plates are here concealed under the dermis. In the type two to three of the interactinal plates bear a single small blunt central spine, none have spinules.
The adambulacral plates are concealed by the dermis, which continues upward on the bases of the outer row of spines and forms a short continuous web between them.
Each adambulacral plate bears an inner small furrow-spine, and on the proximal part of the groove, mostly two larger, a lit- tle thicker, and somewhat flattened, oblong or slightly clavate blunt spines. Where not too crowded, these stand side by side, but they often stand more or less obliquely, and there may be but one on some plates. Near the distal end of the groove, they become more slender and more oblique, due to crowding. Near the jaw-plates a few plates may have two inner spines.
The adoral and peroral spines of the jaws are similar to those of the adambulacral plates, but slightly stouter, especially the apical ones. West Indies, special locality not known. (Coll. A. EK. Verrill). Also taken by the Albatross.
This is probably immature, but I am unable to refer it to any described species.
Suborder VELATA Perrier. (See above).
Family PTERASTERIDe Perrier.
Pterasteride Perrier, op. cit., 1875; op. cit., 1884; op. cit. p. 180, 1894. Ver- rill, op. cit., 1899. Pterasteride (pars) Sladen, Voy. Challenger, vol. xxx, p. 468, 1889. Pterasteride Fisher, op. cit., 1911b, p. 343 Verrill, op. cit., 1914a, p. 266. This is one of the most peculiar groups of starfishes hitherto discovered. It shows in general characters, a remarkably high degree of specialization not found in any other group. Most of the genera and species are from the deep sea. Disk usually plump; rays 5-8, usually five, rarerly 6-9; upper
WEST INDIAN STARFISHES 81
surface covered by a supradorsal membrane supported by the tips of long, slender, divergent, often webbed, paxillary spin- ules and pierced by small concentric pores or ‘‘spiracles’’ often closed and invisible in preserved specimens and usually with a central osculum. Beneath this membrane is a ‘‘nidamental eav- ity’’ or gonocodium, traversed by the columnar pseudopaxille, and containing the papule. In this cavity the eggs are retained, and also the young, till they assume the adult form, often 8 to 10™™ in diameter. The dorsal skeleton ossicles are lobed or eruci- form, loosely reticulated.
The adambulacral spines usually form transverse webbed combs or fans (not much webbed in Hymenasterine).
Series of slender, divergent spines more or less appressed and attached to the under surface, or imbedded in it, and us- ually webbed to the adambulacral fans in Pterasterine, are al- ways present. These peculiar spines, called ‘‘actino-lateral spines’’ by Sladen. should rather be called retroambulacrals or outer adambulacrals, for they are attached to the outer end of the adambulacral plates. They are sometimes short, and do not reach the margin, except distally, on the rays, but in Pteraster and some other genera their tips usually form the marginal edge of the disk.
Between the bases of these there is usually a small slit or pore (actinal spiracle or ‘‘segmental aperture’’) furnished with a partly caleareous valve. Interactinal plates are lacking. Pedi- cellariz have not been found. Jaws have a series of adoral spines, usually webbed. Epioral spines of large size, in one to three pairs, often without webs, stand perpendicular to the jaw; those of one pair are sometimes specialized and partially hyaline. Ambulacral feet are large, in two to four rows, with large apical suckers.
The supradorsal membrane may be thin and translucent, with thin muscular fibres, or thick and muscular. In some cases it contains calcareous spicules, muscular and cartilaginous fibers in the form of a network, and often abundant mucous glands. The copious mucus is phosphorescent in some cases (Diplop- tereaster.) The intestine and anal pore are well developed. Sex- es are alike externally.
82 NATURAL HISTORY BULLETIN
Subfamily HyMENASTERIN2.
The genera Hymenaster, Cryptaster and their allies form a distinct subfamily, Hymenasterine, differing from Pterasterine in lacking webbed combs of adambulacral spines and in other characters. It is a characteristic deep-sea group. One or two species occur in the West Indies in deep water.
Subfamily PTERASTERINE. (See above).
Genus PTERASTER M. and Trosch.
Pteraster Muller and Troschel, Syst. Aster., p. 128, 1842. Type, P. militaris. Sladen, op. cit., p. 470, 1889. Fisher, op. cit., 1911b, p. 344, (Analy- tical table of all known species, pp. 368-370.) Perrier, op. cit., 1894, p. 181. Clark, op. cit., 1908, p. 286 (Analytical Table.) Verrill, op. cit., 1914a, p. 267.
Hexaster Perrier, Comptes-rendus, vol. 112, p. 1227, 1891. Type, H. obscurus
Perrier. Temnaster Verrill, Proc. U. S. Nat. Mus., vol. 17, p. 275, 1894. Type T. hexactis V. = H. obscurus P.
Disk large; rays 5 to 8; ambulacral grooves turn up distally, more or less reaching to the upper side of the rays.
Adambulacral spines form large, webbed combs. Retroambu- lacral or ‘‘actinomarginal’’ spines are closely appressed, ending in a fringe or fold. One pair of specialized epioral spines; these are often translucent distally. Supradorsal membrane furnished with a central contractile oseulum, guarded by five or more groups of webbed, projecting spines. It usually contains irregular muscular or elastic fibers for its support and sometimes calcareous deposits. The muscular bands usually do not form a regular network and are generally very inconspicuous externally. Usually two rows of ambulacral sucker-feet.
PTERASTER CARIBBZUS Perrier.
Pteraster militaris A. Agassiz, op. cit. (non Miller.) Pteraster caribbeus Perrier, op. cit., 1881, p. 13; Etoiles de Mer, p. 216, 1884. Plate vil; figure 4.
The radii of the type were 15™™ and 30™™; ratio, 1:2. One be- fore me has them 8™™ and 12™™. The adambulacral comb of very slender hyaline, webbed spines contains six or seven erect spines, increasing in length outwardly, and another, the retro-
WEST INDIAN STARFISHES 83
ambulacral, much larger and longer, recumbent, extending be- yond the border of the ray and forming a webbed margin. The dorsal parapaxille, as seen below the supradorsal membrane, are mostly cross-shaped or four-lobed at the base, two of the lobes are often longer than the others, and somewhat enlarged distally. These lobes overlap and underlap those of the adjacent plates. From the center arises a very slender and rather long column, longer than the longest basal lobes, and about three or four times higher than thick. It is enlarged or subeapitate at the summit. Each bears about eight to ten long and very slender divergent, hyaline, webbed spinules, which enter and support the supradorsal membrane.
The jaws are prominent, subconical; each half has about six very slender marginal spines, becoming longer adorally.
The specialized epioral spine is relatively large, long, flatish, wider near the tip, or sublanceolate, nearly vitreous or transluc- ent, and subhyaline in its distal half. It reflects prismatic colors in some lights.
The madreporic plate is hemispherical, raised on a stalk about as high as broad. The dorsal membrane is very thin and delicate and rises into many small conical or subacute angular elevations when dry.
This is a delicate and fragile species closely related to P. mili- taris of northern and arctic seas. Its supradorsal membrane is more delicate and its supporting parapaxille are longer and much more slender, with longer and more thread-like spinules.
This species was dredged by the Blake Exped. at four stations off Florida and among the Lesser Antilles, in 151 to 422 fathoms.
The Albatross dredged it at Station 2667, off Southern Georgia, N. lat. 30° 53’, in 273 fathoms (No. 18,415, deseribed above.) It was not obtained by the Bahama Expedition.
Order PHANEROZONA Sladen (emended).
Phanerozona (emended) Sladen, Voyage Challenger, Zool., vol. xxx, Introd. pp. xxviii, xli, 1889. Verrill, Revision, Trans. Conn. Acad., x, p. 200, 1899. Fisher, op. cit., 1911b, p. 17. Verrill, op. cit., 1914a, p. 280.
Valvate + Pasillose (as orders), Perrier, in Mem. Etoiles de Mer, An- tilles, etc., p. 154, 1884.
Valvata + Pazillosa Perrier, Exped. Trav. et Talism., pp. 22, 23, 28, 29, 1894; Resultats Camp. Scientif., Albert I, fase. xi, p. 16, 1896.
84 NATURAL HISTORY BULLETIN
This order includes starfishes varying in form from pentagonal to stellate, with long rays. They have two rows of large and thick marginal plates (rarely but one distinct row, as in Luidia), which are usually in contact and form a thick, nearly perpendic- ular margin. They are nearly always larger and thicker than the dorsal and actinal plates. Papular pores are nearly always confined to the upper surface and are usually placed singly, often in special areas. Dorsal skeleton various. The dorsal sur- face may be closely covered with angular tesselated plates, which may be naked, granulated, or covered with a smooth skin, or they may take the form of pseudopaxille, protopaxille or true paxille especially in the Paxillosa.
The various forms of paxilliform plates as distinguished in 1894, by the writer, are as follows:
True paxille are columnar or hour-glass-shaped ossicles, usu- ally with isolated, circular or lobed bases, which bear at summit a group of small spinules, of which the marginal series are dif- ferent from the rest and divergent, so as to cover the interven- ing spaces between the spines, thus forming fascioles or channels for the circulation of water to the papule. These are highly de- veloped in Astropecten and Iwidia. Those called spinopaxille are of the same general structure, but the center of the summit is occupied by a distinct spine, or by more than one. Such forms occur on some species of Luidia, Pontaster, ete.
Parapaczille are rounded or stellate ossicles, or angular plates, with a raised central portion, tabulate, truncate, or like a low cone. They may be either isolated or articulated by their bases. The summit is covered with small, differentiated spinules, like those of true paxillea. Those on the dorsal surface of Lophaster, Solaster, and Mediaster are examples. They sometimes bear a central spine.
Protopaxille are similar, but-less elevated, convex ossicles or plates, covered with round or angular granules, with the mar- ginal series differentiated and more or less covering the grooves between the plates. As in the preceding, there may be a central spine in some eases. This form occurs on Chewraster, many species of Goniasteride, ete. The transition from this last kind to simple, uniformly granulated plates is easy, when the grooves between the plates become obsolete.
WEST INDIAN STARFISHES 85
Pseudopaxilie are articulated plates with flattened, usually lobed, and often overlapping bases, which bear a group of slender, fascicled or divergent spinules on the more or less raised central or subcentral area or boss. These have no differentiated mar- ginal series of spinules. This form is seen in Henricia, ete.
In some families the plates are reticulated ossicles, smooth or bearing large spines or tubercles as in Oreasteride. In other cases a thick skin may cover and obscure the plates.
The interactinal plates may be few in slender stellate forms or many in the pentagonal forms. In the latter cases they are usually arranged in definite rows, most often subparallel with the adambulacral plates, the latest formed ones being situated next the median, interradial, marginal plates, but they also form rows running from the adambulacral plates to the mar- ginals, and often define fasciolated grooves.
They are most commonly tesselated and granulated, or else in the form of pseudopaxille or protopaxille; but they are some- times spinose. They may be more or less imbricated by their edges or lobes. |
Adambulacral plates are not compressed, rather large, usually oblong or quadrangular, and commonly bear numerous small spines, various in arrangement.
Ambulaeral plates are rather large, not compressed nor crowd- ed. Ambulacral feet are in two rows, with terminal suckers in Valvulosa; but in the Pazillosa they are large and pointed, with- out distinct suckers.
Pedicellariz are sometimes lacking, but usually present and sometimes large. They may be bivalved, sessile, and seated over a pore (foraminate), or implanted in special pits on the plates (fossate) ; or else papilliform or fasciculate, composed of two to four or more modified, convergent spinules; or pectinate, con- sisting of two comb-like groups of spinules convergent over a suture between two adjacent plates, asin Lwidiaster, or seated on a’single plate.
The sessile valvular pedicellarie with a pore between the valves are called foraminate; they may have two, or more than two, valves (bivalvular and trivalvular, ete.). In the suborder Valvulosa they are often provided with a pair of pits or fosse, into which the valves fit when widely open. The valves in these
86 NATURAL HISTORY BULLETIN
are often spatulate, spoon-shaped, or suger-tongs shaped, or bat- tledore-shaped, and usually may best be called plataleiform or spatulate. When furnished with receiving pits, they have been called excavate; but fossate seems to be a more appropriate term.
The peculiar papilliform pedicellarie of the Pazillosa, with two to four or more slender valves, apparently formed by modi- fied spinules or granules, are also found in certain of the Valvu- losa. Sometimes such pedicellarie are found associated with larger valvular pedicellarize on the same specimen, as in the genus Nymphaster, and in a few other genera.
The following is the arrangement of the suborders and fam- ilies adopted by the writer (op. cit., 1914a) :
Order PHANEROZONA.
Suborder I. VauvuLosa Verrill = Vatvata Perrier. (Sense extended. )
Family I. OpHIpIASTERIDZ Verrill, 1867 = LincK1apz Per- rier, 1875.
Family II. Vatvastreripa Fisher, 1911.
Family III. Asreropmw Fisher, 1911 = GyMNATERIDz (pars) of authors. ,
Family IV. Oreasterma Fisher = Pentacerotide Gray (re- stricted).
Family V. MIMAsTERIDa Verrill.
Family VI. GontasTerIpa Forbes (restricted), includes An- theneide Perrier (restricted). ,
Family VII. Oponrasteripz Verrill = Gnathasterine Perrier (pars).
Family VIII. Cuaztasteri& Ludwig.
Family IX. ARCHASTERIDZ Ver. (restricted to Archasier).
Suborder II. Noromyota Ludwig (as an order). Family X. BeNTHOPECTINIDZ Verrill; Benthopectinine Ver- rill, 1894. Suborder III. Paxmuosa Perrier (sense restricted).
Family XI. PoRcELLANASTERIDZ Sladen.
Family XII. GoniopectiInD& Verrill.
Family XIII. Astropectinipz@ Gray (restricted). Family XIV. Luimp# Verrill = Lu:dune Sladen.
WEST INDIAN STARFISHES 87
in the West Indian area, but some of them occur only in deep water.
Subfamilies of GONIASTERID2.
Subfamily I. GONIASTERINZE Verrill = Pentagonasterine Sladen (pars).
Subfamily II. PSEUDARCHASTERIN& Sladen.
Subfamily III. Meprasrertnz Verrill.
Subfamily IV. Necrrunz Per.; Fisher.
Subfamily V. Hippasterunz Verrill.
Subfamily VI. ANTHENEINZ Fisher (sense restricted).
Subfamily VII. LeproGoNAsTERINZ Perrier.
Subfamily VIII. CurronasterIn& Fisher.
Those subfamilies represented in the West Indian shallow- water fauna are the first, third and seventh. But the seventh is essentially a deep-sea group; only a few stray examples occur- ring in less than 150 fathoms. The third and fifth families are well represented in the deeper waters. The others are chiefly extralimital.
Suborder VALvuLosa Verrill.
Vawulate (order) Perrier, Nouv. Arch. Mus. Hist. Nat., vi, 1885. Valvata (order) Perrier, Exped. Trav. et Talism., pp. 22, 23, 28, 29, 1894. Valwata (suborder) Verrill, Revision, Trans. Conn. Acad., x, p. 200, 1899. Valvulosa Verrill, North Pacific Starfishes, p. 284, 1914a.
These starfishes are stellate or pentagonal and usually rather stiff in appearance, and are nearly always five-rayed. The dorsal plates are most commonly tesselated and granulated, or else take on the character of protopaxille or pseudopaxille. Owing to the generally close union of the plates, especially of the large marginal plates, most of these starfishes are rather rigid, showing much less flexibility than in most other groups. The Ophidias- ‘teride are exceptional in this respect, for they are mostly flexible in life.
Ambulacral feet are large, with well developed suckers.
Pedicellariz are often lacking; when present they are usually valvular, foraminate and sessile, or fossate. The valves may be either granuliform, or large and broad, or narrow and spoon-
88 NATURAL HISTORY BULLETIN
shaped, plataleiform, spatulate, or battledore-shaped, rarely pa- piulliform. There may be two to five valves. The valvular form of the pedicellarie first led to the establishment of this group.
The various kinds of dorsal plates pass into each other by various intermediate forms, so that it is impossible to draw any very strong or sharp family lines on this character alone, though the character of the plating may generally be taken as of generic value. They are more frequently polygonal and tesselated; often in the form of protopaxille or parapaxille; sometimes deeply lobed and openly reticulated ; sometimes imbricated.
Sometimes they are naked; often spinose; very rarely truly paxilliform ; sometimes covered by a thick, smooth, or granulated integument.
Both rows of marginal plates are usually well developed and form a stout margin to the disk and rays.
Interactinal plates are usually rather numerous and tesselated or imbricated. A distinct dorsal pore is usually present. Gen- ital pores are generally ten, usually all separate, situated in the dorsal interradial areas; sometimes serial on the sides of the rays.
Superambulacral plates are present in some genera, absent in others. The existence of superambulacral plates, formerly sup- posed to be characteristic of Astropectinde, occurs in several of the genera now referred to Valvulosa (viz., Medwaster, Dytas- ter, Pseudarchaster, ete.), and in other groups.
Adambulacral plates wide, with a longitudinal group of small furrow-spines and with one or several spines on the actinal side, often becoming much longer distally. Jaw-plates are large and usually convex on the actinal surface, bearing numerous oral and epioral spines.
This is a very extensive group, found in all seas.
Family OPHIDIASTERIDE Verrill.
Ophidiasteride Verrill, Trans, Conn. Acad. Sci., i, part 2, p. 344, 1867; op. cit., p. 308, 1914a.
Linckiade Perrier, Revision Stellerides, p. 117 [381], 1875.
Linckiide Sladen (pars), op. cit., 1889, p. 397. Fisher, op. cit., 19110, p. 240,
Linckiade Viguier, Squelette des Stell., p. 144, pl. ix, figs. 1-13; pl. x, figs. 1-7, 14-19 (structure), 1879.
WEST INDIAN STARFISHES 89
Form stellate, usually with five slender, flexible rays, but the number is variable in some autotomous species.
Abactinal and marginal plates usually tesselated or subim- bricated, in rows or irregular; in most cases granulose (covered with rather thick, smooth skin in Levaster), sometimes tuber- culated, rarely spinose.
Marginal plates are usually well developed, in two regular rows, but often not conspicuous and rarely so thickened as to form a rigid margin. They are similar to the dorsal plates, as to granulation.
Papule usually numerous, mostly abactinal and lateral and in groups, sometimes single; in some of the genera they occur also between or below the marginals, or between the interactinal plates.
Adambulacral plates small, covered with either granules or spinules, the margin bearing one or two rows of small spines, no more.
Pedicellarie often lacking; when present, usually bivalve, sometimes spatulate and fossate. Superambulacral plates often present.
Genus OPHIDIASTER Agassiz (restricted).
Ophidiaster (pars) L. Agassiz, Prod. Monog., 1834. Gray, op. cit., 1840, p- 283; Synopsis, 1866. Mill. and Trosch. (pars), Syst., p. 28, 1842.
Ophidiaster (emended) Liitken, op. cit., p. 163, 1864. Perrier, Revision, p- 384, 1875; op. cit., 1884, p. 221 (deser.)
Linckia (pars) Nardo, 1834. Von Martens, op. cit., p. 351, 1865.
Disk small; rays slender, subterete, closely granulated. The dorsal plates are in about three to five regular rows, leaving squarish or angular papular areas between them in regular rows. Papular areas sometimes occur, also, between the marginal rows; rarely, also, in one submarginal row. The marginal plates, which form two regular rows, are not unlike the dorsals, and closely ‘granulated. Interactinals are smaller, angular, and usually form one or two narrow rows. Adambulacral plates usually have two unequal small spines on the inner edge, and one just back of them in the second series decidedly larger, stout, but not much elongat- ed. Superambulacrals are usually present but small. Fossate pedicellariz, usually with spatulate valves, occur on most species.
90 NATURAL HISTORY BULLETIN
OPHIDIASTER GUILDINGI Gray.
Ophidiaster guildingit Gray, op. cit., 1840, p. 284; Synopsis, p. 13, 1866 (description inadequate). Perrier, Revision, p. 387, 1875 (no descrip- tion). Liitken, op. cit., 1871.
Scytaster millert Duchassaing, Ann. Rad. des Antilles, 1850 (t. Perrier from types.)
Ophidiaster flaccidus Litken, Vidensk, Meddel., p. 86, 1859 (good descrip- tion. )
Rays five, with a weak skeleton, roundish, slender. Radii of a medium sized specimen 5™™ and 60™™; ratio 1:12. Other speci- mens have shorter rays. The dorsal and lateral surfaces are cov- ered with about nine rows of plates, between which there are eight equally spaced and pretty regular rows of large papular areas, each with about five to twelve papular pores. There is no median area destitute of papule. Marginal plates are much like the dorsals. All these plates are covered with small, unequal, squam- miform granules, some of them minute.
The adambulacral plates bear two rows of small obtuse spin- ules; those of the inner or furrow series are small, equal, and crowded; those in the adjacent outer row are decidedly larger, about half as numerous, with some granules between their bases.
The color, in life, commonly varies from orange-red, with ir- regular yellow markings, to purplish brown, with blotches of dirty white (Clark).
Liitken (op. cit., 1871) recognized that his O. flaccidus is the same as O. guildingu Gray.
Locality of Gray’s types was unknown. Perrier, who exam- ined the types, states that they are identical with O. flaccidus.
St. Thomas (Liitken; Perrier). Fort Reef, Port Antonio, Jamaica (Clark.)
OPHIDIASTER FLORIDZ Perrier. Ophidiaster floriade Perrier, op. cit., 1881, p. 9; Mem. Etoiles de Mer, p. 221, -pl. iv, fig. 1.
This was described from a single young specimen (radii 5 and 33™™) from off Florida, in 2014 fathoms.
It is, perhaps, the young of O. guildingti, the common shallow water West Indian species.
According to Perrier it is formed much like a Linckia. The dorsal and marginal plates form seven regular rows of elongated
WEST INDIAN STARFISHES SE
oval plates, covered with a granulation much like that of the actinal side. The plates are rather prominent and distinct.
No papulz are found below the inferomarginal plates; between the marginal and dorsal rows there are six regular rows of pap- ular areas, each having two to four pores.
The disk has ten distinct radial and interradial plates.
Interactinal plates are smaller than the marginals, arranged in few regular rows, without papule between them, but defined by the rather large rounded granules that closely and regularly cover them.
The adambulacral plates bear two contiguous rows of spines; those of the inner marginal row are small, regular, and equal; in the outer row each group has a much larger, obtuse median spine and a small flat one each side of it.
On the interradial areas are two rows of interactinal plates that bear spines similar to those of the outer adambulacral spines but smaller; some of them bear fossate pedicellarix, having spat- ulate valves. The second row is, however, in the young speci- men, feebly developed with few plates.
Straits of Florida, in 123 feet (Perrier, Coll. Mus. Comp. Zool.). Probably 123 fathoms was intended. Off E. Florida, 277 fathoms, Albatross collection.
OPHIDIASTER ALEXANDRI Verrill, sp. nov. Plate xiii; figures 3, 3a, 3b. Details. Plate xxv; figure 2. Type.
Disk small, rays regularly tapered, subacute. Radii of the type 11™™ and 60™";; ratio, 1:5.5. The upper and lateral surfaces of the rays have five very regular and nearly equal rows of some- what prominent plates, closely united radially, but so openly and regularly united transversely as to leave nearly square pap- ular areas, which form very regular rows. They contain about eight to ten pores.
The disk is somewhat convex. It has a set of basal radial and interradial plates, and a central, larger than those of the rays, and a little swollen.
The madreporic plate is large, round, flat, covered with fine radiating gyri. The whole surface of the plates and the papular areas is covered with a fine even granulation, above and below.
92 NATURAL HISTORY BULLETIN
The interactinal rows of plates, beneath, are so closely united and densely granulated that their outlines are not visible with- out preparation. When denuded they are seen to form three rows. The three plates in a transverse series are imbricated. These plates are ovate, except the one next to the adambulacral plate, which is truncated at the suture, and corresponds in width and position to the adambulacral that it joins. Two transverse series correspond to one inferomarginal. The latter are much larger and four-lobed.
The small transversely oblong adambulacral plates are finely granulated and bear a simple row of very small and short, fiat, blunt marginal spinelets, two to a plate; and an outer row of much larger and longer, stout, tapered spines, alternating with very small acute spinelets, one of each kind on each plate.
Dredged by the Bahama Expedition at stations 56 and 57 on the Pourtales Plateau, off Florida, in 200 to 225 fathoms. Taken by the Albatross at station 2416 in 276 fathoms, coral sand and shells, N. lat. 31° 26’, off Georgia, several specimens. (Nat. Mus.; Yale Mus.)
Perrier (1884, p. 172, and 1894, p. 37) listed an undescribed species of this family as ‘‘Fromia meaxicana’’ from station 207, off Martinique, in 96 fathoms. This seems not to have ever been described, and is to be regarded as a MSS. name.
Genus Linck1a Nardo.
Linckia Nardo, Okens Isis, 1834, p. 717. Gray, op. cit., 1840, p. 284; Synopsis, p. 13, 1866. Liitken, Vidensk. Meddel., pp. 163, 164, 1864 (emended); op. cit., p. 266, 1871. Viguier, Arch. Zool. Exper. et Gen., vol. vii, p. 147, pl. x, figs. 1-6 (structure), 1878. Verrill, op. cit., 1867, p 285; op. cit., p. 309, 1914a. Perrier, op. cit., p. 399, 1875; Etoiles des Mer, p. 226, 1884. Sladen, op. cit., vol. xxx, p. 408, 1889. Fisher, 1911b, p. 242.
Ophidiaster (pars) Miller and Troschel, Syst. Ast., p. 28, 1842.
Disk small; rays long, slender, nearly terete, usually five, var- iable in number in some autotomous species. Adambulacral plates bear granule-like structures, in two or three close series, those in the second row alternately larger. Pedicellariz not ob- served. Two madreporites in autotomous species.
Interactinal plates usually form two or more close rows. Dorsal plates are irregularly arranged, not forming definite
WEST INDIAN STARFISHES 93
radial rows; closely granulated. Papule are numerous, in clusters on the dorsal side, but are lacking below the marginal plates.
LINCKIA NODOSA Perrier.
Linckia nodosa Perrier, Revision, p. 417, 1875; Etoiles des Mer, p. 226, 1884. Sladen, Voy. Chall., pp. 409, 786, 1889. Fisher, op. cit., 1906, p- 1088. Plate xiii; figures 2—2a. Plate xxix; figures la, 1b.
The type of this species had the radii 11™™ and 91™; ratio, 1 :8.3. .
The dorsal plates form about three irregular crowded rows; some of these plates, irregularly disposed, are larger and more convex than the others, or even subnodose; similar swollen plates also occur upon the disk.
The papular areas are small and well defined; they mostly have six to eight pores; fewer on the disk. Madreporic plate large, flat, with numerous fine, sinuous gyri.
The marginal plates are squarish, covered with granules a little smaller than those of the interactinal plates; between the two rows are roundish papular areas, with about twelve pores.
There are several rows of interactinal plates proximally, the inner ones short; the outer or peractinal row extends nearly or quite to the tip of the rays. These plates are all covered with small rounded granules, those near the adambulacral plates are a little the larger.
The adambulacral spines are almost granuliform; they form two rows; those of the margin of the groove are alternately larger and smaller; the larger are clavate, the smaller cylindric. Close to these are the spines of the outer row; each of these is opposite one of the smaller spines of the inner row; they are rather larger than the larger of the inner row. External to these spines is a row of granules larger than those of adjacent plates and also some intermediate small granules. There are no actinal papular pores. :
A larger specimen before me was taken by the Albatross off Pensacola, Fla. The radii are 13™™ and 125™™; ratio, 1:9.6.
The rays are well rounded above, flattened below, and taper very gradually.
94 NATURAL HISTORY BULLETIN
The dorsal surface of the rays is covered with a broad, very irregular band of large, swollen, evenly convex, unequal, round- ish plates, mostly in contact radially to a variable extent. These, where most regular, seem referable to three rows, by reason of the large, irregular papular areas between them. The plates that seem to belong to the median row average larger and more prom- inent than the others, but are unequal and seem to be crowded out of position by the growth of others.
Outside of those described, and on the sides of the ray, there is a much more regular row of dorsolateral plates, which are also roundish and strongly convex, but less so than those above. They are in serial contact, but are joined to the row above by small connective ossicles and large intervening papular areas, which form here a regular row. The papule are small and very numerous in these and the dorsal groups. Twenty to thirty or more can be counted.
All these plates, as well as those below, are covered closely with small, nearly uniform rounded granules, which also sur- round the papular pores, but there become angular, so that four or five around each pore look like small valves.
The two rows of marginal plates are very distinct and nearly equal and regular. Their plates are less convex and not so large, about five corresponding to four in the row above. They are broadly in contact serially, but the two rows are separated by a regular row of large papular areas, like those above.
Below the inferomarginals and closely joined to them and to each other there are three or four rows of squarish interactinal plates, so closely united and compactly granulated that their out- lines are concealed. Their granules gradually become coarser and angular near the adambulacral spines.
The adambulacral plates are small and concealed by granules. Each bears two unequal inner erect spines; both are flattened and obtuse. The larger is more than twice as large as the other, but only a little higher. Outside these, but close to them, and alternating with them, is a single larger, obtuse, flattened, erect spine, about twice as large as the larger inner ones, and about as broad as high. These, like the inner ones, form a regular con- tinuous row, very distinct from the adjacent granules.
These rows continue to the jaw plates with but slight change,
WEST INDIAN STARFISHES 99
but three or four of the most proximal ones of the outer row, and especially the first, become larger, higher and more obtuse, and stand a little farther back from the inner row, the first pair be- coming the epioral spines on the small jaws. The inner spines on these few proximal plates also become a little more elongated. About eight somewhat more slender spines in a continuous row, form the peroral armature of the angle of the combined jaw plates.
The granules in the adoral area are larger and higher than elsewhere.
The ocular plate is small, roundish or broad-ovate, convex or mammiform, not bilobed and not granulated. It is turned up- ward.
The madreporic plate is large, situated below the basal inter- radial plate and