The Shallow Water Amphiurid Brittle Stars (Echinodermata, Ophiuroidea) of Florida! Lowell P

THE SHALLOW WATER AMPHIURID BRITTLE STARS (ECHINODERMATA, OPHIUROIDEA) OF FLORIDA! LOWELL P. THOMAS Institute of Marine Science, University of Miami

ABSTRACT The shallow water amphiurid brittle stars of Florida are described in detail. A survey of literature pertaining to tropical western Atlantic species is given, along with a brief discussion of general biology and morphology. Nine genera and twenty-one species are reported upon, including illus- trations and synonymies of each species. The material studied includes that collected by the author and the collections of the Museum of Comparative Zoology at Harvard, the Peabody Museum at Yale, the Allan Hancock Foundation of the University of Southern California, and the United States National Museum. A mphiodia repens (Lyman) is synonymous with Amphiodia pulchel/a H. L. Clark, Amphiodia tymbara H. L. Clark is synonymous with Amphio- dia trychna H. L. Clark, and A mphiodia rhabdota H. L. Clark is synonymous with Ophiophragmus pulcher H. L. Clark. Amphiodia trychna and Amphipholis pachybactera H. L. Clark are recorded for the first time from Florida and the U. S.

INTRODUCTION Amphiurids are long-armed mud-dwelling serpent stars which have long been known as common inhabitants of oceans and bays. They are found in all seas from the intertidal zone to a depth of several thousand meters and often occur in large populations. The importance of amphiurids in the level bottom community is well known. Thorson (1957) discussed Amphiura and Amphiodia-Amphioplus communities. In such communities, amphiurids may occur in numbers up to 500 individuals per square meter. Barnard and Ziesenhenne (1961) found that, along the California coast, A mphiodia urtica is the " ... most abundant and widespread animal in any phylum on the coastal shelves ... " It averages 422 specimens per square meter. Such communities have been found in Biscayne Bay near the Institute of Marine Science. An Amphioplus community is present with up to a hundred specimens per square meter (unpublished data, Bottom Community project). A similar Ophionephthys amphiurid community has also been found. Amphiurids, although common in Florida waters, are difficult to identify. The present study is a revision of the Florida amphiurid fauna and provides a basis for the identification of the shallow water

IContribution No. 425 from The Institute of Marine Science, University of Miami. Portions of this paper were submitted in partial fulfillment of the degree of Master of Science, University of Miami, Coral Gables, Florida. 624 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) amphiurid brittle stars of Florida and adjacent waters. The writer wishes to thank the many people who have assisted him during this study. Dr. Elisabeth Deichmann of the Museum of Comparative Zoology at Harvard has loaned specimens from the collections of the Museum and has generously given her time to answer questions. She was also instrumental in providing the author with a visiting curatorship which enabled him to study the excellent M.C.Z. collection. This curatorship was financed by funds from National Science Foundation grant G-5183. Dr. Ailsa M. Clark of the British Museum has helped to solve several systematic problems. Mr. Fred Ziesenhenne of the Allan Hancock Foundation of the University of Southern California has assisted greatly by the loan of a specimen of Ophiophragmus sept us and by his readiness to help with various problems. Dr. J. B. Balinsky, of the Univer- sity of the Witwatersrand, South Africa, generously sent specimens of his new species, Ophionephthys africana and Ophiophragmus sacensis. Dr. Willard D. Hartman of the Peabody Museum of Yale University arranged for a visiting curatorship to that institution, allowing the author to ,examine A. E. Verrill's collections. This curatorship was financed by funds from National Science Foundation grant G-10772. The completion of this paper was further aided by work in inverte- brate systematics which the author did in connection with the following grants: A National Geographic Society grant for coral reef ecology, National Science Foundation grants G-14521 and G-5941, Dingell-Johnson project funds (Project No. F-2-R-2), and a Florida Bay Estuary Ecology grant financed by the Florida State Board of Conservation. Final drafting and completion of the manuscript has been done under National Science Foundation grant G-23649. The writer wishes to express his thanks to Fenner A. Chace, Jr., Curator of Marine Invertebrates, United States National Museum and to Charles E. Cutress, Associate Curator, as well as othe'r members of the staff for their kindness and assistance during a visit to that institution. He also wishes to thank his fellow workers at The Marine Labo- ratory for their assistance during the ,course of this study, especially Robert C. Work,' J. Kneeland McNulty, Raymond B. Manning, Donald R. Moore, Anthony J. Provenzano and Durbin C. Tabb. He wishes especially to thank Dr. Gilbert L. Voss, Curator of the Marine Museum, who directed the study, Dr. Frederick M. Bayer, 1962] Thomas: Amphiurid Brittle Stars 625 Editor of the Bulletin of Marine Science of the Gulf and Caribbean, who critically reviewed the manuscript, and Drs. John E. Randall and C. Richard Robins, also of The Marine Laboratory, who made special efforts to collect material for the writer. Finally he wishes to thank his wife, Shirley B. Thomas, for her limitless aid and en- couragement. HISTORICALRESUME The first paper to mention Florida ophiuroids, by Thomas Say ( 1825), contained descriptions of many Florida echinoderms including the original descriptions of two amphiurids, Hemipholis elongata and Ophiostigma isacanthum. A second paper, by William Stimpson (1852), contained the original description of the common West Indian species, Amphipholis gracillima. Some of the most important work dealing with the ophiuroids of the West Indian region was done between the years 1867-1879 as a result of the dredging operations of the U.S. Coast Survey ships CORWIN,HASSLER,and BLAKE.During this period Theodore Lyman of Harvard and Christian F. Uitken of Copenhagen were pre- eminent in the study of West Indian species. LUtken (1859) described three species from St. Thomas, Virgin Islands, Amphiura scabriuscula, A. stimpsoni, and A. septus, in the second part of a three part work dealing largely with Caribbean forms. In 1865 Lyman described three genera, Ophiophragmus, Ophiocnida, and Hemipholis, which were based, in part, on material from Florida and the West Indies. LUtken (1869) described two new species from St. Thomas, Ophionephthys limicola and Ophionema intricata, and in the same year Lyman's report on the ophiuroids dredged in the deep water between Cuba and the Florida reef was published. In addition to describing Amphiura pulchella, Lyman provided a section on the faunal distribution of the relatively few ophiuroids known from Florida and the West Indies. In 1871 A. E. Verrill described Amphipholis abdita, a species which is common in Florida waters. Lyman's report on the ophiuroids of the HASSLERexpedition contained descriptions of two new species from the west coast of Florida, Ophiophragmus filograneus and Am- phiura repens. However, A. repens is identical with Amphiura pulchella which he had named six years previously. Lyman's paper also contains ,eighty-four figures including many of West Indian forms. Lyman's report on the ophiuroids of the CHALLENGERexpedition, published in 1882, is important to the study of the group. It included a list of all the species accepted at that time and all the generic and 626 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) specific names which had been used up to the date of publication. In 1899 the descriptions of two new West Indian amphiurid genera, Amphiodia and Amphioplus, appeared in each of two similar papers by A. E. Verrill (1899, 1899a). A faunal list of West Indian ophiuroids is included in one of these papers (Verrill, 1899a) . H. L. Clark (1901) discussed eleven species of amphiurids from Puerto Rico, and Koehler (1914) elaborated upon species collected mainly at Barbados. Koehler's 1914 paper is perhaps more important in that he reported upon a much larger number of specimens contained in the United States National Museum. Twelve of the twenty- one species in the present study were included, and three species, Hemipholis elongata (Say), Ophiocnida scabriuscula (Llitken), and Amphipholis gracillima (Stimpson), were reported from Florida for the first time. A mphiodia erecta, which Koehler described in this paper, was later identified by H. L. Clark as Ophiophragmus septus (Llitken) . A review of the Ophiuroidea by H. L. Clark (1915) is of great general importance. Called a monograph by many authors, it includes a list of all known species. A new Florida species, Ophiophragmus brachyactis, is described in this publication and Amphiura stimpsoni Llitken is recorded from the west coast of the state. In 1918 and 1919 two more papers by H. L. Clark appeared. The first deals, in part, with a collection of specimens from the Dry Tortugas and Tobago, B.W.I., and takes into account six new species important to the present work. The second deals entirely with the faunal distribution of Caribbean shallow-water echinoderms. Another faunal paper by A. H. Clark, concerned mainly with the echinoderms of Barbados and Antigua, appeared in 1921. The latest important work is H. L. Clark's (1933) Puerto Rican study which includes most of the West Indian shallow-water echinoderms and discusses the geographical range of many. The only paper dealing exclusively with the ophiuroid fauna of South Florida is that of Pearson (1936) in which the ecology of the amphiurids of Biscayne Bay is discussed. Six amphiurid species are included, but no systematic treatment is given. A recent paper by Fell (1960) gives keys to all the genera of recent ophiuroids. This monumental work will prove to be a great aid to future workers. Another work by Fell (1962), revising most of the genera of amphiurids, reached my hands after the present paper 1962] Thomas: A mphiurid Brittle Stars 627 went to press. It has therefore been impossible to comment upon it in the following pages. METHODS The present study is an enlargement of what was originally planned as a study of the shallow-water amphiurids of South Florida. Although "shallow water" in this case refers to depths from the intertidal zone to 20-30 feet, the author has made no distinction between true shallow-water species and specimens of deep-water species which have strayed into shallow water. Too little is known concerning the distribution of the West Indian Amphiuridae to make such a dis- crimination in more than a few species; however, suspected deep- water species are noted as such in the text. The study area, from Ft. Myers on the west coast to Melbourne on the east coast, has yielded all but three of the known Florida species. The systematic positions of these three are questionable and they are discussed in detail below. Besides those species previously known from Florida, two species heretofore known only from Tobago, Amphipholis pachybactera H. L. Clark and A mphiodia trychna H. L. Clark, have been collected. Two other species, Ophionephthys limicola Liitken and Ophiophrag- mus pulcher H. L. Clark, formerly unknown north of the Dry Tor- tugas, have been found to be common in Biscayne Bay. Ophioneph- thys limicola in particular is the dominant species in the level bottom community (personal communication with Moore, Work, and Mc- Nulty, see below). A fifth species, Ophiophragmus septus Liitken, taken off Miami Beach, has previously been recorded only from Cape Hatteras, Tobago, and St. Thomas. Collections were made at Melbourne, Lake Worth, and Biscayne Bay on the east coast, and at Matecumbe in the Keys. On the west coast, collections were made at Clam Slough near Marco, and Estero Bay at Fort Myers Beach. The author also collected specimens from Whitewater Bay, Coot Bay, and the Cape Sable-Flamingo region. Finally, a ten-day collecting trip to St. John in the Virgin Islands provided valuable specimens and observations, particularly regarding the genera Ophiophragmus and Ophionephthys. A great number of specimens have been made available by Dr. Hilary B. Moore, Robert C. Work, and Kneeland McNulty, of the Marine Laboratory, who have routinely collected large samples of amphiurids in the course of their Biscayne Bay Bottom Survey 628 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) project. Their deep water exploratory work, which extended to about 200 meters, has been especially valuable. The collection of amphiurid brittle stars involves removing them from the sand or mud in which they live. The author accomplished this by digging up mud with a shovel or his hands and carefully sifting the animals out, using a screen sifting box or his hands to wash the mud away. A considerable amount of collecting and observing was done at depths of from one to five meters, using aqua- lung or skin diving techniques. A sixteen-foot outboard boat was used in Biscayne Bay, and collections were made as far north as Government Cut and as far south as the Ragged Keys and Goulds Canal. All the animals captured were either permanently preserved in 70 per cent ethyl alcohol, or were placed in a jar half filled with mud and topped off with sea water until they could be properly treated with alcohol. Such handling pr,events excessive arm breakage. Formalin softens the calcareous skeleton of the arms and disk, causing the specimens to fragment, and never should be used to preserve ophiuroids. The abbreviations USNM for United States National Museum, UMML for the University of Miami Marine Laboratory, and MCZ for The Museum of Comparative Zoology at Harvard are used throughout the text.

TERMINOLOGY The terminology employed has been adopted from various sources including A. E. Verrill (1899a), Hyman (1955), Fell (1960) and A. M. Clark (1955). As an aid to the use of the present paper a list of terms and definitions is given at the end of this section. Furthermore, a hypothetical amphiurid illustrating most of these terms is shown in Figure 1. As drying sometimes distorts specimens, the author has attempted to examine fresh and alcoholic as well as dried specimens of the species included. The style of the descriptions follows, in part, Lyman's (1865) outline for describing an ophiuroid. Disk diameter is the diameter of the usually round disk. In mutilated or misshapen specimens the diameter has been estimated from several measurements. Arm length has been given as an average when the arms are all nearly the same length. Greatest and least arm length is given where there is an obvious discrepancy between the arm lengths of a single specimen. Dorsal and ventral arm plates, unless 1962J Thomas: Amphiurid Brittle Stars 629

....:., .. a .' ':., ' ~ "'f<":D:~ ~ C B

FIGURE 1. Diagram of the external features of an amphiurid brittle star. A.- Dorsal view: a, dorsal arm plate; b, arm spine; c, disk; d, accessory shield; e, papilIa; I, primitive plate; g, radial shield; h, scales. B.-Ventral view: a, adoral plate; b, half jaw; c, ventral arm plate; d, e, I, g, oral papillae; h, oral plates; i, j, oral tentacle scales; k, side arm plate; /, teeth; m, tentacle scales; n, genital plate. C.-5ide view of an arm: a, first arm spine. 630 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) otherwise indicated, include those plates on the proximal one-third of the arm, not those near the distal end. The ventral arm plates may have from four to six or more sides; the distal side known as the outer side, the proximal known as the inner side, and two lateral sides. In addition there may be diagonal outer and inner lateral sides bringing the possible total to eight (Fig. IB, c). The numbering of the arm spines begins with the dorsalmost spine and proceeds ventrally (Fig. 1C, a). The question of oral papillae and oral tentacle scales is a bother- some one. Each of the five jaws is derived from "vertebrae" of the two arms adjacent to it. Consequently, there are two pairs of tentacles homologous with the tentacles of the arm (Fig. IB, 0). Tentacle scales are often present at the base of these oral tentacles and are spoken of as oral tentacle scales (Fig. IB, i). It is sometimes very difficult to distinguish an oral tentacle scale from an oral papilla (Fig. IB, d, e, f, g), since the tentacle scales may be in line with the papillae. Lyman (1869) notes that in Amphiura semiermis the papilla at the outer corner of the mouth slit is the tentacle scale of the second pair of mouth tentacles (Fig. IB, j). Therefore, the group including A. semiermis, A. stimpsoni, and A. chiajii should be spoken of as having one pair of oral or mouth papillae to each jaw instead of three. The author has generally referred to these oral tentacle scales as papillae in the following descriptions.

GLOSSARY adoral plates: the pair of plates joining the proximal lateral sides of the oral plate. Fig. IB, a. arm spines: the spines borne by the side arm plates. Fig. lA, b. bursae: the pouches which open onto the ventral disk surface on either side of the arm (see genital pouch). disk: the round or pentagonal body to which are attached the arms and the mouth frame. Fig. lA, c. disk scales: (see scales). distal: farthest from the center of the disk; the portion of the jaw farthest from the center of the ventral disk surface. dorsal: referring to the aboral surface. dorsal arm plates: the series of plates forming the upper arm surface. Fig. lA, a. genital plate: the plates lying alongside the genital pouch. Fig. IB, n. 1962] Thomas: A mphiurid Brittle Stars 631 genital pouch: the pouches which open onto the ventral surface of the disk, usually on either side of the arm, as genital slits. half jaws: a pair of stout plates which, with the teeth and oral papillae, forms the angle of the jaw. infradental papillae: the pair of block-like oral papi11ae which lies at the proximal angle of the jaw, just below the teeth. Fig. 1B, g, f. interbrachial: the area of the disk between the arms. jaw apex: the proximal portion of the jaw. opercular: descriptive of oral papi11ae which tend to close off the mouth slit, often touching their counterparts on the adjacent jaws, as in Amphipholis. oral papillae: some or all of the papi11ae which arm the jaws. Fig. lB, d, e, f, g, j. oral plate: the single, often large, plate at the distal end of the jaw. Fig. lB, h. oral tentacles: tentacles (or podia), usually four to each jaw, which arise from the jaw and are homologous to those of the arm. oral tentacle scale: the tentacle scale of the oral tentacle, often resembling an oral papilla in amphiurids. Fig. lB, i. papillae: calcareous projections of the disk or jaws. Fig. lA, e. primitive plates: the six plates which form the aboral surface of the juvenile and may persist in the adult as a rosette. Fig. lA, f. proximal: nearest the center of the disk. radial shields: the five pairs of plates found on the aboral surface of the disk, which articulate with the genital plates. Fig. lA, g. scales: flattened calcareous plates, often free at one end, found at the base of the tentacles and on the disk in many ophiuroids. Fig. lA, h; and lB, m. side arm plates: the pair of plates making up the left and right sides of the arms and bearing the arm spines. Fig. lB, k. teeth: the wide plates which lie under the two proximal (infradental) papi11ae, and extend down into the oral cavity. Fig. 1B, 1. tentacle scales: the scales which often occur at the base of the arm tentacles, Fig. lB, m. ventral: referring to the oral surface. ventral arm plates: the series of plates forming the underside or oral side of the arm. Fig. IB, c. vertebrae: the series of internal ossicles which form the arm segments and allow arm movement. 632 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) tentacles: the pair of podia borne on the oral surface of each arm segment (on distal arm segment in Fig. IB). tentacle pore: the opening, usually guarded by a scale, through which a tentacle emerges. A KEY TO THE GENERA OF THE SHALLOWWATER AMPHIURID BRITTLE STARS OF FLORIDA The following key is intended for use in identifying the genera of amphiurid brittle stars of Florida and the West Indies as listed by H. L. Clark (1933). 1. Oral papillae not present at jaw apex; ventral surface of disk naked except

in young when it may be papillose 0 ••• 0 0 •• 0 •• 0 0 0 0 o' Hemipholis, po 686 1. Oral papillae present at jaw apex (Fig. lBog, f), ventral surface of disk

naked, scaled, or papillose 0 0 • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • • 2 2. Disk covered with short spines or papillae dorsally ...... 3 2. Disk with scales or naked, dorsal spines lacking or restricted to the edge

of the disk 0 ••• 0 0 0 0 0 ••• 0 0 0 0 0 0 0 0 •••••• 0 0 • • • • • • • • • • • • 4 3. Disk covered with spines, radial shields naked, some scales present dorsally; distal oral papillae not opercular Ophiocn ida, p. 683 3. Disk covered with short papillae, radial shields at least partially covered with papillae, no scales evident dorsally; distal oral papillae opercular

• •••••••••••••••••••••••••••••••••••••••• 0 •• 0 Ophiostigma, p. 689 4. A fence of short spines or papillae present around the edge of the disk, oral papillae fairly low and solid, not thin and fan-like · Ophiophragmus, p. 664 4. No distinct papillae around disk edge ...... 5 5. Disk almost naked dorsally 6 5. Disk completely covered with scales, dorsally 7 6. Scales only around radial shields and a single row in each interbrachius

• ••••••••••• 0 •• 0 • • • • • • • • • • • • • • • • • • • • • • • • •• Ophionephthys, p. 680 6. No disk scales present, no tentacle scales Ophionema (Not yet reported from Florida, see H. L. Clark, 1933; 46-47) 7. Distal oral papillae long, opercular, often capable of entirely closing off the gap between the jaws A mphipholis, p. 657 7. Distal oral papillae not opercular, usually rounded or spiniform ..... 8 8. Oral papillae three pairs, infradentals separated by a gap from the next papillae (which are higher and actually represent oral tentacle scales), arm spines four to seven, rarely three Amphiura, p. 633 8. Oral papillae three pairs, papillae of each side continuous; or four to five pairs of papillae to each jaw, in a discontinuous row; in either case arm spines three 9 9. Oral papillae three pairs, upright, continuous, no oral tentacle scale, dorsal arm plates nearly two times as wide as long or wider .. Amphiodia, p. 640 9. Four or five pairs of papillae to each jaw, varying somewhat in size and shape, dorsal arm plates about as wide as long, not over 1V2 times wider than long Amphioplus, p. 651 Family AMPHIURIDAELjungman, 1867 The Amphiuridae have slender arms bearing short erect spines. 1962] Thomas: Amphiurid Brittle Stars 633 Most have prominent radial shields, and the disc may be scaled, partially scaled, naked, or covered with papillae or spines. There are two distinct infradental oral papillae at the apex of each jaw (with the exception of Hemipholis) and a varying number of distal papillae. No tooth papillae are pr.esent. Genus Amphiura Forbes, 1843 Amphiura Forbes, 1843, p. 149 (in part).-Liitken, 1859, p. 114 (section Bin part).-Verrill, 1899, p. 23, 1899a, p. 306. There is one pair of oral papillae (infradentals at the proximal angle of the jaw, usually one or more spiniform or flattened papillae at the distal jaw angle, and a median (oral tentacle scale) papilla in the mouth slit above the level of the infradentals. The arm spines vary from seven or more to four and occasionally three. The upper arm plates are often as long as wide. The radial shields are usually divergent. Type species.-Amphiura chiajii Forbes, 1843 (by subsequent designation: Verrill, 1899, p. 24). The following key has been adapted from that of H. L. Clark (1933). KEY TO THE FLORIDA SPECIES OF Amphiura 1. Tentacle scales 2; tip of arm spines with prominent tooth on distal edge only palmeri, p. 633 1. Tentacle scales 1 (occasionally 2 on basal pores), arm spines not toothed or toothed on both proximal and distal edges 2 2. Oral shields as long as wide or longer; arm spines 7, decreasing to 5; dorsal arm plates elliptical or nearly circular; some arm spines toothed both distally and proximally fibulata, p. 638 2. Oral shields as wide as long or wider; arm spines 5 or 4, not toothed; dorsal arm plates rounded distally with proximal angle truncate . stimpsoni, p. 636 Amphiura palmeri Lyman, 1882 Fig. 2 Amphiura flexuosa: Lyman, 1875, p. 17, pI. 3, figs. 35-37 (non flexuosa Ljungman, 1867). Amphiura palmeri Lyman, 1882, p. 143.-Verrill, 1899a, p. 309.-Koehler, 1914, p. 55, pI. 18, figs. 1-3.-H. L. Clark, 1915, p. 230; 1933, p. 44. Amphiura kiikenthali Koehler, 1913, p. 356, pI. 20, figs. 1-4; 1914, p. 59. Material studied.-2 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 20 July, 1958; UMML 41.75.-2 spec.; Soldier Key, Biscayne Bay, Florida; May 1935; UMML 41.18.-1 spec.; Ragged Key No.3, Biscayne Bay, Florida; Thomas, 4 April, 1959; UMML 41.76.-3 spec.; Soldier Key, Biscayne Bay, Florida; Thomas, 18 April, 1959.-1 spec.; off Cape Florida, Biscayne Bay, Florida; Thomas, 13 October, 1960.- 1 spec.; Barbados, University of Iowa Exp.; 634 Bulletin of Marine Science of the Gulf and Caribbean [12 (4 )

FrGURE 2. Amphiura palmeri Lyman, 1882: A.-Disk and one arm, dorsal view, X 15. B.-Two jaws and one arm, ventral view, X 25.

USNM E. 506.-1 spec.; off Puerto Rico, Johnson-Smithsonian Exp. sta. 35; 9 Feb. 1933; USNM E. 5339.-1 spec. (as A. kiikenthali); Key West; Hemp- hill; USNM 33883. Diagnosis.-Each jaw with three pairs of papillae, infradentals block- 1962] Thomas: Amphiurid Brittle Stars 635 like, slightly wider rounded papillae in distal corners of jaws, spiniform oral tentacle scales above the infradentals; most arm spines with lateral "beak" at apex; radial shields slender, lunate, proximally separated; tentacle scales two. Description.-The diameter of the disk of one specimen measures 5.7 mm. The arms average 46 mm in length. The jaws bear three papillae on each side: a heavy, block-like, almost square infradental papilla; a slightly larger, flat, subcircular papilla at the distal angle; and a spiniform oral tentacle scale which is somewhat above the infradentals. The oral shields are ovate, slightly longer than wide, and somewhat truncate on their distal ends. The adoral shields are slightly separated at the proximal end of the oral shields. The six-sided ventral arm plates have two short inner lateral sides and are as long as wide, with their angles broadly rounded. The tentacle scale borne on the side arm plate is about twice as wide as the one on the ventral arm plate. Each narrow, band-like side arm plate bears six or seven arm spines, the first five or six equal and slightly longer than the length of a dorsal arm plate. The sixth spine is considerably longer than the others. Most of the spines, with the exception of the first and sixth or seventh, are tipped with a crest of very short spinules. On the distal edge of these spines, just below the spinules, is a large pointed beak which extends at right angles from the spine. In addition there may be a much smaller beak on the proximal side of spine number five. The dorsal arm plates nearest the disk are subcircular, the remaining ones are oval, about 1.75 times as wide as long, and overlap slightly. The disk is rounded and completely covered with scales which are very small ventrally, within the interbrachial portions of the dorsal surface, becoming larger near the radial shields. The radial shields join distally and are widely separated thereafter by several rows of scales. They are more than 3 times longer than wide and are acute proximally. This species is light tan when alive but, in alcohol, soon fades to white except for a few darker scales which probably represent primitive plates. There is usually a spot of brown pigment at the base of each arm spine. The arms may be dorsally blotched with light brown. Type.-Museum of Comparative Zoology. 636 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) Type locality.-Off Barbados, 100 fathoms. Distribution.-In deep water off Georgia, Barbados and Puerto Rico. Shallow water at Florida, the Dry Tortugas, and St. Thomas. Discussion.-Koehler (1914) examined the type of A. flexuosa Ljungman and found that it possessed simple arm spines. The specimens which Lyman described as A. flexuosa (1875) had toothed arm spines and therefore were not conspecific with Ljungman's species, but were actually specimens of Amphiura palmeri Lyman. A further point of confusion has been demonstrated by H. L. Clark (1933) who showed that Koehler's earlier descriptions (1913) of A. kukenthali and A. palmeri were in error and that the two are synonymous. As Clark pointed out, the separation of these two species was based on Koehler's assumption that A. palmeri did not possess ventral disk scales, and Koehler himself ignores this claim in 1914. Koehler's (1913) figur,e of A. ki1kenthali agrees with my specimens. Remarks.-I have taken this species with Amphioplus abditus, A. thrombodes, and Ophiocnida scabriuscula entangled in the roots of Thalassia. The numerous spines give the arms a distinctive, robust appearance. Amphiura stimpsoni Ltitken, 1859 Fig. 3 Amphiura stimpsoni Li.itken, 1859, p. 116;-Ljungman, 1866, p. 320; 1871, pp. 631-634.-Lyman, 1875, p. 4, pI. 5, fig. 66; 1882, pp. 124-143; 1883, p. 252.-Marktanner, 1887, p. 299.-H. L. Clark, 1901, p. 247.-Koehler, 1914, p. 64, pI. 7, figs. 1-2.-H. L. Clark, 1933, p. 45.-Engel, 1939, p. 8.-Fountaine, 1953, p. 200. Amphiura vivipara H. L. Clark, 1918, p. 268; pI. 1, figs. 1, 2. Material studied.-2 spec.; (locality and collector unknown); UMML 41.16. -1 spec.; Spanish water, Curacao; C. J. van der Horst, 1920; USNM E. 615. Diagnosis.-J aws with three pairs of papillae, outermost prominent and slender, infradental large and block-like, middle spiniform, above infradental; oral plates oval, lobed distally, one tentacle scale; arm spines 4 to 5; disc completely scaled; radial shields 3 times as long as wide. Description.-The arms of a specimen with a disk diameter of 3.4 mm are about 11.8 mm long. Those of a smaller specimen have a similar proportion to the disk diameter. Each jaw bears three pairs of papillae: a long, slender, distal pair; a pair of similarly shaped oral tentacle scales which lie deep in 1962J Thomas: Amvhiurid Brittle Stars 637

B .'

FIGURE 3. Amphiura stimpsoni Liitken, 1859: A.-Disk and one arm, dorsal view, X 26. B.-Two jaws and one arm, ventral view, X 40. the oral grooves almost directly under the distal papillae; and two large rounded infradentals. The oral shield is sub-oval and wider than long. In the two dried 638 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) UMML specimens the adoral shields are slightly separated at their proximal ends. The ventral arm plates are about one-fourth longer than wide and superficially appear to be rectangular. There are, however, two inner lateral sides which appear on the distal plates. A single tentacle scale lies in the angle of the ventral and side arm plates. Each side arm plate bears four or five slender spines which, near the base of the arm where there are five, decrease in length from the fifth to the third. The first and second spines are about the same length as the fifth. When there are only four spines they decrease in length from the fourth to the second, with the first about equal in length to the fourth. The dorsal arm plates are slightly wider than long, broadly rounded distally, truncate proximally, and slightly overlapping. The disk is covered with delicate irregularly overlapping scales. The radial shields are about 3 times longer than wide, taper to a point, and are entirely separated except along their distal third or fourth. Both specimens were dirty white with no visible color pattern. Type.-Zoological Museum, Copenhagen(?) Type locality.-St. Thomas, V. I. Distribution.-West coast of Florida; the Dry Tortugas; St. Thomas; Barbados; Tobago; Curacao; and Brazil. Discussion.-Unfortunately I have not taken any samples of this well-known species. Although H. L. Clark (1933) found stimpsoni in coralline algae at the Dry Tortugas, I have found no specimens in a similar habitat along the Florida coast. A mphiura fibulata Koehler, 1913 Fig. 4 Amphiura fihulata Koehler, 1913, p. 359; 1914, p. 56, pI. 7, figs. 3-5.- H. L. Clark, 1915, p. 234, 1933, p. 45. Material studied.-l spec. (holotype); Gulf of Mexico, FISH HAWK sta. 7295, 10 metrs, 24 Feb. 1920; USNM 32293.-2 spec.; the Dry Tortugas, Florida, 2-3 meters; MCZ 4736. Diagnosis.-Oral papillae three pairs, infradentals closely appressed, rather small, a second pair (oral tentacle scales) slightly higher and extending proximally to the infradentals, the distal pa1Jillae small. arising from the adoral shields near the proximal end of the oral shield; oral shield longer than wide, rhomboidal; adorals not meeting 1962] Thomas: A mphiurid Brittle Stars 639 in front of the oral shield; tentacle scale single; arm spines five to seven, where seven are present the fifth bears a single tooth on both edges; upper arm plates almost round; disk scaled dorsally and ventrally; radial shields united distally. Description.-A specimen with the disk 8 mm in diameter (MCZ 4736) has the arms broken but they appear to have been quite long in life. The infradental papillae are small and rather close together. The distal papillae are slightly smaller than the infradentals. A third papilla (oral tentacle scale) lies higher in the oral cavity between the infradental and the distal papillae. The distal papillae arise from the adoral shields near the proximal end of the oral shield and tend to be bluntly pointed at their free ends. The middle (oral tentacle scales) papillae are longer than either of the other two pairs and are rather

FIGURE 4. Amphiura fibulata Koehler, 1913 (MCZ 4736): A.-Portions of the disk and an arm, dorsal view, X 12. B.-One jaw, X 25. C.-Ventral arm surface and a 5th arm spine from segment number 22. acutely pointed. The oral shields are roughly rhomboidal with broadly rounded angles, the distal angle being more acute than the proximal. The adoral shields are separated proximally in all of the specimens examined. The ventral arm plates are rectangular with rounded corners and are slightly longer than wide. At the angle of junction of the side and ventral arm plates there is a single tentacle scale. The narrow side arm 640 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) plates bear five to seven flattened, rather blunt arm spines. Usually each of the two spines immediately above the ventralmost spine is provided with two terminal teeth. These teeth project laterally from the spine and are on opposite sides. The dorsal arm plates are almost round at the proximal end of the arm, becoming more rectangular distally. The disk is covered by rather small imbricated scales which are smallest on the ventral surface. The radial shields are about 3 times longer than wide and are bluntly rounded at both ends. They touch distally but are separated by about their own width proximally. Both dried and alcoholic specimens lack a distinctive color pattern. Type.-U.S.N.M., cat. no. 32293.

Type locality.-FISH HAWK station 7295, Gulf of Mexico, Lat. 24°38'40"N.; Long. 81°56'26"W., 5th fathoms. Distribution.-Type locality and the Dry Tortugas, Florida. Discussion.-Koehler (1914) gives a detailed description of this speCIes. Genus Amphiodia Verrill, 1899 Amphiodia Verrill, 1899, p. 25; 1899a, pp. 306, 313. There are three (rarely four) pairs of similar oral papillae in a regular series, which are either block-like or erect, never opercular. The jaw slits are prominent, and no oral tentacle scales are visible. Three (rarely four) arm spines are present on each side arm plate. The upper arm plates are usually wider than long. The dorsal surface of the disk is scaled, bearing radial shields which often are joined. Several genera are very similar to A mphiodia. An example is Ophiophragmus, which has a fence of papillae in addition to Amphi- odia characters. Ophiocnida has the disk covered by short spines. The validity of these features as taxonomic characters is questionable. For example, the author has seen specimens of Ophiophragmus pulcher and O. filograneus which completely lacked fence papillae and differed in no external way from an A mphiodia. As a further confusing point, the adult Ophiocnida cubana has a fence of papillae just as an Ophio- phragmus, while the disk of the juvenile is completely covered by short spinules. There is no question that a great deal of work must be done before the true affinities of the various species in the above genera are understood. 1962] Thomas: Amphiurid Brittle Stars 641 Type-species.-Amphiura pulchelia Lyman (by subsequent designation: Verrill, 1899, p. 25).

KEY TO THE FLORIDA SPECIES OF A mphiodia I. Tentacle scales 1 pulchella, p. 641 1. Tentacle scales 2 2 2. Disk scales usually thick and swollen, adorals meet proximal to ora] shield ...... trychna, p. 645 2. Disk scales not swollen, adorals do not meet proximal to ora] shield ...... planispina, p. 648 A mphiodia pulchelia (Lyman, 1869) Fig. 5 Amphiura pulchelia Lyman, 1869, p. 337.-Ljungman, 1871, p. 648.- Lyman, 1875, pI. 5, fig. 75; 1882, pp. 125, 147. Amphillra repens Lyman, 1875, p. 18, pI. 3. figs. 38-40. Amphiodia Plilchella: Verrill, 1899, p. 25; 1899a, p. 313.-H. L. Clark, 1901, p. 248; 1915, p. 250; 1933, p. 54.-Koehler, 1914, p. 70. Amphiodia repens: Verrill, 1899, p. 25; 1899a, p. 313.-H. L. Clark, 1915, p. 249; 1933, p. 54.-Fontaine, 1953, p. 201. Material studied.-l spec.; Marine Lab. Beach, Virginia Key, Florida; Thomas, 27 August, 1958; UMML 41.79.-3 spec.; W. of Virginia Key, Biscayne Bay, Florida; Thomas, 31 March, 1958; UMML 41.78.-17 spec.; Marine Lab. Beach, Virginia Key, Florida; Thomas, ] September, 1958.-1 spec. (as Am- phil/ra 'repens); between Miss. River Delta and Cedar Key, Fla.; ALBATROSS sta. 2373, 1885; USNM 15341.-1 spec. (as Amphiura repens); between Miss. River Delta and Cedar Keys, Florida; ALBATROSSsta. 2372, 1885; USNM 15407.-2 spec. (as Amphiura repens); off Cape Catoche, Yucatan; ALBATROSS sta. 2362, 1885; USNM 15427.-2 spec.; off Rio de la Plata, Argentina; AL- BATROSSsta. 2765, 12 Jan. 1888; USNM 33886.-1 spec.; San Antonio Bridge, Puerto Rico; FISH HAWK, 12 Jan. 1899; USNM 21342.-11 spec.; (as Am- phiodia repens); Biscayne Bay, Florida; H. L. Clark, April 1937; MCZ 5510. Diagnosis.-Oral papillae three pairs, distal two pairs erect, similar; oral shield lobed distally, rounded or bluntly pointed proximally; adoral shields not meeting proximally; tentacle scale single, arm spines three, somewhat flattened, bluntly rounded; disk scaled dorsally and ventrally, although ventral squamation may be fine. Description.-A specimen 5 mm in diameter has arms varying from 43 to 47 mm in length. The largest specimen has a disk 5.2 mm in diameter. Each narrow jaw bears three pairs of contiguous oral papillae, the block-like infradentals and the distal two, erect, rounded, and faintly toothed. The oral shields are rounded or slightly pointed proximally and lobed distally. in the shape of a classic arrowhead or spade. The ventral arm plates, longer than wide and pentagonal, possess a 642 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

FIGURE 5. Amphiodia pulchella (Lyman, 1869): Upper.-Disk and one arm, dorsal view, X 15. Lower.-Two jaws and one arm, ventral view, X 30. slightly concave outer side, lateral sides, and inner lateral sides. There is one prominent tentacle scale on the ventral arm plate at the angle of the side arm plate. The three arm spines are somewhat flattened 1962] Thomas: Amphiurid Brittle Stars 643 and very finely toothed, particularly the middle one, which is bluntly rounded or truncate. It is also the shortest, and is flattened horizon- tally, whereas the other two spines are flattened vertically. The dorsal arm plates bear a straight, slightly undulate, or convex outer side, decidedly convex inner side, and are about 1.5 times as wide as they are long. The disk is covered with numerous small scales, among which the six primitive plates are almost always present in specimens of 4 mm disk diameter or less, but disappear in larger individuals. The radial shields, over twice as long as wide, are joined almost their entire length, being separated proximally by a single small scale. The disk is scaled ventrally although the scales may be very small and widely spaced in some specimens. The disks of alcoholic specimens remain gray but the arms fade to whitish. In both dried and living material the arms are usually blotched with dark gray. Often a faint light stripe extends down the dorsal surfa.ce of the arm. In some alcoholic specimens this stripe is secondary to a series of light brown or gray spots, one of which graces the outer edge of each dorsal arm plate. Type.-Museum of Comparative Zoology (Dr. Elisabeth Deichmann informs me that the holotype cannot be found). Type locality.-Straits of Florida, 72 meters. Distribution.-Bermuda; Florida; Gulf of Mexico; Jamaica; Puerto Rico; St. Lucia; Tobago; Brazil; and off Rio de la Plata, Argentina. Discussion.-All of the specimens which I have examined possess scales on the ventral surface of the disk, although ventral scalation may be difficult to see unless the specimen is dried. The majority of specimens under 4 mm in disk diameter bear a rosette of primitive plates. Of the 21 specimens in the Marine Laboratory .collections, only one with a disk diameter of 4 mm lacks primitive plates, and four specimens (including one of 5 mm) larger than 4 mm have primitive plates. The oral shields are spade-shaped, as Lyman (1875) described and figured for A. repens. The arm spines tend to be flattened, but in smaller specimens this is not obvious. In view of the differences between large and small specimens, I believe that repens is merely the adult of what Lyman described in 1869 as pulchella. Small specimens are more heavily scaled ventrally and have primitive plates and slender arm spines. As they grow the ventral disk scaling becomes sparse, the primitive plates disappear, and the arm spines flatten and become 644 Bulletin of Marine Science of the Gulf and Caribbean [J 2 (4)

FIGURE 6. Amphiodia trychna H. L. Clark, 1918: A.-Disk and one arm, dorsal view, X 16. B.-Two jaws and one arm, ventral view, X 24. 1962] Thomas: Amphiurid Brittle Stars 645 blunt. Dr. Elisabeth Deichmann (personal communication) has reached the same conclusions regarding the status of Amphiodia repens. It appears certain that Lyman was mistaken in separating these larger specimens as A. repens. Therefore this species must be declared synonymous with A mphiodia pulchella. A mphiodia trychna H. L. Clark, 1918 Figs. 6, 7 Amphiodia trychna H. L. Clark, 1918, p. 289, pI. 3, figs. 1-3; 1933, p. 54. Amphiodia tymbara H. L. Clark, 1918, p. 290, pI. 2, fig. 6; 1933, p. 53. Material studied.-l spec.; off Key Biscayne, Florida, 38 meters; Robert Work and J. Kneeland McKnulty, 20 January, 1957; UMML 4] .105.-1 spec. (holotype); British West Indies, Tobago, Buccoo Bay, Sandy Point, sandy mud, 1 meter; April, 1916; MCZ 4144.-1 spec.; Biscayne Bay, Florida; April, 1927; MCZ 5504.-1 spec.; 2 mi. S.W. of La Parguera, Puerto Rico, 2 meters, sandy mud; Norman Mattox, 9 July, 1957; Allan Hancock Foundation, cat. no. 7061. -1 spec. (holotype of A. tymbara); Sandy Point, Buccoo Bay, Tobago, British West Indies, 1 to 3 ft., sandy mud; April, 1916; MCZ 4145.-1 spec. (as A. tymhara); off Boynton, Florida, about 160 meters; 1941 (diskless, identification questionable); MCZ 6639.-2 spec. (as A. planispina); the Dry Tortugas, Florida, about 3 meters; MCZ 4734. Diagnosis.-Oral papillae three pairs, contiguous, thick; oral shields diamond-shaped; tentacle scales two; arm spines three, flattened, as long as width of ventral arm plate; disk with thick, coarse scales often inflated; primitive plates may be prominent; radial shields about twice as long as wide, joined over half their length. Distal end of dorsal arm plates white, proximal to this a dusky band. Description.-The disk of the dried specimen at the Marine Labora- tory measures 4.8 mm in diameter. The total arm length cannot be determined, as the arms are too badly broken to permit an estimate. The longest broken arm is 36 mm. Each half jaw bears three thick, contiguous oral papillae of which the distal one is the largest. The infradental papillae are widely separated. Pairs of short half jaws, thick adoral shields, and a single thick, diamond-shaped oral shield slightly longer than wide complete the oral aspect. The ventral arm plates are about 1.5 times wider than long and bear one of the two prominent tentacle scales. The second and largest of these scales is borne by the side arm plate. Of the three rather long, blunt arm spines, the third is the longest, the first and second being about equal, somewhat longer than the length of a dorsal arm plate. All the spines tend to be quite flattened. The dorsal arm plates are 646 Bulletin pf Marine Science of the Gulf and Caribbean [12(4)

~i .. . ..-~. ;1.. .. . '

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B FIGURE 7. A.-Amphiodia trychna H. L. Clark, 1918 (holotype, MCZ 4144): 1, a single jaw, X 30; 2, portions of the disk and an arm, dorsal view, X 18; 3, ventral arm surface and arm spines. B.-Amphiodia tymbara H. L. Clark, 1918 (holotype, MCZ 4145): 1, a single jaw, X 18; 2, portions of the disk and an arm, dorsal view, X 12; 3, ventral arm surface and arm spines. about 2.5 times wider than long and very slightly convex, sometimes with a reentering central concavity. The rather inflated disk is entirely covered by thick, irregular scales which often overlap. Near the edge of the disk, where the ventral scales meet the dorsal scales, there may be as many as three or four scales piled on one another. Smaller scales partially separate the large radial shields which are joined two-thirds of their length. Although the color of the Marine Laboratory specimen has faded, the pattern has been retained. The disk and arms are light tan. Each 1962] Thomas: Amphiurid Brittle Stars 647 dorsal arm plate has, except near the disk, a fairly broad transverse line of light tan just inside its outer edge. Ther,e is a border of white along the outer edge. Periodically the tan bands darken as do the cor- responding dorsal shields. A secondary banding is thus produced. Larger specimens (MCZ 4734 and 4145) differ in certain respects from smaller ones. Their disk scales do not overlap to a great extent. The upper arm plates widen greatly in proportion to their length and may become slightly concave along the distal edge. The ventral arm plates widen with age. The oral aspect remains relatively unchanged except for an elongation of the oral shield which may be lobed at its distal end (Fig. 7). The arm color apparently fades somewhat with age. Type.-Museum of Comparative Zoology, ,cat. no. 4144. Type locality.-Sandy Point, Buccoo Bay, Tobago, British West Indies. Distribution.-This species was first discovered at Tobago (H. L. Clark, 1918). It has since been taken along the Florida east coast and at La Parguera, Puerto Rico. Discussion.-One of the major problems confronting the student of brittle stars is the identification of juvenile specimens. A. M. Clark (1953) pointed out the difficulty in distinguishing between mere growth stages and valid specific characters in the genus Ophionereis. The same problem undoubtedly occurs in most ophiuroid genera. An example of misinterpretation of characters is the A mphiodia trychna -A mphiodia tymbara problem. H. L. Clark (1918) described A mphiodia trychna from a specimen with a disk diameter of only 3.5 mm. On the following page of the same publication he described A. tymbara from a specimen with a disk diameter of 8 mm, stating that the diameter would have probably been greater had the disk not been recently regenerated. Dorsally the two specimens do not appear very similar (Fig. 7). The disk scales are comparatively thicker and less numerous in the type of A. trychna and the radial shields are narrow,er proximally. However, we cannot depend on the type of A. tymbara to have a "typical" disk in view of the obvious recent regeneration. The dorsal arm plates of A. tymbara are wide but those of A. trychna are only slightly wider than long. This difference may be explained by the stage of development. Young specimens of some amphiurids tend to have much narrower dorsal arm plates than do the larger, older ones. 648 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) There is a very evident similarity between the oral papillae, jaws, and adoral shields of the two specimens. The oral shields of A. tymbara are comparatively longer than those of A. trychna, but the shape falls within a reasonable limit of individual variation and may be considered a growth difference. The ventral arm plates of the two are very similar, as are the arm spines. In both the arm spines are thick, somewhat flattened, and have bluntly rounded ends. The ventral spine is the longest. The color pattern on the upper arm plates of A. trychna is more pronounced than, but identical with, that of A. tymbara. It is difficult to see why H. L. Clark (1918, pp. 289-290) says of A. trychna, "It is unlike any other West Indian brittle-star ... in coloration," when one compares the color description he gives for the two specimens. Of A. trychna he says, "... every upper arm plate, regardless of its ground color, has the distal margin white and is crossed, immediately behind this white margin by a very distinct band of deep gray; on plates which are themselves deep gray, this band is almost black," of A. tymbara, " ... the upper arm plates show much diversity and no sort of se- quence in ground color, but the distal margin is generally white and often back of that is an indistinct dusky transverse band." That the color pattern may vary greatly in intensity within a species of brittle star is to be expected. This may be observed in almost any large collection. The important fact is that the basic pattern usually remains constant. In view of the evidence, I feel that the type specimen of A. tymbara is conspecific with that of A. trychna, the latter being a young specimen. I choose Amphiodia trychna as the valid species name in order to avoid the type of A. tymbara which appears to have an imperfect disk. Remarks.-Two MCZ specimens listed as A. pl'lnispina (MCZ 4734) were reidentified by A. M. Clark as A. tymbara. The larger of these has coarse disk scales like the type of A. trychna, while the smaller one has smaller, thinner scales.

Amphiodia planispina (von Martens, 1867) Fig. 8 Amphiura planispina von Martens, 1867, p. 347.-Lyman, 1882, p. 146. Amphiodia planispina: Verrill, 1899, p. 313.-H. L. Clark, 1915, p. 248. pI. 8, figs. 8-9; 1918, p. 287; 1933, p. 53.-A. H. Clark, 1921, p. 53. Material studied.-3 spec.; E. coast of South America, 22 meters; 12 Jan., 1888, ALBATROSSsta. 2764; USNM 39009 (smallest one questionable).- 1962] Thomas: Amphiurid Brittle Stars 649 1 spec.; off Rio de la Plata, Argentina, 20 meters; 12 Jan., 1888, ALBATROSS sta. 2765; USNM 33983.-1 spec. (cotype); Rio de Janiero, Brazil; MCZ 1449. 1 spec. (cotype?); Rio de Janeiro, Brazil; von Martens; MCZ 695.-1 spec.; MCZ 1531.-1 spec. (questionable); Porto Seguro, Brazil; Thayer expedition; MCZ 1450.-1 spec. (questionable); the Dry Tortugas, Florida; H. L. Clark, June 1917; MCZ 4236. Diagnosis.-Disk covered with well defined, imbricated scales; radial shields joined over half their length, slightly longer than wide; arm spines three, quite flattened, blunt; dorsal arm plates 3 times as wide as long; two tentacle scales; oral shield diamond shaped, adorals do not meet proximal to oral shield; three pairs of contiguous oral papillae, distal papillae the largest, infradentals smallest. Description.-The arm length is about 10 times the disk diameter, which is 10.4 mm in MCZ 1449. Each half jaw bears three contiguous papillae: the infradental, which is the smallest and widely separated from its mate; the somewhat larger second one; and the very large distal papilla, which is about twice as large as either of the others.

FIGURE8. Amphiodia planispina (von Martens, 1867) (syntype, MCZ 1449): A, a single jaw, X 13. B, ventral arm surface, X 7; C, arm spines from: 1, the arm base: 2, the 10th segment; 3, the 55th segment. D, portion of the disk and an arm, dorsal view, X 11. 650 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) The oral shields are diamond-shaped and longer than wide. There is a tendency for the distal end to be lobed. The adorals are straight along their proximal edge and extend laterally well past the edge of the largest oral papillae. They do not join proximal to the oral shield and may disappear under its end. The v.entral arm plates are rectangular with very broadly rounded corners. There are two tentacle scales, which are wider than long and close together. The disk is covered by prominent, imbricated scales. Those on the dorsal surface meet those of the ventral surface to form a rather sharp angle. The radial shields are 1.25 times longer than wide and joined for two thirds of their length. The dorsal arm plates are at least 3 times wider than long and tend to be slightly concave along the distal edg,e. There are three very flattened, blunt arm spines. Near the disk the uppermost spine is longest, with the second and third shorter respectively. By the tenth segment, the first and second spines are about the same len~th while the lowermost is slightly longer. At the fifty-fifth segment the uppermost spine is by far the shortest while the second and third are about the same length. The arms are irregularly banded with dusky purplish-gray. Type.-Syntypes at the Museum of Comparative Zoology, MCZ 695 and 1449 (the former probably type material but questionable). Type locality.-Rio de Janeiro, Brazil. Distribution.-Rio de la Plata, Argentina; northward along the Brazilian coast. The Dry Tortugas, Florida (?). Discussion.-Those features which define A. planispina and distinguish it from similar sp.ecies are the broad, flattened, bluntly rounded arm spines and the separated adoral plates. If these characters are non-conservative, A. planispina might well be an extreme variation of another species of Amphiodia. However, it is likely that planispina is a valid species, for specimens taken from stations over a thousand miles apart (MCZ 1449 and USNM 39009) show little variation of characters. It is unfortunate that the species has no distinctive color pattern to aid in identification. Although the arms are mottled and banded, this is a feature common to many amphiurids. The other West Indian Amphiodia differ in various respects from A. planispina. Amphiodia trychna has a distinctive color pattern, more slender arm spines, and adoral shields which are in broad con- 1962] Thomas: Amphiurid Brittle Stars 651 tact proximally. A mphiodia atra, A. gyraspis, and A. limbata (all closely related, if not conspecific) have slender arm spines and are quite different in other respects. A single specimen from Rio de Janeiro labeled A. rUsei (MCZ 1455) has slender spines and five-sided v,entral arm plates. A mphiodia pulchella has but a single tentacle scale. Remarks.-The .card catalogue in the Museum of Comparative Zo- ology lists the dry co-type (MCZ 1449) as having a disk which is detached, but the box contained only arm fragments. In an adjacent box (1450) there were two oral frames and a single disk. The card file listed MCZ 1450 as being a single diskless specimen. r found that the arm fragments in MCZ 1449 matched the disk and one of the oral frames in MCZ 1450. These were put with the arm fragments in MCZ 1449 and explanatory notes were added. An alcoholic specimen, apparently the second cotype mentioned by H. L. Clark in 1915, is identical with the specimen which I removed from MCZ 1450 and placed, correctly, I believe, in MCZ 1449. Genus A mphioplus Verrill, 1899 Amphioplus Verrill, 1899, p. 25; 1899a, pp. 306, 314. The following generic description is taken directly from Verrill (1899): "Four or five small unequal oral papillae, none operculiform, usually arranged in a discontinuous series, of which the outermost, at least, arises from the adoral shield and is really a distal oral tentacle- scale. Arm-spines thr.ee (rarely four). Radial shields generally quite separated. Disk scales naked." Type species.-Amphiura tumida Lyman (by subsequent designation: Verrill, 1899, p. 25) KEY TO THE FLORIDA SPECIES OF Amphioplus 1. Tentacle scales 1; dorsal disk scales swollen at their distal edge · ...... thrombodes, p. 654 1. Tentacle scales 2 2 2. Ventral disk surface naked; second arm spine not flattened · coniortodes, p. 656 2. Ventral disk surface scaled, second arm spine flattened, somewhat truncate · abditus, p. 651 Amphioplus abditus (Verrill, 1871) Fig. 9 Amphipholis abdita Verrill, 1871, p. 132.-Lyman, 1875, pI. 5, fig. 82. Amphiura abdita: Lyman, 1882, p. 148.-Koehler, 1907, p. 306, pI. 11, figs. 24, 25.-Coe, 1912, p. 82, pI. 19, fig.!. 652 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) Amphiura macilenta Verrill, 1882, p. 142. Amphioplus macilenta: Verrill, 1899, p. 314.-Coe, 1912, p. 84, pI. 19, fig. 1. Amphioplus abditus: Verrill, 1899a, p. 314.-Koehler, 1914, p. 71.-H. L. Clark, 1915, p. 251; 1933, p. 55. Material studied.-l spec.; Clam Slough, Marco Beach, Florida; Thomas, 30 December, 1958; UMML 41.99.-41 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 2 April, 1958; UMML 41.100.-14 spec.; Lower Matecumbe Key, Florida; Thomas, 2 September, 1958; UMML 41.101.-2 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 12 July, 1958.-50 + spec. (syntypes); Noank Harbor, Conn.; 25 August, 1874; USNM 29574.-10 -+- spec. (syntypes of Amphiura maci/enta Verrill); off Martha's Vineyard, Mass.; SPEEDWELLsta. 920. 1880; USNM 18007.-4 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 13 July, 1958.-1 spec.; West end of Virginia Key, Biscayne Bay, Florida; Thomas, 31 March, 1958.-1 spec. (syntype of A. maci/enta); MCZ 1473.-5 spec. (syntypes of A. macilentus); U.S.F.C. 14792, sta. 920; Peabody Museum, Yale University. Diagnosis.-Entire disk finely scaled; radial shields straight, two and one-half times longer than wide, separated except at base; arm spines three, second spine largest, flattened, broadly rounded at tip; arm length 10 times disk diameter; tentacle scales two; each jaw with five pairs of papillae. Description.-The arm length is ten times the diameter of the disk which, in a large specimen, may be 10 mm. Each jaw bears five pairs of papillae. The infradental papillae are the largest, block-like, and usually widely separated. They are fol- lowed in close order by three (sometimes two) similar papillae and, distally removed from the others, a fifth papilla which is occasionally double. The oral shields are variable, more or less diamond-shaped, and usually 1.5 to 2 times longer than wide. The ventral arm plates are p.entagonal, with two inner lateral sides. There are two tentacle s.cales, the larger arising from the side arm plate, the smallest from the ventral arm plate. The disk is somewhat angular and completely covered with small, closely appressed scales which are largest near the radial shields. A faint line of very slightly raised scales ,extends around the edge of the dorsal disk surface in ,each interbrachius. The straight radial shields are about 2.5 times longer than wide, touch distally, and are separated the remaining length by two or more scales. The dorsal arm plates are broadly elliptical and barely touch each other. Each side arm plate extends almost halfway across the dorsal arm surface and bears three spines of which the second is largest, usually flattened, broadly rounded distally, and .covered with short 1962] ThDmas: Amphiurid Brittle Stars 653

FIGURE 9. Amphiopilis abdillis (Verrill, 1871): A, disk and one arm, dorsal view, X 9; B, two jaws and one arm, ventral view, X 18. 654 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) conical spinules which are visible in dried material. This species lacks a distinctive color pattern. The disk and arms are light gray, the latter blotched with darker gray and the former often peppered with small, dark gray spots. Type.-US.N.M., cat. no. 18007. Type locality.-Verrilliists "... off Thimble Islands and Savin Rock, near New Haven [Connecticut] ... " as the typ.e locality but all the syntypes I have examined came from Noank Harbor, Connecticut. Distribution.-East coast of the United States to West Florida. One record from Panama (Clark, 1933). Discussion.-In many respects this species r,esembles A. coniortodes and A. thrombodes. It differs, however, in that the disk is completely scaled, there are two subequal tentacle scales (A. thrombodes has but one), and the middle arm spine lacks the acute point which character- izes the two other species. Koehler (1914) synonymized Amphioplus abditus and Amphioplus macilentus Verrill (1882). I have examined Verrill's syntypes of both species and agree that they appear to be identical. Remarks.-This is the most common intertidal amphiurid of South Florida. It lives among the roots of marine angiosperms in the intertidal zone and down to about 140 meters (Clark, 1933). Amphioplus thrombodes H. L. Clark, 1918 Fig. 10 A Amphioplus thrombodes H. L. Clark, 1918, p. 292, pI. 7, figs. 1, 2; 1933, p.56. Material studied.-2 spec.; Estero Bay, Ft. Myers, Florida; Thomas, 12 April, 1959.-1 spec.; Lake Worth, Florida; Thomas, 11 April, 1959.-35 spec.; Clam Slough, Marco Beach, Florida; Thomas, 30 December, 1958; UMML 41.97.-1 spec.; west side of Virginia Key, Biscayne Bay, Florida; 31 March, 1959; UMML 41.98.-7 spec.; two miles south of Cutler Power Plant, Bis- cayne Bay, Florida; Thomas, 11 April, 1958.-6 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 2 April, 1958.-1 spec. (holotype); Key West, Florida; H. L. Clark, June, 1917; MCZ 4218. Diagnosis.-Disk completely covered with scaLes; scales of dorsal surface swollen on their distal edges producing prominent raised "papillae"; radial shields joined distally, straight, twice as long as wide; each jaw bears five pairs of papillae; one tentacle scale. Description.-A specimen with the disk 6 mm in diameter has arms 65 mm long. The oral armature of this species is so much like that of A. abditus that to describe it would be repetitious. The oral shields tend to be 1962] 655

FIGURE 10. A.-Amphioplus thrombodes H. L. Clark, 1918: two jaws and one arm, ventral view, X 22. B.-Amphioplus coniortodes H. L. Clark, ]918: two jaws and one arm, ventral view, X 30. 656 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) truncate distally. A single rather small tentacle scale arms the pentagonal ventral arm plates. The three arm spines are slender and pointed, the second the largest, but possessing an acute point as do the first and third. The dorsal disk scales are swollen at their distal edge, giving the disk surface a distinctly papillose app.earance. The scales of the ventral disk surface are very small and widely scattered. The radial shields are straight, joined distally, and 1.5 to 2.5 times longer than wide. The disk of this species is often light gray. The arms are tan, blotched with dark gray. Type.-Museum of Comparative Zoology, cat. no. 4218. Type locality.-Key West, Florida. Distribution.-Ft. Myers; Biscayne Bay; and Key West, Florida. Discussion.-This species differs from both A. abditus and A. conior- fodes in having but one tentacle scale and in the faintly papillose disk surface. Like A. coniortodes, the second arm spine is round and acute. Remarks.-I have taken this species only with A. abditus, never with A. coniortodes. Amphioplus coniortodes H. L. Clark, 1918 Fig. 10 B Amphioplus coniortodes H. L. Clark, 1918, p. 291, pI. 7, figs. 3, 4; 1933, p.56. Material studied.-ll spec.; south side of Virginia Key, Biscayne Bay, Florida; Thomas, August, 1958; UMML 41.95.-5 spec.; southwest side of Virginia Key, Biscayne Bay, Florida; Thomas, 31 March, 1958; UMML 41.96.-1 spec. (holotype); Key West, Florida; H. L. Clark, June, 1917; MCZ 4217. Diagnosis.-Disk finely scaled dorsally, naked ventrally; radial shields joined distally, separate proximally, slightly curved inward, four or more times longer than wide; arm spines three, similar; longest arm 20 times disk diameter; five pairs of papillae on each jaw; tentacle scales two, sub-equal. Description.-A specimen with the disk 6 mm in diameter has arms 128 mm in length. The second proximal papilla is rather spiniform, otherwise the dentition is nearly identical to that of A. abditus. The oral plates are almost oval and tend to be about as long as wide, although there is considerable variation. There are two tentacle scales, the one on the side arm plate the larger. The ventral arm plates are pentagonal. The side arm plates each bear three similar acute spines. The second or middle spine is the 1962] Thomas: Amphiurid Brittle Stars 657 thickest, but, like the first and third, tapers to a point. The disk is distinctly pentagonal, and its dorsal surface is completely covered by fine scales. A prominent row of larger, somewhat raised scales extends along the lateral dorsal surface. The radial shields are from 3.5 to 5 or more times longer than wide and touch distally. They are not straight, curving toward ,each other proximally. The disk is gray. The arms are tan, blotched with dark gray. Type.-Museum of Comparative Zoology, ,cat. no. 4217. Type locality.-Key West, Florida. Distribution.-Type locality and Biscayne Bay, Florida. Discussion.-This species may be recognized by its very long arms and naked ventral disk surface. It may be distinguished, even after losing its disk, by its three slender acute spines, paired tentacle scales, and somewhat oval oral shields. Remarks.-A. coniortodes is commonly associated with Ophioneph- thys limicola in soft mud. It does not often occur in the intertidal zone, as does A. abditus, but prefers deeper water. Genus Amphipholis Ljungman, 1866 Amphipholis Ljungman, 1866, pp. 165, 311, 644 (in part).-Verrill, 1899, p. 24; 1899a, pp. 306, 311. There are three or four oral papillae in a continuous series, of which the distal most is flat and opercular, tending to cover the oral slit. The arm spines vary from three to four and are usually small and pointed. The radial shields are often in close contact along most of their length. Type-species.-A mphipholis januarii Ljungman, 1866=Ophiolepis gracillima Stimpson, 1852 (by monotypy). A KEY TO THE FLORIDA SPECIES OF Amphipholis 1. Arm spines flattened, armed with large terminal teeth pachyhactera, p. 657 1. Arm spines not armed, but rounded and bluntly or acutely pointed . . . 2 2. Radial shields 3 times longer than wide, very slender; arms long, slender, more than 10 times disk diameter gracillima, p. 660 2. Radial shields about 2 times longer than wide; arms short, at best 5 or 6 times disk diameter squamata, p. 662 A mphipholis pachybactera H. L. Clark, 1918 Fig. 11 Amphipholis pachyhactera H. L. Clark, 1918, p. 284, pI. 1, figs. 3-5; 1933. p.49. Material studied.-6 spec.; Coral Harbor, St. John, V.I.; Thomas, 19 December, 1958; UMML 41.72.-1 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 12 658 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) July, 1958; UMML 41.83.-6 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 20 July, 1958; UMML 41.84.-2 spec.; Bear Cut, Biscayne Bay, Florida, from peat deposit; Thomas, 23 January, 1959; UMML 41.85.-2 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 22 January, 1959; UMML 41.86.-1 spec.; Key West, Florida; H. Hemphill, 1885; USNM 34012 (as Amphipholis sp.). -1 spec. (as Amphipholis gracillima); 7 miles off Little River Inlet, South Carolina; A. S. Pearse, 17 August, 1949; USNM E. 7860.-1 spec. (holotype); Sandy Point, Buccoo Bay, Tobago, British West Indies; H. L. Clark, April, 1916; MCZ 4140.-6 spec. (paratypes); Sandy Point, Buccoo Bay, Tobago, British West Indies; H. L. Clark, April, 1916. MCZ 4141. Diagnosis.-Three pairs of oral papilla.e, the outermost wide, opercular; disk completely scaled; radial shields twice as long as wide, slender, joined along distal two thirds; arm spines three or four, second or third spine respectively has toothed lateral edges, two tentacle scales. Description.-One specimen measures 4.2 mm across the diameter of the disk and has arms ranging from 42 to 50 mm long. Ther.e are three pairs of oral papillae: the infradental pair, which are elongate and widely separated from each other; the middle pair, which are smallest at the base and wider distally; and the wide distal pair, which are opercular. The diamond-shaped oral shields are as long as wide and have broadly rounded angles. The ventral arm plates are pentagonal, having a convex outer side, two lateral sides, and two inner lateral sides. One wide tentacle scale is attached to the ventral arm plate, and the second smaller scale lies against the side arm plate. The two are joined at right angles to each other. The slightly overlapping dorsal arm plates are twice as wide as long and roughly elliptical. Proximally there are four arm spines as far as the tenth to seventeenth arm segment, where the spines are reduced to three. Depending upon whether there are four or three arm spines, the third or second spine resp.ectively is flattened and truncate, the distal lateral edges each bearing one, two, or several prominent teeth. Under close examination the entirety of the lateral edges is seen to be finely serrate. The remaining arm spines are round, acute, and somewhat shorter than the toothed spine. Small, distinct scales cover both dorsal and ventral surfaces of the disk. The radial shields are slender, about 3 times longer than wide, and are separated their proximal one-third to one-fourth by one or two narrow scales. The disk and arms are light tan, the arms mottled with darker gray. Type.-Museum of Comparative Zoology, cat. no. 4140. 1962] Thomas: Amphiurid Brittle Stars 659

'. .. . ' .... '. ..." ,' . . -. . , ..... • ==C0

FIGURE 11. Amphipholis pachybactera H. L. Clark, 1918: A, disk and one arm, dorsal view, X 20; B, two jaws and one arm, ventral view, X 60; C, arm spine, X 100. 660 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) Type locality.-Sandy Point, Buccoo Bay, Tobago, British West Indies. Distribution.-Off Little River, South Carolina; Biscayne Bay, Fla.; St. John, V.I.; and Buccoo Bay, Tobago, B.W.!. Discussion.-My Florida specimens agree with Clark's holotype and paratypes. Remarks.-Formerly, this species was known only from the type locality. In Biscayne Bay it occurs with Ophiophragmus pulcher, Ophiocnida scabriuscula, and Amphioplus abditus in the roots of marine angiosperms, particularly in Thalassia. I found a very char- acteristic specimen of A. pachybactera in the United States National Museum collections (USNM E.7860) labeled as Amphipholis gracillima. The locality given is off the South Carolina coast, and if this is correct the species extends well northward. A mphipholis gracillima (Stimpson, 1852) Fig. 12 Ophiolepis gracillima Stimpson, 1852, p. 224. Llitken 1859, p. 219. Amphiura gracillima: Lyman, 1865, p. 116. Amphipholis januarii Ljungman, 1866, p. 165. Amphipholis gracillima: Ljungman, 1867, p. 314.-Lyman, 1875, pI. 1, fig. 70; 1882, p. 146.-Koehler, 1914, p. 66.-H. L. Clark, 1915, p. 241, pI. 6, figs. 5, 6; 1933, p. 49.-Dilwyn-John and A. M. Clark, 1954, p. 155, fig. 8. Amphipholis goesii: H. L. Clark, 1933, p. 52. Material studied.-l spec.; North Biscayne Bay, Florida; J. Kneeland McNulty, (date unknown); UMML 41.108.-2 spec.; off Aguadilla, Puerto Rico; FISH HAWK (?) sta. 6055, 1899; USNM 21348.-7 spec.; Coral Harbor, St. John, V. 1.; Thomas, 20 December, 1958; UMML 41.70.-4 spec.; Coral Harbor, St. John, V. 1.; Thomas, ]9 December, ]958; UMML 41.73. Diagnosis.-Three oral papillae, the outermost wide, opercular; disk completely covered with fine scales; radial shields 3 times longer than wide, very slender, joined one-half their length; three similar acute arm spines (four or five near disk); two narrow tentacle scales joined at right angles. Description.-One dried specimen measured 5.5 mm across the disk. None of the arms were intact, but originally they must have ,extended to 20 times the disk diameter. There are three pairs of oral papillae. The infradentals are small, block-like, and equal to the middle pair. The distal pair are opercular, wide at their proximal end and tapering to a point at their distal end; the whole papilla forms an acute isosceles triangle with the base proximal. 19621 Thomas: Amphiurid Brittle Stars 661

FIGURE 12. Amphipholis gracillima (Stimpson, 1852): A, disk and one arm, dorsal view, X 16; B, two jaws and one arm, ventral view, X 40. 662 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) The oral shields are longer than wide and roughly diamond-shaped, with the proximal end bluntly pointed and the distal end roundly lobed. The ventral arm plates, about as wide as long, are pentagonal, with an outer side, lateral sides, and inner lateral sides. Those plates close to the disk may also bear an inner side, thus becoming hexagonal. There are two tentacle scales, a slender scale which extends the full length of the lateral side of the ventral arm plate and, at right angles to it, a second smaller scale attached to the side arm plate. The dorsal arm plates are a little more than half as long as wide. Their outer side is slightly convex, the inner side pronouncedly convex. Of the three (four or five near the disk) slender arm spines, the second is the largest. All are round and acute. Both dorsal and ventral surfaces are covered by fine s.cales. There is no trace of the primitiv,e plates. The radial shields are very long and narrow. In the specimen with the disk 5.5 mm in diameter they were between 1.8 and 1.9 mm long and 0.3 mm wide, the ratio of length to width being about 5 or 6 to 1. The disk and arms are tan, the arms indistinctly banded with light brown. Type.-Not traced. Type locality.-South Carolina. Distribution.-South Carolina; Florida; Bermuda; St. Thomas; To- bago, B.W.I.; British Honduras; and Brazil. Remarks.-This species is not common in Biscayne Bay, Florida. At St. Thomas it is very common with Ophionephthys limicola. It easily may be mistaken for A mphioplus at first glance. H. L. Clark (1933, p. 52) indicated that A. gracillima is synonymous with A. goesii Ljungman. Amphipholis squamata (Delle Chiaje, 1829) Fig. 13 Asterias squamata Delle Chiaje, 1829, p. 74. Ophiocoma neglecta; Forbes, 1841, p. 30, text fig. Ophiolepis squamata; Muller and Trosche1, 1842, p. 92. Amphipholis squamata; Verrill, 1899a, p. 312.-Coe, 19]2, p. 81, pl. 17.- Koehler, 1914, p. 66.-Clark, 1933, p. 50. Material studied.-3 spec.; Lake Worth, Florida; Thomas, April 11, 1959; UMML 41.106.-4 spec.; Woods Hole, Massachusetts; 28 July, 1943; UMML 41.105.-2 spec. (as Amphipholis goesii); Mayaguez Harbor, Puerto Rico; FISH HAWK sta. 6061, 1899; USNM 21353. Diagnosis.-Oral papillae three pairs, distal pair opercular; oral shield 1962] Thomas: Amphiurid Brittle Stars 663 ~ ..

FIGURE 13. Amphipholis squamata (Delle Chiaje, 1829): Upper.-Disk and one arm, dorsal view, X 27; Lower.- Two jaws and one arm, ventral view, X 48. 664 Bulletin of Marine Science of the Gulf and Caribbean r12 (4) as wide or wider than long; oral arm plates lack inner side, bear two converging inner lateral sides instead; tentacle scales two, large, joined; arm spines three; dorsal arm plates not overlapping; arms short, four or five times disk diameter; disk s.caleslarge; radial shields joined, total length about 2 times width, distal end white. Description.-The disk of the largest Florida specimen is 2.8 mm in diameter. The arms vary from 10 to 11 mm in length. There are three contiguous pairs of oral papillae, the two proximal pairs similar in size and shape, the distal pair long and opercular. The infradental papillae are slightly or not at all separated. The oral shield is roughly diamond-shaped and as wide as, or slightly wider than, long. The adoral shields are large, joined at their proximal ends, and bear the opercular distal oral papillae. The ventral arm plates, widely separated by the side arm plates. are somewhat longer than wide and are five-sided, having an outer, two lateral, and two inner lateral sides. There are two large contiguous tentacle scales, one on the lateral side of the ventral arm plate and one on the side arm plate. The side arm plates are large and visibly encircle the entire arm. Each plate bears three short, rounded spines. The dorsal arm plates, widely separated by the joining side arm plates, are slightly wider than long, V-shaped proximally, and slightly convex distally. The disk is round and entirely covered with large, overlapping scales. There is a distinct line of demarcation betw,een the scales of the dorsal and ventral surfaces. The prominent radial shields, joined their entire length, are about twice as long as wide. This species is light gray with the distal tips of the radial shields white. Type.-Not traced. Type locality.-Not traced. Distribution.-Cosmopol itan. Discussion.-This species is perhaps the best-studied member of the family. Hyman (1955) summarized much of the work dealing with it. Remarks.-The three Florida specimens were collected on a small piece of sponge lying on muddy bottom in only a few inches of water. Although none were taken in Biscayne Bay their small size (seldom over 3 mm disk diameter) and secretive habits (Clark, 1933) make them difficult to collect, and they are possibly much more common than indicated. I have seen two specimens, apparently of this species, in which the ventral surface of the disk is spinulose. Both w,ere from 1962] Thomas: Amphiurid Brittle Stars 665 Mayaguez Harbor, Puerto Rico (USNM 21353). Genus Ophiophragmus Lyman, 1865 Ophiophragmus Lyman, 1865, p. 131.-H. L. Clark, 1918, p. 270. Each jaw bears three oral papillae which are commonly contiguous. The dorsal arm plates are usually wider than long. Each side arm plate bears three short arm spines. The disk has a border or fence of erect papillae and the radial shields are partially or entirely joined. All three of the species which Lyman originally placed in this genus (Amphiura wurdemanii Lyman, Amphiura septus Liitken, and Am- ph iura marginata Liitken) have distinct, spinulose papillae bordering the edge of the disk. In the years following, several authors, including Lyman, plae,ed species of Ophiophragmus in other genera. Conversely, Matsumoto (1915) and others described species of A mphiodia as Ophiophragmus. H. L. Clark (1918) revised the genus Ophiophrag- mus, limiting the Western Atlantic species to six. However, due to lack of material for .comparison, Clark's key and descriptions are somewhat incomplete. The present author has endeavored to present a more accurate key, and has included all the Western Atlantic species. As the number of disk papillae, the arm length, and the shape of the radial shields vary considerably in this genus, they ar.e used only as supplementary key characters, secondary to the more diagnostic features such as the shape of arm spines and oral shields, and in some cases, coloration. Type-species.-A mphiura wurdemani Lyman (by subsequent designation: H. L. Clark, 1915, p. 238). Remarks.-All of the species in the following key have been examined by the writer; specimens of Ophiophragmus lutkeni (Ljungman) identified by H. L. Clark wer.e loaned by Dr. Elisabeth Deichmann of the Museum of Comparative Zoology. Later the author was able to visit the Museum and examine all the specimens of Ophiophragmus.

KEY TO THE WESTERN ATLANTIC SPECIES OF Ophiophragmus I. Second arm spines near arm base (under the disk) bear 2 or 3 stout terminal hooks or teeth brachyactis, p. 666 1. Second arm spines lack stout hooks, but may bear many fine spinules which are easily seen in dry specimens 2 2. Second arm spine (distal to the disk) more or less truncate; radial shields 2 or more times longer than wide; oral shields diamond-shaped; broken or entire black line extending the length of dorsal arm surface septus, p. 669 2. Arm spines similar, second truncate, arm not striped, or striped with 666 Bulletin ot Marine Science of the Gulf and Caribbean [12 (4) other than black 3 3. Disk with papillae on ventral surface, marginal disk papillae bluntly rounded, not conical and pointed, color gray, arms indistinctly banded ...... filograneus, p. 677 3. Disk lacking papillae on ventral surface, or if present, arms sharply banded 4 4. Marginal spines acute or conical lutkeni (Not found in Florida.) 4. Marginal spines bluntly rounded, not acute ...... 5 5. Green or red longitudinal stripe and green bands on dorsal arm surface ...... pulcher. p. 671 5. No stripe on arm, basal tentacle scales may be widely separated; arm spines bluntly rounded, color white, arms distinctly banded with black, occasionally spines on ventral surface wurdemani, p. 675 Ophiophragmus brachyactis H. L. Clark, 1915 Fig. 14 Ophiophragmus brachyactis H. L. Clark, 1915, p. 238, pi. 10, figs. 13, 14; 1918, p. 278. Material studied.-l spec.; off the Sea Buoy, Miami, Florida, 37 meters; Rob- ert Work and J. Kneeland McNulty, 25 July, 1957; UMML 41.107.-1 spec.; off Sombrero Light, Florida, 61 meters; MCZ 6683.-1 spec. (holotype); off Sombrero Key, Florida, about 87 meters; W. Stimpson, April 2, 1872; MCZ 1120. Diagnosis.-Three blunt contiguous oral papillae each side of jaw; adoral plates broadly in contact at apex of oral plate, joining first arm plate to form continuous chain of plates around mouth; arm spines very short, middle basal spine toothed at tip; two similar tentacle scales; fence of very short, blunt papillae; dorsal disk scales flat, overlapping but little. Description.-The single UMML specimen has a disk 8.8 mm in diameter and an arm length of about 50 mm. Each jaw bears three pairs of papillae, the closely appressed pairs of distal papillae separated by a small gap from the block-like infradental papillae, which extend far into the oral cavity and are widely separated from each other. The short, thick adoral plates are broadly joined at their proximal angles as well as at the first ventral arm plate; they form a regular circle of plates around the oral frame. The oral plates are roughly diamond-shaped and longer than wide, with an acute proximal angle and a long, rounded distal end. The ventral arm plates are not quite twice as wide as long; although they are superficially r.ectangular, actually they have two inner lateral sides which, with the inner side, may form an almost straight line. The two prominent tentacle scales are equal or subequal and slightly 1962J Thomas: Amphiurid Brittle Stars 667

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B FIGURE 14. Ophiophragmus brachyactis H. L. Clark, 1915: A, disk and one arm, dorsal view, X 10; B, two jaws and one arm, ventral view, X 20. 668 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) separated. Three short, somewhat flattened spines are borne on each side arm plate. The first spine is the shortest, the second longer, and the third the longest, slightly shorter than the length of a dorsal arm plate. The second spines are unique in that those at the base of the arm often bear two or three short, acute teeth (Fig. 14). The dorsal arm plates are 3.5 to 4.0 times wider than long, slightly convex distally and concave proximally. There is often a slight depression in the conv,ex distal side, making a re-entering curve. The outline of the disk edge has five broad scallops and is bordered by a fence containing twenty to twenty-five short, bluntly rounded papillae in each lateral interbrachius. The dorsal disk scales are large and prominent. Those in the center of the disk do not overlap, but lie side by side like tiles on a floor, while those towards the sides imbricate slightly. The radial shields are variable, long and slender in small specimens and round to oval in large specimens. The disk is white, and the arms are blotched with light reddish brown. The oral surface is white. Type.-Museum of Comparative Zoology, cat. no. 1120. Type locality.-Off Sombrero Key, Florida, 87 meters. Distribution.-Off Miami Beach; off Sombrero Key, Fla.; and the Gulf of Mexico (Behr,e, 1950). Discussion.-Clark (1915) apparently did not notice the toothed basal arm spines. This character appears to be a better diagnostic character than the arm length, which is often unobtainable due to breakage. Although Clark mentions the possibility of a relationship with O. wurdemani, I believe that the oral skeleton and tentacle scale structure are much more similar to those of O. pulcher and O. septus than to O. wurdemani. In both the MCZ specimens which I examined, the proximal portions of the radial shields are uncovered, making them longer than wide. There seems to be a good deal of variation in this feature however, for the UMML (41.107) specimen has the proximal portion of the radial shields covered by disk scales so the radial shields are about as wide as long. Clark (1918) states that ". . . no case is known of radial shields becoming ... shorter and wider with age" but, as the width does not increase in the present case, the change of shape is probably not unique. 1962] Thomas: Amphiurid Brittle Stars 669 Remarks.-Since this species has been tak.en in relatively shallow water (37 meters) there is a good chance that it will appear in even shallower bay water. Behre (1950) records O. brachyactis from the Gulf of M.exico (near Louisiana?) but her specimens are no longer extant (Dr. H. J. Bennet, Louisiana State University, personal communication). A specimen from Brazil labeled O. brachyactis in the National Museum collection (USNM 39784) is not typical. It is very close to A mphiodia trychna in appearance except for the fence of papillae. The second specimen in a lot of two from Panama (USNM 15406) labeled A mphiodia riisei is also very close to O. brachyactis but lacks a disk, making positive identification impossible. Ophiophragmus septus (Li.itk.en, 1859) Fig. 15 A mphillra septa Liitken, 1859, p. 120. Ophiophragmlls septlls: Lyman, 1865, p. 132.-H. L. Clark, 1915, p. 239; 1918, p. 275; 1933, p. 48. AmphipllOlis septa: Liitken, 1872, pi. 2, figs. 3a, 3b. Amphiodia erecta Koehler, 1914, p. 67, pi. 6, figs. 4-7. Material stlldied.-1 spec.; Cape La Vela, Colombia, 39-41 meters, 1939; Allan Hancock Foundation.-l spec.; off Miami Beach, Florida, 32 meters; 2 July, 1958. UMML 41.87.-2 spec.; off Cape Hatteras, North Carolina, 100 meters, 29 February, 1884; ALBATROSS sta. 2136. USNM 32295, (the larger specimen is the type of Amphiodia erecta Koehler).-1 spec.; Buccoo Bay, Tobago, B.W.I.; H. L. Clark; MCZ 4135. Diagnosis.-Oral papillae three, blunt, contiguous; oral shields more or less rectangular; disk with a border of blunt papillae; radial shields 2 to 2.5 times longer than wide; complete or broken black line on dorsal arm surface, similar line may be present on ventral surface; two similar tentacle scales. Description.-Although the arms are not entire, a specimen with the disk 8 mm in diameter has an arm length of about 63 mm. There are three contiguous, thick oral papillae on each side of a jaw; the infradental which is block-like; the triangular distal one; and a small papilla lying between the apical and distal ones. The distal papilla, arising from the adoral plate, is the largest. The two proximal papillae ar,e borne on a rather small half-jaw. The adoral plates form an acute isosceles triangle with their apices meeting at the proximal end of the diamond-shaped oral shields. The ventral arm plates are wider than long, rather flat, and bear one or two equal or slightly subequal contiguous tentacle scales. These plates are pentagonal with straight outer, lateral, inner lateral, and inner sides connected by rounded corners. The side arm plates are 670 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

arm, 1962] Thomas: Amphiurid Brittle Stars 671 fairly stout and bear three equally heavy, flattened spines. The middle spine is shortest and does not have a bluntly rounded tip as the first and third spines, but instead its tip is angular proximad and rounded distad. This gives it the appearance of being bent back toward the disk. The dorsal arm plates form a rectangle, twice as wide as long, with evenly rounded corners. The disk scales are all about equal in size, and primitive plates mayor may not be pr,esent. The radial shields, 2.0 to 2.5 times longer than wide, are joined for two-thirds of their length. They are separated along the proximal third by two or three small scales. A margin of blunt papillae extends around the disk and, to some extent, onto the ventral disk surface at each radial shield pair. The .color of this species is striking. In the Colombian specimen, which is dried and twenty years old, the disk is light tan, and the arms are tan with several areas of light gray entirely or half covering the dorsal arm plate. The spines are light. Most distinctive is the single dark line extending the entire length of the dorsal surface of the arms. A similar, though fainter, line extends over most of the ventral surface. The Florida specimen is much lighter, greenish-brown in color instead of tan on the dorsal arm surface, and the dark arm line is broken dorsally and nonexistent ventrally. The black dorsal arm stripe is almost completely faded in the USNM specimens. The MCZ specimen has a prominent black line on the ventral arm surface. Type.-Zoological Museum, Copenhagen(?) Type locality.-St. Thomas, V.I. Distribution.-Cape Hatteras; Florida; St. Thomas; Tobago; Cape La Vela, Colombia. Discussion.-This species may be distinguished from all other species of Ophiophragmus by the shape and size of the middle arm spine and the black dorsal arm line. The side arm plates are stout and not extended as in O. wurdemani, nor are the tentacle scales separated. O. septus is similar to O. brachyactis and O. pulchella in having contiguous, thick oral papillae, and in the structure of the tentacle scales and side arm plates. Remarks.-Although I have not taken this species from shallow water, it apparently occurs there in the West Indies (H. L. Clark, 1918, 1933). O. septus is another species which does not seem to app,ear in shallow water along the east coast of the United States, although it has been taken in deeper water. The color is variable and it is possible 672 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) that specimens lacking the dorsal line may be taken. The Florida specimen has this line quite reduced and broken. Ophiophragmus pulcher H. L. Clark, 1918 Fig. 16, 17 Ophiophragmus pulcher H. L. Clark, 1918, p. 274, pI. 8, fig. 1; 1933, p. 4R. -Engel, 1939, p. 8. Amphiodia rhabdota H. L. Clark, 1918, p. 288, pI. 8, fig. 4; 1933, p. 53. Material studied.-l spec.; Bear Cut, Biscayne Bay, Florida: Thomas, 22 January, 1958; UMML 41.90.-1 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 12 July, J 958; UMML 41.91.-2 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 22 January, 1958; UMML 41.92.-2 spec.; S. of Virginia Key, Florida; Thomas, August 1958; UMML 41.93.-1 spec. (holotype); near Loggerhead Key, Dry Tortugas; H. L. Clark, June, 1917; MCZ 4210.-4 spec. (paratypes); Bird Key, Dry Tortugas, Florida; MCZ 4215.-1 spec. (holotvpe of A mphiodia rhabdota); Bush Key, Dry Tortugas, Florida, June 1917; MCZ 4216. Diagnosis.-Oral papillae three pairs; papillae of fence on dorsal surface blunt, often widely spaced; ventral interbrachius covered with large well-formed scales; tentacle scales two; arms banded with green, and a green or red stripe the length of dorsal arm surface. Description.-The arms of a specimen with the disk 7 mm in diameter are from 70 to 80 mm long. Each jaw possesses three pairs of oral papillae, the block-like infradental pair which are separated from each other, the oval second pair, and a triangular or flattened distal pair. The three papillae of .each half-jaw are contiguous. The rhomboidal oral shields are prominent, somewhat longer than wide, and are bluntly lobed distally. Proximally they vary from bluntly rounded to sharply rounded. The ventral arm plates are slightly wider than long, and hexagonal, having short inner lateral and inner sides. One of the two overlapping tentacle scales arises from the ventral arm plate, the other from the side arm plate. The dorsal arm plates are roughly rectangular and are considerably wider than long. There are three short, blunt spines on each of the rather inconspicuous side arm plates. The disk is entirely covered by fairly large, approximately equal, scales. A fence of six to nine short, stout, white papillae may be present in each interbrachius. However, the fenc,e may be entirely missing or composed of fewer spines than indicated above. There may be an additional papilla at the distal tip of each radial shield just above the arm. The radial shields are twice as long as wide and are 1962] Thomas: Amphiurid Brittle Stars 673

FIGURE 16. Ophiophragmus pulcher H. L. Clark, 1918: A, disk and one arm, dorsal view, X 10; B, two jaws and one arm, ventral view, X 30. 674 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

FIGURE 17. Amphiodia rhabdota H. L. Clark, 1918, (holotype, MCZ 4216): A, a single jaw, X 30; B, portion of disk and arm, dorsal view, X 20. in contact for half or two-thirds of their length. The uppermost portion is separated by two or more small scales. The color of this species is usually striking. The disk scales are, to a greater or lesser degree, peppered with light green, while the radial shields are light tan or whitish. A faint, five-armed cross of green may thus be formed by the scaled portion of the disk. Every fourth or fifth dorsal arm plate is entirely green or brownish, contrast- ing with the pale yellow-white ground color of the arm and producing a banded effect. There is also a thin green or red dorsal line running the entire length of the arm. It is usually red in young specimens (Clark, 1918). There were traces of red in the specimen which has been described. In alcohol the green coloring is retained for some time, but the red fades immediately. All ranges of intensity exist and occasional specimens may be almost devoid of color. Type.-Museum of Comparative Zoology, cat. no. 4210. Type locality.-The Dry Tortugas, Florida. Distribution.-Biscayne Bay, Florida; the Dry Tortugas; Aruba, Dutch West Indies (?). Discussion.-This species greatly resembles O. septus, but differs in having longer, more pointed arm spines, a somewhat different color pattern, more regular disk scaling, and wider radial shields. It is my opinion that Amphiodia rhabdota H. L. Clark (Fig. 17) is merely Ophiophragmus pulcher with a poorly developed color pattern and 1962] Thomas: A mphiurid Brittle Stars 675 weak fence of papillae. The oral armature falls within the range I have observed for O. pulcher as do the rather slender radial shields. The disk scales and arms spines are typical O. pulcher. There is an indication of a disk fence which Clark does not mention. Remarks.-Ophiophragmus pulcher is one of the most heautiful ophiuroids of the Biscayne Bay region. I found it in large numbers in the flats south of Key Biscayne, where several may be taken in a single handful of Halimeda sediment. Within the Bay it appears to be associated with Amphipholis thrombodes and Amphipholis pachybactera among the roots of Thalassia and Diplanthera. This colorful Ophiophragmus has been recorded only once since Clark (1918) named the species from material taken at the Dry Tortugas. This single record, from Aruba (Engel, 1939), is questionable, as the color given by Engel is atypical. It is interesting to note that young specimens of Ophionereis reticulata may have a color pattern very similar to that of Ophiophragmus pulcher. Ophiophragmus wurdemani (Lyman, 1860) Fig. 18 Amphiura wurdemanii Lyman, 1860, p. 169.-Ljungman, 1871, p. 648. Ophiophragmus wurdemanii: Lyman, 1865, p. 42, pI. 8, figs. 1, 2; 1875, p. 21; 1882, p. 159.-H. L. Clark, 1915, p. 239; 1918, p. 273; 1933, p. 47; (non 0phiophragmus wundermanii, Koehler, 1914, p. 42). Material studied.-l spec.; Beaufort, North Carolina; MCZ 1118.-1 spec. (as Ophiocnida cubana); Fort Macon Beach, Beaufort, North Carolina; USNM E. 5892.-1 spec. (as O. septus); Sandy Point, Tobago; MCZ 4135 (identification uncertain, probably O. wurdemani; location probably incorrect).- 2 spec.; Ft. Myers Beach, Florida; Robert Work and J. Kneeland McNulty, 8 May, 1959; UMML 41.109.-1 spec. (holotype); Charlotte Harbor, Florida: MCZ 1119. Diagnosis.-Three pairs of papillae to each jaw, infradentals largest, middle pair often spiniform, distal pair variab~e. sometimes rounded; ventral arm plates wider than long, distal edge of basal plates undulate; side arm plates project laterally, two tentacle scales often widely separated; disk entirely scaled, encircled by a fence of bluntly rounded papillae; radial shields large, longer than wide; color white, arms banded with dark brown or black. Description.-There are three oral papillae on each half jaw; the infradental, which is largest and contiguous with its mate; the long blunt or spiniform middle papilla which is usually separated from the infradental; and the distal one, which is often rounded. The small oral shields are variable, longer than wide, rounded proximally and distally, and widest proximally. There is a prominent 676 [12 (4)

B FIGURE 18. Ophiophragmus wurdemani (Lyman, 1860): A, disk and one arm, dorsal view, X 9; B, two jaws and one arm, ventral view, X 25. 1962] Thomas: Amphiurid Brittle Stars 677 pit at the proximal end of the oral shields betwe.en the two half-jaws. The adoral shields are comparatively large, about as long as the oral shields, and concave proximally. The ventral arm plates are wider than long and often have a prominent undulate distal edge. The first eight plates may be swollen laterally. The side arm plates are produced laterally to a considerable degree and bear one of the two tentacle scales. The tentacle por,e is large, becoming largest toward the proximal end of the arm. There are thr,ee bluntly pointed arm spines which may be acute on the basal portion of the arm under the disk. The dorsal arm plates are about 2.5 to 3.0 times wider than long, and are rounded laterally. The disk is completely scaled and bears a dorsal 'border of short, bluntly rounded papilIae, which are usually noncontiguous. Dorsally the scales are prominent and overlap slightly. The radial shields are large, joined almost their entire length, and about 1.5 times longer than wide. They are separated proximally by one or more disk scales. The disks of the dried specimens observed are tan; their arms are tan and banded with darker gray. Both the specimens from Ft. Myers are very light, the arms white, with scattered bands of black. The disk is lightly peppered with light gray. The oral shields are almost black, as are the proximal ventral disk scales and the half-jaws. Type.-Museum of Comparative Zoology, cat. no. 1119. Type locality.-Captiva Island, Florida. Distribution.-The West Coast of Florida and Beaufort, N.C. Discussion.-This species is very similar to O. filograneus in many respects. It differs in usually lacking the ventral disk papillae and in its very light color and prominent arm bands. The proximal arm spines (under the disk) are acute and smooth rather than finely toothed as O. filograneus. H. L. Clark (1918) noticed the striking similarity in other respects, and I must agree with him. A specimen from Beaufort, North Carolina (USNM E.5892) has ventral disk papillae just as O. filograneus. Remarks.-The two specimens of O. wurdemani present in the Marine Laboratory museum were coUected in very fine quartz sand on the sand bar of Ft. Myers beach. This species appears to be the stenoha- line counterpart of O. filograneus. Ophiophragmus filograneus (Lyman, 1875) Fig. 19 Ophiocnida [ilogranea Lyman, 1875, p. 20, figs. 88, 89.-Verrill, 1899, pp. 317, 377.-Koehler, 1914, p. 42.-H. L. Clark, 1915, p. 250. 678 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

FIGURE 19. Ophiophragmus filograneus (Lyman, 1875): A, disk and one arm, dorsal view, X 9; B, two jaws and one arm, ventral view, X 27. 1962] Thomas: Amphiurid Brittle Stars 679 Ophiophragrnus filograneus: H. L. Clark, 1918, p. 274, pI. 2, figs. 4, 5; 1933, p. 47.-Pearson, 1936, p. 70. Material studied.-2 spec.; Clam Slough, Marco Beach, Florida; Thomas, 30 December, 1958; UMML 41.94.-12 spec.; two miles S. of Cutler Power Plant, Biscayne Bay, Florida; Thomas, 31 March, 1958.--4 spec.; Cocoplum Beach, Florida; 4 February, 1936; UMML 41.19.-2 spec.; Lake Worth, Florida; Thomas, 11 April, 1959; UMML 41.104.-27 spec.; Estero Bay, Ft. Myers, Florida; Thomas, 12 April, 1959.-6 spec.; Indian River at Melbourne, Florida; Thomas, 11 April, 1960. A large number of specimens from east Whitewater Bay, Florida were collected and studied during July to December 1958. Diagnosis.-Oral papillae three; fence on dorsal surface with 20 to 40 small bluntly rounded papillae in each interbrachius; scales of dorsal surface large, prominent; scales of ventral interbrachius covered by conical granules; three bluntly rounded arm spines; proximal ventral arm plates laterally swollen; proximal second arm spines often with many fine teeth. Description.-A large specimen has a disk 9 mm in diameter and arms 87.5 mm long. Each side of a jaw possesses three oral papillae, of which the infradental is block-like and largest; the middle is the smallest, higher than wide, often acute; and the outermost is blunt and usually wider than high. The diamond shaped oral shields are longer than wide, with their distal angle drawn out into a lobe. At the inner angle of each oral shield and between the two half-jaws is a prominent pit. The ventral arm plates are pentagonal, with two inner lateral sides which form a rather obtuse inner angle. Near the disk the lateral edges of these plates are swollen, and the plates are more nearly rectangular than pentagonal. There ar.e two tentacle scales at right angles to each other, one borne on the ventral arm plate, one on the side arm plate. Each side arm plate bears three short stout arm spines, the first the shortest, the third the longest. The second spine, at the proximal end of the arm (under the disk) may terminate in three to five or more small acute teeth which are best seen in dried material. The third is bluntly rounded. The upper arm plates are at least 3 times wider than long, slightly concave distally, and convex proximally. The fence of papillae encircling the dorsal disk surface contains an unbroken row of 20 to 40 short, bluntly rounded papillae in each interbrachius. Below this fence the scales of the ventral disk surface are closely covered by papilliform granules. The scales of the dorsal 680 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) surface are large and broadly overlapping. The radial shields are broad, less than twice as long as wide, and in contact almost their entire Length. At their innermost extreme they are divided by one to several scales. The disk is brown or gray, and each scale is edged by a lighter gray or dirty white. The radial shields are gray excepting their outermost ends which are white, and the arms are dark gray, blotched and banded with lighter color. Type.-Museum of Comparative Zoology. Type locality.-Cedar Key, Florida. Distribution.-From Lake Worth on the east coast around the southern tip of Florida and up the west coast to the Tampa-Cedar Key regIon. Discussion.-Clark (1918) noticed the similarity between O. filograneus and O. wurdemani. However, the difference in coloration and the usual absence of oral disk papillae are sufficient to identify O. wurdemani. Remarks.-Previous to Pearson's (1936) record from the Biscayne Bay region, this ophiuroid had been taken only from the west coast of Florida. I have taken O. filograneus from Biscayne Bay to Mel- bourne on the east coast, Ft. Myers and Marco on the w,est coast, and Whitewater Bay at the southern tip of the state. In the Whitewater Bay region I had the opportunity to observe it periodically for five months (Thomas, 1961). Here it lives in shallow water entangled in the roots of a marine angiosperm, Diplanthera wrightii (Ascherson), and may be exposed to salinities below 1OJ{ ( for several days or even weeks at a time. Genus Ophionephthys Ltitken, 1869 Ophionephthys Liitken, 1869, pp. 7, 93.-H. L. Clark, 1918, p. 278. There is a single pair of infradental papillae at the jaw apex and two or three papillae along each half-jaw. The arms are very long with three to five spines on each side arm plate, and bear very small tentacle scales. The aboral surface of the disk is bare except for a patch of scales at the tip of each radial shield and a group of large, flat scales at the edge of the disk. Koehler (1922) suggested suppressing the genus Ophionephthys and placing all the species assigned to that genus (H. L. Clark, 1918, 1938, Mortensen, 1936, etc.) in the genus Amphiura. H. L. Clark 1962] Thomas: Amphiurid Brittle Stars 681 (1918) also mentioned that certain members of the genus differ considerably from the type species. It is likely that many of the species assigned to Ophionephthys do not actually belong in that genus. Type-species.-Ophionephthys limicola Ltitken (by monotypy). Ophionephthys limicola Ltitken, 1869 Fig. 20 Ophionephythys limicola Llitken, 1869, pp. 24, 25, text fig.-H. L. Clark, 1915, p. 239; 1918, p. 280; 1933, p. 46.-Neilson, 1931, p. 266, fig. 8. Material studied. 6 spec.; Coral Harbor, St. John, V.I.; Thomas, 20 December, 1958; UMML 41.68.-1 spec.; W. end of Virginia Key, Biscayne Bay, Florida; Thomas, August 1957; UMML 41.103.-4 spec.; W. end of Virginia Key, Biscayne Bay, Florida; Thomas, 21 June, 1958; UMML 41.102.-1 spec.; Lake Worth, Florida; Thomas, 11 April, 1959.-9 spec.; W. end of Virginia Key, Biscayne Bay, Florida; Thomas, 3 July, 1958. Diagnosis.-Arms very long, 16 to 20 times disk diameter; disk devoid of scales except proximal end of long narrow radial shields and single row in margin of interbrachius; three to five slender, acute arm spines, single small tentacle scal.e; four or five pairs of papillae to each jaw. Description.-The arms are very long, usually 16 to 20 times the diameter of the disk; a length of 250 mm is not uncommon in specimens with the disk 12 mm in diameter. Each jaw bears four or five pairs of papillae; a pair of large infradental papillae, two or three pairs of much smaller triangular papillae which are contiguous with each other but separated from the proximal papillae, and a spiniform oral tentacle scale which is high in the oral groove between the proximal papilla and the three distal papillae. The oral shields are suboval and smallest distally. The adoral shields meet proximal to the oral shields. The ventral arm plates are slightly longer than wide and, near the disk, are more or less rectangular, the outer side being slightly concave. Farther from the disk they become pentagonal, having two inner lateral sides. A single small tentacle scale is borne in the angle formed by the oral and side arm plate. The narrow, bandlike side arm plates usually bear five spines near the disk, then four, and finally three toward the distal end of the arm. The ventral arm plates, slightly wider than long, are distinctly convex distally, and have two straight inner lateral sides. The disk is naked except for a cluster of scales at the proximal end of each radial shield and a single row of large overlapping scales, which extends from the outer distal edge of ,each radial shield to the center of the interbrachial edge of the disk. The radial shields are 682 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

FIGURE 20. Ophionephthys limicola Liitken, 1869: A, disk and one arm, dorsal view, X 9; B, two jaws and one arm, ventral view, X 25. 1962] Thomas: A mphiurid Brittle Stars 683 about 3 times longer than wide and are separated except at the distal end. Fresh or live specimens from Florida waters are strikingly marked. The disk is usually a rather drab olive brown in contrast to the light tan arms. Each of the arms bears a continuous or partly broken line of very dark blue or black its full dorsal length. At irregularly spaced intervals this longitudinal line is crossed or intersected by lateral lines of either the same dark shade or bright red. Often the lateral lines are doubled and very close together. Occasionally specimens are bright pink instead of tan, or lack markings altogether. Type.-Zoological Museum, Copenhagen(?) Type locality.-St. Thomas, Virgin Islands. Distribution.-Lake Worth and Biscayne Bay, Florida; the Dry Tor- tugas; St. Thomas, Virgin Islands. Discussion.-H. L. Clark (1918) took this species in shallow water in the Dry Tortugas. In both his 1918 review of the genus and his 1933 paper he indicates that the mouth parts are similar to those of Amphiura. The jaw papillae do not, however, occur at the distal jaw angle as in Amphiura, but are found on the half-jaws much the same as in Amphioplus. Some of the Pacific forms of the genus are more like Amphiura and probably should be removed from Ophionephthys. Remarks.-This is one of the most common of all the amphiurids of Biscayne Bay. It may be found occasionally with almost all of the other amphiurids in the various habitats, but occurs in very large numbers in soft muddy bottom devoid of macros.copic plant life. There it is usually associated with Amphioplus coniortodes. Exceed- ingly long arms allow O. limicola to live eight or nine inches deep in the mud with just the tips of its arms protruding. It is very active and is difficult to capture intact, almost always casting its disk during or shortly after capture. The color pattern of this species is rather variable. Specimens from S1. John, V.I., are darker, have a transverse black stripe on almost every aboral arm plate, and lack the dorsal line. Genus Ophiocnida Lyman, 1865 Ophiocnida Lyman, 1865, p. 133.-Verrill, 1899a, p. 315 (in part). There are three rather thick contiguous oral papillae. The dorsal surface of the disk has divergent naked radial shields and scales which are covered by short spinules. The arms bear two tentacle scales (rarely one), and three to five 684 Bulletin of Marine Science of the Gulf and Caribbean [12 (4 ) spines on each side arm plate. Type-species.-Ophiolepis hispida Le Conte (by subsequent designation: Verrill, 1899, p. 317). Ophiocnida scabriuscula (Ltitken, 1859) Fig. 21 Amphiura scabriuscula Liitken, 1859, pp. 216, 220, pI. 3, fig. 4, a, b, c. Ophiocnida scabriuscula: Lyman, 1865, p. 135; 1882, p. I55.-Verrill, 1899a, p. 317.-Koehler, 1913, p. 360; 1914, p. 43.-H. L. Clark, 1915, p. 251, pI. 9, figs. 3, 4.-Boone, 1928, p. 14.-H. L. Clark, 1933, p. 55.-Engel, 1939, p. 8.-Fontaine, 1953, p. 201. Material studied. 1 spec.; Bear Cut, Biscayne Bay, Florida, in Thalassia roots, sandy mud; Thomas, 12 July, 1958; UMML 41.88.-1 spec.; Brazil; W. L. Schmitt, 1927; USNM E. 4959.-1 spec.; Brazil; W. L. Schmitt, 1927; USNM E. 4960.-1 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 22 January, 1959; UMML 41.89.-2 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 20 July, 1958. Diagnosis.-Oral papillae three, similar; oral shields diamond-shaped, longer than wide; disk partially covered with short spines; radial shields 3 times longer than wide, naked, separated proximally, bordered by naked scales; arm spines three, short, blunt; tentacle scales two. . Description.-One specimen with the disk 6.5 mm in diameter has a single unbroken arm slightly over 50 mm long. Each side of a jaw bears three oral papillae; a block-like infradental papilla and two smaller truncate papillae, which are .equal or slightly subequal. The members of the infradental pair are separated. The oral shields are diamond shaped and longer than wide, with the distal angle slightly pinched to form a lobe. The adoral shields touch proximally or are separated by only a very small space. The ventral arm plates, particularly near the disk, are rather rectangular and wider than long. Farther from the disk the inner lateral sides appear, making the shields pentagonal. There are two similar, contiguous tentacle scales. The side arm plates bear three bluntly rounded arm spines, which are about as long as the width of the ventral arm plates. The dorsal arm plates are almost 3 times as wide as long and are roughly rectangular, with a trace of a concavity at their outer side. The disk surface is covered by a scattering of short papillae or spines except near the radial shields, where the scales are prominent. The radial shields are about one-fifth as long as the diameter of the disk, and are joined or nearly joined distally and widely separated 1962] Thomas: Amphiurid Brittle Stars 685

FIGURE 21. Ophiocnida scabriuscula (Li.itken, 1859): A, disk and one arm, dorsal view, X 14; B, two jaws and one arm, ventral view, X 28. 686 Bulletin of Marine Science of the Gulf and Caribbean fJ 2 (4) proximally. They are 3 times as long as wide and taper to a blunt point at their proximal end. The color is light golden brown with a touch of yellow or pink in live specimens, fading to dull brown or white in alcoholic specimens. Type.-Zoological Museum, Copenhagen(?) Type locality.-St. Thomas, V.I. Distribution.-From Bermuda, Florida, and the Dry Tortugas to Tobago and Brazil. Discussion.-Llitken's (1859) figure shows an amphiurid which is somewhat more spiny than the specimens that I have observed. The shape of the upper arm plates and two tentacle scales is, however, very similar to that of my specimens, as is the shape of the radial shields. A diskless specimen in the National Museum collection (USNM 4623), identified as Amphiodia planispina, is probably O. scabriuscula. Unlike most of the specimens I have examined, this one has the infradental papillae quite close together. H. L. Clark (1933) separates this species from the similar O. cubana A. H. Clark on the basis of the two tentacle scales and long, slender radial shields. Unfortunately, the holotype of O. cubana is a very young specimen and quite undeveloped in many respects. The adult is more similar to Ophiophragmus and certainly would have been placed in that genus by H. L. Clark. I hope to redescribe O. cubana in the near future. Genus Hemipholis Lyman 1865 (Agassiz MS.) Hemipholis Lyman, 1865 (Agassiz MS.), p. 137. Ther;e are two large round papillae at the distal angle of each jaw. Infradental papillae are not present at the jaw apex. The adoral shields form a continuous ring around the mouth. Each side arm plate bears three short spines. The disk is regularly covered with scales above, naked below, and bears prominent, partially joined radial shields. Fell (1960) has placed Hemipholis in the family Ophiactidae, where it probably belongs. Type-species.-Hemipholis cordifera Lyman (not Bose) = Hemi- pholis elongata (Say) Hemipholis elongata (Say, 1825) Fig. 22 Ophiura elongata Say, 1825, p. 16. Amphiura elongata: Liitken, 1859, pp. 215, 217, pI. 2, fig. la, b, c. 1962] Thomas: Amphiurid Brittle Stars 687 Hemipholis cordi/era: Lyman, 1865, p. 137, pI. 1, figs. 1-3 (non cordi/era Bose, 1830). Hemipholis elongata: Koehler, 19]4, p. 154.-Boone, L., 1933, p. 106, pI. 62.-H. L. Clark, 1933, p. 46; 1939, p. 446. Material studied.-I spec.; May River, South Carolina; USNM 33979.-1 spec.; upper Biscayne Bay, Florida; J. Kneeland McNulty, (date unknown); UMML 41.74.-1 spec.; disk diameter 2.6 mm; Lake Worth, Florida; Thomas, 11 April, ]959; UMML 41.77.-26 spec.; Trinidad; ALBATROSS(sta?), 1884; USNM 7490.-2 spec.; Rio de Janiero, Brazil; von Martens; USNM 17744. Diagnosis.-One pair of oral papillae, very large and flattened; disk with large scales above, naked below; radial shields prominent, as wide as long; tentacle scale single; arm spines three; dorsal arm plates 3 times as wide as long. Description.-The largest Florida specimen studied has a disk 9 mm in diameter. The arms are not intact; howev,er, the longest fragment measures 66 mm, and the total arm length must have been about 75 mm. A single large, rounded tentacle scale lies at the distal angle of each jaw. The oral tentacle pores are v,ery large. There are no infradental papillae at the apex of the jaw. The adoral plates are bluntly lunate; they touch or almost touch each other both at the inner apex of the oral shield and abov,e the first ventral arm plate, so that a continuous border of adoral plates encircles the oral aperture and jaws. The oblong oral plates are much wider than long. The ventral arm plates are roughly rectangular, 1.5 times as wide as long and have broadly rounded comers. The side arm plates bear a single large tentacle scale which guards a very large tentacle pore. The dorsal arm plates are hexagonal, with a broad slightly convex outer side, a very short lateral side, a somewhat longer inner lateral side, and an inner side equaling about one-half of the plate width. The plates ar:e 3 times as wide as long and touch each other. The three fairly slender pointed arm spines are about as long as the length of a dorsal arm plate and are nearly equal. The disk surface is covered by well-defined scales, among which the primitive plates are easily discernible. The scalation ends abruptly at the periphery and the entire ventral surface is usually naked. In small specimens, the naked ventral surface is somewhat papillated. The two radial shields are long,er than wide and tear-drop shaped, separated proximally by a large round scale and distally by one or two long slender ones which extend almost the length of the shields. There may be sev,eral small scales between these separating scales. Distally the shields mayor may not touch. 688 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

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FIGURE 22. Hemipholis elongata (Say, 1825): A, disk and one arm, dorsal view, X 9; B, two jaws and one arm, ventral view, X 20. 1962] Thomas: Amphiurid Brittle Stars 689 The outline of these shield pairs is distinctly cordate, and each pair bears four to six short blunt papillae at its distal end. Every third to sixth dorsal arm plate is dark, almost black. The ground color of the arms and disk is light tan or brown. Because of the dark plates the arms appear banded. Type.-Academy of Natural Science of Philadelphia(?) Type locality.-Charleston, South Carolina. Distribution.-This sp.ecies has been found along the east coast of the United States from South Carolina to Florida, and at Puerto Rico, Trinidad, and Brazil. There are no records for it between Florida and Puerto Rico. Discussion.-Boone's (1933) illustration and description are apparently incorrect, for the specimen is described and figured as having seven arm spines instead of three. It is possible that Boone did not have a Hemipholis, but a species of Ophiactis. Remarks.-The small spe.cimen from Lake Worth (disk diameter 2.8 mm) differs in certain respects from the large specimen from Biscayne Bay. The arm bands are light yellow on a darker yellow background. The radial shields are more widely separated and the shield pairs are consequently not cordate. Finally, the oral disk surface bears a sprinkling of very small, faintly spinulose scales or papillae. These papillae are also present on a rather large specimen from Trinidad (USNM 7490). Genus Ophiostigma Liitken, 1856 Ophiostigma Liltken, 1856, p. 13.-Lyman, 1865, p. 103. There are three pairs of oral papillae of which the distal pair is long and opercular. The adoral plates are thick, forming a ring around the oral aperture. There are three short arm spines. The disk is entirely covered by short papillae which usually extend onto the radial shields. Type-species.-Ophiostigma tenue LUtken (by monotypy). Ophiostigma isacanthum (Say, 1825) Fig. 23 Ophiura isacanthum Say, 1825, p. 150. Ophiocoma isacantha: Milller and Troschel, 1843, p. 103. Ophiostigma moniliforme Liitken, 1859, p. 132. Ophiostigma isacanthum: Ljungman, 1866, p. 317 (as isocanthum).-Ly- man, 1865, p. 103.-Ljungman, 1871, p. 636.-Lyman, 1875, p. 5; 1878, p. 224; 1882, p. 165.-Koehler, 1907, p. 298; 1913, p. 363, pI. 20, figs. 690 Bulletin of Marine Science of the Gulf and Caribbean [12 (4)

FIGURE 23. Ophiostigma isacanthum (Say, 1825): A, disk and one arm, dorsal view, X 16; B, two jaws and one arm, ventral view, X 40. 1962] Thomas: Amphiurid Brittle Stars 691 6,7; 1914, p. 154.-H. L. Clark, 1915, p. 244; 1933, p. 50; 1939, p. 446. -Engel, 1939, p. 4, 8.-Fontaine, 1953, p. 20l. Material studied.-4 spec.; Ragged Key No.3, Biscayne Bay, Florida; Thomas, 18 April, 1959.-1 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 22 January, 1959; UMML 41.81.-1 spec.; off Pines Canal, Key Biscayne, Florida; Robert Work, 21 March, 1958; UMML 41.80.-1 spec.; Bear Cut, Biscayne Bay, Florida; Thomas, 2 April, 1958; UMML 41.82.-1 spec.; Key West, Florida; Hemphill, 1885; USNM 34000 (as Amphipholis sp.).-l spec.; Ensenada de St. Rosa, Cuba; Tomas Barrera Expedition, sta. 7, May 1914; USNM 34782. -4 spec.; off St. Thomas, V. I.; FISHHAWKsta. 6079, 1899; USNM 26660. Diagnosis.-Outermast aral papillae elongate, capable af almo.st completely closing outer half of mouth slit; disk cavered with short blunt spines; no. scales evident dorsally; arms fairly short and stout. Description.-A specimen with an arm length of 17.8 mm to 15.3 mm has a disk 4.5 mm in diameter. Each side af a jaw bears three oral papillae, the distal most elongate and, with its adjacent fellow, capable of almost ,entirely closing off the mauth slit. The infradental and second oral papillae are much smaller than the distal, and the infradental, smallest of the three, is roughly rectangular. The oral shields are shart, slightly wider than long, and rounded on their distal side. The adaral shields are proximally cancave, distally convex, and rounded at each end. The ventral arm plates are rounded distally and as wide as long, becoming much narrower and long.er as they near the arm tips. There are two tentacle scales nearly identical in size and shape. The side arm plates are praminent from both dorsal and ventral views as in Ophiactis. Each side arm plate bears three similar arm spines, all of which are shorter than the lengths of the dorsal arm plates. The dorsal arm plates are wider than lang near the disk, becoming narraw- er and cone-shaped toward the proximal ,end of the arm. No. scales are visible an the disk, there being instead short blunt spines cavering bath darsal and ventral surfaces. The radial shields mayor may not be covered. One or two spines longer than the rest may usually be faund around the interbrachial edge of each radial shield. The disk and arms are light tan, the disk aften mottled with darker colar. Every second or third dorsal arm plate is slightly darker than the shields bardering it, giving the entire arm a faintly banded appearance. In addition to this, each dorsal arm plate possesses a faint oval line of darker color in its center which is joined from one arm plate to the next by a shart dark line, forming a .chain which extends the entire 692 Bulletin of Marine Science of the Gulf and Caribbean [12 (4) length of the arm. This chain is clearly seen in the photograph by Koehler (1913). Type.-Not traced. Type locality.-Coast of Florida. Distribution.-Bermuda; Florida; and the West Indies. Discussion.-H. L. Clark (1933) states that some specimens of isacanthum have six arms. In this respect, as well as in superficial appearance, O. isacanthum looks considerably like an ophiactid. I have found specimens under sponges as well as buried in sandy mud and in the roots of marine angiosperms. Remarks.-This species is common in Biscayne Bay. LITERA TORE CITED BARNARD,J. L. ANDZIESENHENNE,F. C. 1960. Ophiuroid communities of southern California coastal bottoms. Pacif. Nat., 2(2): 131-152. BEHRE,E. H. 1950. Annotated list of the fauna of the Grand Isle region 1829-1946. Occ. Pap. Mar. Lab. La Univ., (6): 1-66. BOONE,L. 1928. Echinodermata from the tropical East America Seas. Bull. Bing. oceanogr. Coli., 1(4): 1-22, Pis. 1-8, Figs. 1-8. 1933. Scientific results of cruises of the yachts EAGLEand ARA ... Coelen- terata, Echinodermata, and MolIusca. Bull. Vanderbilt oceanogr. (Mar.) Mus., 4: 1-217, Pis. 1-133. Bose, L. A. G. 1830. Histoire Naturelle des Verso Suites a Buffon. II. 1830, p. 138. CLARK,A. H. 1921. Report on the ophiurans collected by the Brabados-Antigua expedition ... in 1918. Stud. nat. Hist. Ia Oniv., 9(5): 29-64. CLARK,A. M. 1953. A revision of the genus Ophionereis (Echinodermata, Ophiuroidea). Proc. zool. Soc. Lond., 123(1): 65-94, PIs. 1-3, Figs. 1-12. 1955. Echinodermata of the Gold Coast. J. W. Mr. sci. Ass., 1 (2): 16-56. CLARK,H. L. 1901. The echinoderms of Puerto Rico. Bull. U. S. Fish. Comm., 20 (2): 231-263, PIs. 14-17. 1915. Catalogue of recent ophiurans based on the collection of the Museum of Comparative Zoology. Mem. Mus. camp. Zool. Harv., 25 (4): 165-376, Pis. 1-20. 1918. Brittle stars, new and old. Bull. Mus. compo Zool. Harv., 62 (6): 265-338, Pis. 1-8. 1919. The distribution of the littoral echinoderms of the West Indies. Publ. Carneg. Instn., (281): 49-74. 1933. A handbook of the littoral echinoderms of Porto Rico and the other West Indian Islands. Sci. Surv. P. R. 16 (1): 1-141, PIs. 1-7. 1938. Echinoderms from Australia, an account of collections made in 1929 and 1932. Mem. Mus. compo Zool. Harv., 55: 1-596, PIs. 1-28, 64 text figures. 1962] Thomas: Amphiurid Brittle Stars 693 COE, W.R. 1912. Echinoderms of Connecticut. Bull. Conn. geol. nat. Hist. Surv., (19): 1-152, PIs. 1-32, Figs. 1-28. DELLE CHIAJE, S. 1829. Memoria sulla storia et notomia degli animali senze vertebre del regno di Napoli., 5. DILWYN-JOHN, D. AND CLARK, A. M. 1954. The ROSAURA expedition, 3. The Echinodermata. Bull. Brit. Mus. (nat. Hist.), 2 (6): 139-]62,6 PIs. ENGEL, E. 1939. Echinoderms from Aruba, Curacao, Bonaire, and northern Venezuela. Capita Zoologica, 8 (4). FELL, H. B 1960. Synoptic keys to the genera of Ophiuroidea. Publ. Victoria Mus. Wellington, (26): 1-44, text figures. 1962. A revision of the major genera of amphiurid Ophiuroidea. Trans. roy. Soc. N. Z. (Zool.) , 2 (1): 1-26, pI. 1. FONTAINE, A. 1953. Shallow-water echinoderms of Jamaica. Part 2. The brittlestars (Class Ophiuroidea). Nat. Hist. Notes (nat. Hist. Soc. Jamaica), (61): 3-9, 7 figs. FORBES, E. ]840- A history of British starfish and other animals of the class Echinoder- ]841. mata. London, John Van Voorst. 267 pp., text figures. 1843. On the radiata of the eastern Medeterranean. Part 1. Ophiuridae. Trans. zool. Soc. Lond., 19: 143-153, Pis. 1, 2. HYMAN, L. 1955. The Invertebrata, Vol. 4, The Echinoderms. New York, McGraw- Hill Book Co. KOEHLER, R. 1907. Revision des ophiures du Museum de Paris. Bull. Sci. Fr. Belg., 44: 279-351. 19]3. Ophiures. Zool. Jahrb. Suppl., 11 (3): 351-380, PIs. 20, 21. 1914. A contribution to the study of Ophiurans of the United States Na- tional Museum. Bull. U. S. nat. Mus., 84: 1-173, PIs. 1-18. 1922. Ophiurans of the Philippine Seas and adjacent waters. Bull. U. S. nat. Mus. 100,5: 1-186, PIs. 1-103. LJUNGMAN, A. 1866. Ophiuroidea viventia hue usque eognita. Ofvers. K. Vet. Akad. Forh., (9): 303-336. 1867. Om Nagra nya arter af Ophiuroider. Oivers. K. Vet. Akad. Forh., (23): 163-166. ] 87 I. Om tvanne nya arter Ophiurider. Ofvers. K. Vet. Akad. Forh., (27): 471-475. LUTKEN, C. F. 1856. Oversigt over de Vestindiske Ophiurer. Vidensk. Medd. dansk. naturh. Foren. Kbn. : ]-26. ] 859. Additamenta ad historiam Ophiuridarum, Anden Afdeling. K. danske. vidensk. Selsk., Afd., 8: 24-109, PIs. 1-5. ] 869. Additamenta ad historiam Opbiuridarum, Tredie Afdeling. K. danske. vidensk. Selsk., Afd., 5: 179-271. LYMAN, TH. ] 860. Descriptions on new ophiurans belonging to the Smithsonian Institu- tion and to the Museum of Comparative Zoology at Cambridge. Pree. Boston Soc. nat. Hist., 7: 193-252. 694 Bulletin of Marine Science of the Gulf and Caribbean [J 2 (4) 1865. Ophiuridae and Astrophytidae. lIlus. Cat. Mus. compo Zool. Harv., 1: 1-193, Pis. 1,2. 1869. Preliminary report on the Ophiuroidae and Astrophytidae dredged in deep water between Cuba and the Florida reef ... Bull. Mus. compo Zool. Harv., 1 (10): 309-354. 1875. Zoological results of the Hassler expedition, II. Ophiuridae and Astrophytidae. lIlus. Cat. Mus. compo Zool. Harv., 8: 1-34, PIs. 1-5. 1882. Report on the Ophiuridae. Reports Challenger Zool., 5 (14): 1-386, PIs. 1-48. MARKTANNER-TuRNERETSCHER, G. 1887. Beschreibung neuer Ophiuriden und Bemerkungen zu bekannten. Ann. naturh. (Mus.) Hofmus. Wien., 2: 291-316. MARTENS, E. VON 1867. Ueber vier neue Schlangensterne (Ophiuren) des Kgl. zoologischen Museums. Mber. Akad. Wiss. Berlin, June 1867, (1868?): 345-348. MATSUMOTO, H. 1915. A new classification of the Ophiuroidea: with descriptions of new genera and species. Proc. Acad. nat. Sci., Philad., 67: 43-92. MORTENSEN, TH. 1936. Echinoidea and Ophiuroidea, Discovery Rep., 12: 199-348, PIs. 1-9. MULLER, J. AND TROSCHEL, F. H. 1842. System der Asteriden. Braunschweig, 1842. 12 PIs. NIELSEN, E. 1931. Papers from Dr. Th. Mortensen's Pacific expedition, 1914-16. 59. Ophiurans from the Gulf of Panama, California, and the Strait of Georgia. Vidensk. Medd. naturh. Foren. Kbn.: 241-346, 42 Figs. PEARSON, J. F. W. 1936. Studies of the life zones of marine waters adjacent to Miami, I. The distribution of the Ophiuroidea. Proc. Fla. Acad. ScL, 1: 66-72. SAY, T. 1825. On the species of the Linnaean genus Asterias inhabiting the coast of the United States. J. Acad. nat. Sci. Philad., 5: 141-154. STIMPSON, W 1852. New ophiurans. Proc. Boston Soc. nat. Hist., 4: 224. THOMAS, L. P. 1961. Distribution and salinity to 1era n c e in the amphiurid Brittlestar Ophiophragmus filograneus (Lyman, 1875). Bul!. Mar. Sci. Gulf and Carib., 11 (1): 158-160. THORSON, G. 1957. Bottom communities. Mem. Geo!. Soc. Amer., (67): 461-534. VERRILL, A. E. 1871. Brief contributions to Zoology from the Museum of Yale College. No. 15. Descriptions of starfishes and ophiurans from the Atlantic coasts of America and Africa. Amer. J. Sci., 2: 1-4. 1882. Notice of the remarkable marine fauna occupying the outer banks off the southern coast of New England. No.3. Amer. J. Sci., 23: 135-142. 1899. Report on the Ophiuroidea collected by the Bahama Expedition in 1893. Bull. Labs. nat. Hist. Univ. la. 5 (1): 1-86. 1899a. North American Ophiuroidea. I. Revision of certain families and genera of West Indian ophiurans. II. A faunal catalogue of the known species of West Indian ophiurans. Trans. Conn. Acad. Arts. Sci., 10 (7): 301-385.