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TRANSACTIONS
CONNECTICUT ACADEMY
ARTS AND SCIENCES.
VOLUME X. & 677 as /
NEW HAVEN: Pea etSHED BY THE” AGAD A My 1899-1900.
THE TUTTLE, MOREHOUSE -& TAYLOR CO
OFFICERS OF THE ACADEMY, 1899-1900. President. WILLIAM H. BREWER.
Vice-President.
RUSSELL H. CHITTENDEN.
Secretary.
ALEXANDER -W. EVANS.
—/, ie Librarian. -t.9_ ADDISON VAN NAME. Q . . ¢, Treasurer. \ . MORRIS F. TYLER. * ire
Publishing Committee, GEORGE J. BRUSH. ADDISON E. VERRILL. RUSSELL H. CHITTENDEN. EDWARD 8. DANA. CHARLES A. HASTINGS. CHARLES E. BEECHER. ADDISON VAN NAME. |
Auditing Committee.
ADDISON E. VERRILL. ADDISON VAN NAME.
© ON DeEENS ES:
TAS OWMLADDLTIONS TO THE LIBRARY 22225-25220 525 osee ot eee oe Art. I.—REVISION OF THE NORTH AMERICAN SPECIES OF FRULLA- NIA, A GENUS OF HEPaATICa. By A. W. Evans. Plates 1-15. ART. II.—STUDY OF THE FAMILY PECTINID®, WITH A REVISION OF THE GENERA AND SUBGENERA. By A. E. VERRILL. Plates 16- ART, III.—REVISION OF THE MARINE GASTEROPODS REFERRED TO CYCLOSTREMA, ADEORBIS, VITRINELLA AND RELATED GENERA, WITH DESCRIPTIONS OF “SOME NEW GENERA AND SPECIES BE- LONGING TO THE ATLANTIC FAUNA OF AMERICA. By KATHERINE
ess crammltlalesyoor cos. ne ee se nee ere eee aes ART. IV.—REVISION OF CERTAIN GENERA AND SPECIES OF STAR- FISHES, WITH DESCRIPTIONS OF NEW FORMS. By A. E. VERRILL. JETT PRR SHER Ue ok SS a Rn On ORL RI Nee NE Le Fe ee aS See ART. V.—ON THE DEVELOPMENT OF THE PILIDIUM OF CERTAIN NEMERTHANS. by W.R. Con... Plates 81=35) 22225222222 2.5522 ART. VI.—MATURATION, FERTILIZATION AND EARLY DEVELOPMENT OF THE PLANARIANS. By W. G. VANNAME. Plates 36-41. __ .- ART. VII.—NorTH AMERICAN OPHIUROIDEA. I.—RuEVISION OF CER- TAIN FAMILIES AND GENERA OF WEST INDIAN OPHIURANS. II.— FAUNAL CATALOGUE OF THE KNOWN SPECIES OF WEST INDIAN OpuluRANS: By Av H. VERRILL. Plates 42, 4382-22223 2.222
- Art. VILI.—THE Hawallan HEPATIC OF THE TRIBE JUBULOIDE, Eee bWANS. Plates’ 44-59..-) oe. See ee eo
ArT. [X.—NOTES ON SOME TYPE-SPECIMENS OF MYXOMYCETES IN THE NEW YORK STATE Museum. By W. C. SturGis. Plates
Ni. ils tage cl ee eee eer mei reetege a: Ses Pe ee hs Art. X.—THE AIR-BREATHING MOLLUSCS OF THE BERMUDAS. By LINE WEASEIPSB EY plate: 62.0 525-092 eee Ae eee ArT. XI.—ADDITIONS TO THE ICHTHYOLOGICAL FAUNA OF THE BERMUDAS, FROM THE COLLECTIONS OF THE YALE EXPEDITION Peete yo SAMUEL GARMAN — |...) +.~225. 9300-32) peo ee ArT. XII.— ADDITIONS TO THE MARINE MOLLUSCA OF THE BERMUDAS. By A. &. VERRILL and KATHERINE J. BUSH. Plates 63-65.--- ArT. XIII.—THE NUDIBRANCHS AND NAKED TECTIBRANCHS OF THE BERD as by A. EK. VERRILL.. Plate 66._...--- 22. --2-2 ART. XIV.— ADDITIONS TO THE ANTHOZOA AND HYDROZOA OF THE BERMUDAS. By A. E. VERRILL. Plates 67-69 __...----------- ART. XV.—ADDITIONS TO THE CRUSTACEA AND PYCNOGONIDA OF THE BERMUDAS. By A. E. VERRILL. Plate 70..-.----..-.---- Art. XVI.—ADDITIONS TO THE ECHINODERMS OF THE BERMUDAS. oes Us Plate: 1 Owe. 0) 255 Se ee ee ArT. XVII.—ADDITIONS TO THE TUNICATA AND MOLLUSCOIDEA OF THE BERMUDAS. By A. E. VERRILL. Plate 70__-.---------.-- ArT. XVIII.--ADDITIONS TO THE TURBELLARIA, NEMERTINA AND ANNELIDA OF THE BERMUDAS, WITH REVISIONS OF SOME NEW ENGLAND GENERA AND SPECIES. By A. E. VERRILL. Plate 70.
41
301
387
463
491
595
Per R aly,
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VII.—Norrs American OpnivuromeEa. I.—REVISION OF CERTAIN Famitiges AND GENERA OF West INDIAN OpuiuRANS. II.—A FaunaL CaTALOGUE OF THE KNOWN SPECIES OF WEST InpIAN OpHivurRANs. By A. E. VERRILL.
Part I. Revision of certain Families and Genera of West Indian Ophiurans.
THE numerous shallow water Ophiurans of the West Indian faunal region have been pretty fully studied by several authors,* so that most of the species are fairly well known, and many of them are to be found in most of the larger museums. Nevertheless there is no recent or fairly complete faunal list of the species.
The deep-sea species are also very numerous. These have been collected in large numbers by scientific explorations carried on by the U. 8. Coast Survey Steamer “Blake,” under the supervision of Mr. Alexander Agassiz, and by earlier explorations, under the supervision of Mr. L. F. de Pourtales. A number of deep-sea species were also dredged, in the same region, by the “Challenger.” All the U. 8. Coast Survey collections and those made by the “Chal- lenger” were worked yp and reported upon by Mr. Theodore Lyman in a number of important reports.t
The large collections from this region made by the U. S. Fish Commission steamer “ Albatross” were also studied by Mr. Lyman, but no report upon them has yet been published.
During the present year the writer has published a report{ on a small but interesting collection obtained by a scientific expedition to
*Litken, Addit. ad Hist. Ophiur., Part II; Synop. gen. Ophiur. ver., 1869.
Lyman, North Amer. Ophiuridz, [1l. Catal. Mus. Comp. Zool., I. 1865.
Ljungman, Ophiuroidea viv. hucusque cognita enumerat, Ofvers. Kgl. Veten- skaps-Akad. Forhandlingar, for 1866, 1867..
+ Bulletin of the Mus. Comp. Zoology, Vol. I, No. 10, p. 309, 1869; Vol. V, No. 9, p. 217, 1878; Vol. X, No. 6, 1883; also Vol. V, No. 7, p. 67, 1878, and Vol. VI, No. 2, 1879 (Challenger Coll.).
Iilust. Catal. Mus. Comp. Zool., Vol. VI, 1871; Vol. VIII, No. II, 1875. :
¢ Report on the Ophiuroidea collected by the Bahama Exped., 1893, Nat. Hist. Bulletin, Univ. of Iowa, Vol. V, pp. 1-86, Plates i-viii, 1899.
Trans. Conn. Acap., Vou. X. OctoBER, 1899. 21
302 A. EF. Verrili— North American Ophiuroidea.
the Bahamas and Cuba from the University of Iowa, under the direc- tion of Prof. C. C. Nutting. This collection included only such species as were obtained in less that 260 fathoms.
The present revision and list is based on several collections that I
have studied, but mainly on the following : - I.—The general collections of the Peabody Museum of Yale Uni- versity, in which is included a series of authentically named West Indian species, sent by Dr. Chr. Liitken, from the Museum of Copen- hagen, many years ago.
II.—A pretty full series of deep-sea species dredged by the “Blake” and named by Mr. Lyman, sent by the Museum of Comp. Zoology.
Ill.—The collection made by the Bahama Expedition from the University of Iowa, referred to above.
IV.—The extensive collection made by the U.S. Fish Commission steamers “ Albatross,” “Fishhawk,” and others, under my own supervision, in every year from 1871 to 1887, along the American coast north of Cape Hatteras, and including many deep sea species.
Only a small proportion of those in this last named collection appear to reach the West Indian faunal area, and therefore only a few of the species will be mentioned in this article. A special article on the Ophiurans of the north-eastern coast is, however, well advanced towards completion and will be well illustrated.
In the first part of this paper, I have endeavored to revise some of the larger and more difficult genera and families, and to supply analytical tables, so as to enable students of this group to identify the species without expending such a great amount of time as has been necessary hitherto. The Amphiuride and Ophiacanthide have, therefore, received here more attention than other groups, for they are always the most difficult to deal with.
In this article I have generally used the same names for the organs and parts that were used by Mr. Lyman in his various works on this group, but have made a few changes. I have preferred to use oral shield instead of ‘“‘mouth-shield,” adoral shield instead of “side- mouth-shield,” and oral papille, in place of ‘‘ mouth-papille.” In the genera allied to Amphiura, I have usually called the “ outer mouth- papille ” or papillee of the second oral tentacle, the distal oral tenta- cle-scales to indicate their homology with the ordinary tentacle-scales.
The same idea has been carried out in Ophiacanthide. In the latter ,
group I have designated the apical “mouth-papille” as tooth- papille.
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A. EF. Verrill— North American Ophiuroidea. 308
Class OPHIUROIDEA.
OrpDER I. OPHIURZ: Miiller & Troschel, 1842.
Ophiure Ljungman, Oph. Viv., p. 303, 1867. Verrill, 1899a, p. 4. Ophiuridce Lyman, and many other authors. Zygophiure and Streptophiure Bell, 1892.
Family, PECTINURIDZ Verrill, 1899. Ophiodermatide Ljung., Oph. Viv., p. 87, 1867. Lutk., Addit. Hist. Oph., iii, p. 87, 1869. Pectinuride Verrill, Nat. Hist. Bull. Univ. of Iowa, v, p. 4, 1899a.
The generic name, Ophioderma, is now recognized only as a synonym of Ophiura. Therefore I have changed the name of this family, as is customary in such cases. The name Ophiuride cannot properly be used for the family group here included, because Mr. Lyman and many others have always used it to designate the order Ophiure, or all the Ophiuroidea exclusive of the Huryale.
Family, OPHIOLEPIDZ Ljung., 1866.
Ophiozona nivea Lyman.
Ophiozona nivea Lyman, Ilust. Catal. Mus. Comp. Zool., vol. viii, p. 8, figs. 85-86, 1875; Bull. Mus. Comp. Zool., vol. v, p. 128, 221; Three Cruises of the Blake, ii, p. 110, fig. 390, 1888.
Ophiozona nivea, var. compta Verrill, Nat. Hist. Bull., v, p. 9, pl. iii, fig. 2, 1899. j
Variety, compta Verrill. Prats XLII. Ficures 1, 1a.
The varietal name was given to the variety with distinctly sepa- rated radial shields, regardless of the variations in the oral shields, which happen to be, in both the specimens figured (pl. xxim, figs. 1 and 1a) of the shorter and more ovate form.
A study of a series of specimens sent to me by Mr. Lyman (from sta. 291, 200 fath., Blake Exp.) shows considerable variation in the form of the oral shields. These are sometimes oblong, twice as long as broad, with the outer and inner portions of the same width; in other cases the outer part, beyond the lateral indentations caused by the end of the genital slit, is broader than the inner part; in other specimens the outer part is narrower than the inner. The
304 A. FE. Verrili—North American Ophiuroidea.
number and arrangement of the large angular plates outside the oral shields are variable even on the same specimen. Usually there are three or four of the larger plates, of which two stand side by side, near the margin of the disk.
The radial shields are often separated distally by a row of two or three small angular plates and a large proximal plate as in our figure (pl. xu, fig. 1), but in other specimens the radial shields are in contact distally, but separated proximally by a single large triangu- lar plate, as in Mr. Lyman’s type-specimen of O. nivea. The cen- tral disk-plate is usually closely surrounded by five large angular plates, but in many cases there are small plates intervening more or less irregularly. The variations in the scaling of the disk and in the radial shields are not coincident with the variations of the oral shields.
This species is allied to O. tessellata. It is easily distinguished by the large, irregular disk-plates, wide, oblong, oral shields ; three subequal arm-spines, low down on the sides. There are no marginal spinules outside the radial shields. The upper arm-plates also differ in form.
Off Havana, 110 to 263 fathoms (Bahama Exped.). ‘Taken by the Blake Exped. in 56 to 424 fathoms; off Barbadoes, 200 fath. (Blake Exped.).
Family, OPHIOTHRICHIDZ Ljung.
Ophiothricide Ljung., Oph. Viv., 1866. Ophiothrichide Liitken, Addit., iii, 1869." Verrill, Bahama Exped., p. 18, 1899. Ophiothrichine Ljung., Joseph. Exp., 1871.
The family is characterized by the well defined group of true tooth-papillz ;° by the absence of oral papille; by the usually numerous, long, slender, generally rough and glassy arm-spines; and internally by the complex, interlocking articulations of the arm- bones, and the strong mouth-frames and large radial shields. The peristomial plates, in the typical genera, are in three parts ; of these the middle one is large, like an oral shield. The dental plate or apical jaw-plate is a separate piece.
This family, as now limited, includes the following genera: Ophiothrix, Ophiothela, Ophioenemis, Ophiopsammium, Ophiomaza, Ophiogymna, Ophiocampsis Duncan, Ophiotrichoides Ludw., Ophiop- teron Ludw., Lutkenia Brock, Gymnolophus Brock, Ophiocthiops Brock, Ophiospherea Brock, and Ophiolophus M. Tanner.
A. £. Verrilli— North American Ophiuroidea. 305
The more typical of these genera have the disk-scales covered with slender rough spinules, but the number and length of the spinules may vary considerably, even in the same species of Ophio- thriz. Some of the genera have only granules on the disk-scales, and others have naked scales, and some even smooth skin.
Nearly all the genera and species of this family live clinging closely to various sponges, gorgonian corals, crinoids, hydroids, or even to other ophiuroids. Many of them are more active in their movements than is usual among Ophiuroidea, and many are bright colored when living.
The genus Ophiopteron Ludw. is very remarkable for having a broad membranous web between the arm-spines, and is supposed to be a free-swimming form. It is from Amboina.
The species of this family are mostly found in the warmer seas and in shallow water, and they are most abundant and most diver- sified in the East Indies. Brock enumerated fifty-six species of this family from the Indo-Pacific region and considerable additions have been made to the list by later writers. Several of the genera are known only from the East Indies or Australia. Ophiothriz is the only West Indian genus.
Family, AMPHIURIDZ Lijung., 1867 (emended).
Amphiuride Verrill, Oph. Bahama Exp., Nat. Hist. Bull. Univ. of Iowa, v, p. 23, 1899.
In the report on the Ophiuroidea of the voyage of the Challenger, Mr. Lyman, 1882, recognized about ninety species of Amphiura. In subsequent papers by him and others, about thirty additional species have been described. This very extensive assemblage of species is evidently capable of being divided into several natural groups, in addition to the several minor groups already separated by Mr. Lyman and others. Mr. Ljungman, as long ago as 1867, set off a large number of species as a natural generic group, under the name of Amphipholis. Ata still earlier date, Liitken had indicated this and other natural sections of the genus, without naming them.
Mr. Lyman, however, did not recognize Amphipholis and some other good divisions in any of his works, except as sections of the genus.
The contrast between tle structure of the mouth in typical Am- phipholis and typical Amphiura is very striking. The oral papille in the former can close up the mouth-slits tightly, acting like oper-
306 A, E. Verrill—North American Ophiuroidea.
cula; while in the latter the few slender and widely separated mouth-papillz cannot close the slits, but always leave them widely open. This difference is doubtless directly correlated with impor- tant differences in their mode of feeding and nature of their food.
SUBDIVISIONS OF AMPHIURA.
The species of Amphiura, as adopted by Lyman, mostly fall into four large groups, which seem to be natural divisions of generic value. They are best characterized by the structure, number and arrangement of the mouth parts, as in most other ophiuran families. A few aberrant species, not found in American waters, must be referred to additional groups (V, VI, ete.).
I. Amphiura (restricted). Type, A. Chiajet Forbes.
One apical or subapical oral papilla. One (rarely two) small, distal papilla (oral tentacle-scale) ; middle of jaw-edge without papille; mouth-slits gaping. Four to seven or more (rarely three) arm-spines. Radial shields divergent.
II. Amphipholis (restricted). Type, A. sguamata (or A. elegans.)
Two small lateral oral papille and one broad, operculiform, distal one, forming a continuous series along the entire jaw, and capable of nearly or quite closing the mouth-slits. Radial shields in close contact.
II. Amphiodia Verrill, 1899a. Type, A. pulchella (Lym.).
Three (rarely four) small subequal oral papille, none of them operculiform ; they form a regular series, attached mostly to the side jaw-plate. No distal oral tentacle-scales. Three (rarely four) arm-spines. Radial shields often more or less joined.
IV. Amphioplus Verrill, 1899a. Type, A. twmida (Lym.).
Four or five small unequal oral papilla, 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 three (rarely four). Radial shields generally quite sepa- rated. Disk scales naked.
V. Paramphiura Kehl. Keehler has recently established a new genus, Paramphiura, for A. punctata Forbes and A. bellis, var. tritonis Hoyle.
It is distinguished by having a pair of large supplementary scales or plates, proximal to the adoral plates. There are two small oral papille.
VI. Ctenamphiura Ver., gen. nov. Another special group is represented by A. maxima Lym.
A. FE. Verrill—North American Ophiuroidea. 307
It has three oral papillx in a series, of which the middle one is very large and flat, and the outer one small and spiniform ; the apical one is large. The mouth-shield is so large that it touches the first side-arm plate on each side, while the adoral shields are very small, not meeting within and not embracing the sides of the mouth-shield, as they do in all the other divisions of Amphiura. Two large tentacle-scales. Arm-spines very numerous, ten in the type. Upper and under arm-plates in contact. Disk scales coarse in the type. Radial shields separated.
The type, C. maxima (Lym.), is from the E. Indies in 28 fath.
Amphiura Forbes (restricted sense). Amphiura Forbes, Trans. Linn. Soc., Vol. xix, pp. 149, 150, 1842 (type A. Chiajei). Ljungman, Ophiur. Viv., p. 318, 1867. Amphiura (section B.) Lutken, Addit. Hist. Oph., ii, p. 114, 1859. Amphiura (pars) Lyman, Bull. Mus. Comp. Zool., i, pp. 385, 388; Voy. Chal- lenger, v, pp. 122, 124, 1882.
Amphiura (restr.) Verrill, Ophiur. Bahama Exped., v, p. 24, 1897.
Only one pair of true oral papille to each mouth-slit; they are placed on each side of the apex of the jaw. A single, usually spini- form, papilla, sometimes with a smaller one by its outer side, is situated on each side of the distal end of the mouth-slit, usually attached to the edge of the adoral shield. This is really the outer oral tentacle-scale. —
Owing to the small number of oral papille and their peculiar arrangement, the mouth slits cannot be closed, but appear always gaping, more or less.
The edge of the jaw-plate, along its middle portion, is naked. Higher up in the mouth-slit there is a small spiniform papilla, usually visible from below ; this is the tentacle-scale of the first oral tentacle. It is often shown in published figures as if it were a true oral papilla. Tentacle-scales usually one or two, sometimes lack- ing (section Ophiopelte).
Arm-spines short, usually four to seven or more, rarely three. Radial shields naked, small, generally divergent, with the distal ends either in contact or somewhat separated by small scales. The disk is usually covered with small naked scales.
In one group the under side is without scales (Hemilepis).
In a group referred by Lyman to Ophiocnida, the disk is covered with small spinules, but as the mouth-parts and other organs agree with typical Amphiura, it might better be regarded as a distinct genus, or else as a subgenus of Amphiura. To this Ihave given the name Amphiocnida. (See p. 316.) 5
308 A, E. Verrill—North American Ophiuroidea.
The genus Amphiura, as here adopted, agrees nearly with the typical genus, as restricted by Ljungman in 1867. Mr. Lyman also stated that this should be the typical group, in case the genus were to be divided. This restricted genus still includes over sixty species, with a considerable diversity of structure, as the following table will show. The species are found in all seas and at all depths.
The arms are generally long and slender, tapering very gradually, and very flexible. 3
Many of the species, perhaps nearly all, live buried in the mud and sand of the bottom, or concealed in crevices or under stones, etc. When buried in the mud they usually project the tip of one or more of the arms above the surface of the mud.
This habit of living concealed is doubtless correlated with the absence of disk-spines for protection, and with the lack of special imitative colors.
Many of the species have plain, dull colors, resembling the color of the sea-bottom where they live. A. Otteri, from deep water, is plain salmon or light orange. Such colors are protective in deep water.
Amphiura (restricted): Table of the species inhabiting the West Indies and adjacent waters, and the Atlantic Coast of North America.”
The characters given in this table are those of the adult specimens, or at least of the largest described and figured. The young speci- mens often have fewer arm-spines and differ in other particulars. The number of arm spines given is that of the fully developed joints, towards the base of the arms. The number of spines is also lable to vary in adult specimens. Characters not named in the table, such as the shape of the oral shields, radial shields, arm-plates, etc. are often of more value in determining the species than some of the characters mentioned, and should always be considered. They are not all easily utilized in a condensed table like this.
J.—Disk covered with naked scales.
A.—Tentacle-scales present. Radial shields divergent, their distal ends separated, or scarcely touching. B.—Tentacle-scales two to a pore.
*Tn the table, the species that are entirely northern in their distribution are designated by an asterisk,
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A. E. Verrili— North American Ophiuroidea. 309
a.—Disk covered with scales above and below.
b.—Disk-scales thin and nearly even.
e.—Outer oral papilla flat or squamiform, usually with a small supplementary papilla by its side. Arm-spines 3 to 5, short and stout.
A. incisa Lym., ’83. Arm-spines 3 ;. radial shields wide. Disk-
scales large.
A. Hugenie Liung., 66. Arm-spines 4 or 5. Brazil.
ec.—Outer oral papilla spiniform, prominent. d.—Arm-spines slender, tapered. Radial shield wedge-shaped,
divergent. eé.—Arm-spines 4 or 5; lowest one longest, bent. Disk-scales minute. A. complanata Ljung., 66. Brazil.
ee.—Arm-spines 6 to 8, nearly equal, two or three lower ones usually bent. Disk scales not minute, regular.
A, Otteri Liung., 56. Maine to W. Indies, Portugal.
dd.—Arm-spines 5, short and stout, beaked. Disk-scales minute, obscure beneath. Radial shields narrow, touching distally.
A. Palmeri Lym., ’82. West Indies.
6b.—Disk-seales irregular and swollen. Arm-spines 8. Outer mouth-papilla spiniform.
A. crassipes Ljung., ’66. Brazil.
aa.—Disk naked beneath, or with rudimentary scales only. Radial shields narrow; elongated. Hemilepis Ljung., ’71.
jf.—Arm-spines 4 or 5, stout, subequal. A, semiermis Lym., 69. W. Indies. Jf-—Arm-spines 6, tapered, lowest longest.
A, flexuosa Liung., 66. Brazil.
BB.—Tentacle-scale only one to each pore. Arm-spines 3 to 6.
Radial shields divergent, the distal ends sometimes touch- ing, usually subovate or “ pear-seed shape.” A small sup- plementary papilla or “oral scale” often stands by the side of the outer oral papilla or oral scale.
310 A, E. Verrill—North American Ophiuroidea.
g-—Oral shield transversely elliptical, rhombic, or quadrant- shaped, broader than. long.
h.—Arm-spines 5 or 6, unequal, the lowest longest. Disk-scales minute, not in a rosette. Tentacle scale large, ovate. Oral shield elliptical. A supplementary oral scale is often
present.
A. grandisquama Lym., ’69. W. Indies.
hh.—Arm-spines 3, equal or subequal, tapered. Disk-scales not minute, forming arosette. ‘Tentacle scale small, flat. Oral shield quadrant-shaped. A supplementary oral scale is often present. Radial shields broad, in contact distally.
A, lunaris Lym., ’78. W. Indies.
gg.—Oral shields longer than broad, oblong or ovate. Tentacle- scale minute. Arm-spines 3 to 5, short, subequal. Radial shields small, divergent. . i.—Disk-scales not minute, unequal, with edges rounded and ser- rulate, forming a rosette, oral shield small, subovate. Under arm-plates wide shield-shaped, little longer than broad.
* A, Sundevalli M. and Tr. Gulf of St. Lawrence.
ii. —Disk-scales minute, oral shields rather large, ovate. Under arm-plates oblong, much longer than broad.
A. Stimpsoni Ltk., 59. W. Indies.
AA.—Tentacle-scales absent or rudimentary. j.—Disk destitute of scales below, or only partly covered. Radial shields divergent, pear-seed-shaped or wedge-shaped. = Ophiopelte Sars, Liung.
k.—Arm-spines six or seven, short, straight. Disk-scales minute, not in a rosette.
7.—Arm-spines seven, lowest short and stout, others flattened, subequal. A supplementary outer oral scale. Oral shield
i obtusely angulated or convex distally and proximally. Dorsal arm-plates triangular, with the distal end very con- vex; five middle spines flattened and denticulated.
* A, denticulata Keehl., ’96. Off Newfoundland.
A. E, Verrill— North American Ophiuroidea. 311
7.—Arm-spines six, short. Oral shield acute-angled at both ends. Dorsal arm-plates broad, ovate.
A, Atlantica Liung., 66. Off St. Helena.
kk.—Arm-spines three to five, slender, straight. Radial shields divergent. m.—Arm-spines four or five, short. A rudimentary tentacle-scale sometimes present, usually wanting. “SA. fragilis Verrill, ’85. Arm-spines four or five, subequal, tips rough. U.S. East Coast. mm.—Arm-spines three or four, equal, slender, straight. Radial shields small, touching distally, little divergent. Disk scales small, in rosette. Disk partly naked below.
* A, exigua Verrill, sp. nov. Gulf of St. Lawrence.
jj-—Disk entirely covered with scales below. Radial shields stout, largely joined, not divergent. Arm-spines three or four, equal, straight. Disk-scales rather coarse.
* A. Canadensis Verrill, sp. nov. Gulf of St.- Lawrence.
Il.—Amphiocnida, gen or sub-gen. nov. (see page 316).
Disk-scales bear small acute spinnles. Arm-spines five to ten. Tentacle-scale usually absent. (No American species known.)
AMPHIPHOLIS Ljung. (restr.) Type, A. Januarii Liung.
Amphipholis Ljung., Ofvers. Kongl. Vet. Akad. Férhandl., p. 165, 1866; op. cit., p. 311; op. cit., p. 644, 1871.
Amphiura (pars) Lyman, Illust. Cat. Mus. Comp. Zool., i, p. 115, 1865; Bull. Mus. Comp. Zool., i, pp. 335, 339, 1869; Voy. Chall., v., pp. 122, 125, 1882.
Amphipholis (restr.) Ver., Oph. Bahama Exped., v, p. 24, 1899.
Three or four oral papille form a continuous series along the whole edge of the jaw; of these the distal one is attached more or less to the adoral shield and is operculiform or flat, and often much broader than the others. The two inner are usually small and coni- eal. Disk generally covered with naked scales, but in one species bearing a few spines (Sec. AA). Radial shields naked and usually in close contact along the whole or most of their length. Arm-spines generally three (rarely four), small, slender, tapered. Tentacle-scales one or two, sometimes none.
312 A. E. Verrill—North American Ophiuroidea.
In the more typical species the arms, though slender, are rather short and not very flexible, but in some others they are long, slender and very flexible.*
Table of the species of Amphipholis and Amphiodia from the West Indian region, including Brazil, and from the eastern coast of — North America.
L.—Amphipholis Liung.
Three (rarely four) oral papillae; outer one operculiform or broad and flat, arising partly from the adoral shield. Radial shields joined.
A.—Disk covered with naked scales.
a.—Three arm-spines, rarely four, on basal joints. 6.—Arms of moderate length.
* A. elegans (Leach) Ling. Europe, America. A. tenera (Ltk.) Ljng. W. Indies. A, tenuispina Ling. N. Atlantic. A, limbata (Grube) Ling. Dorsal plates wide, short. Brazil. A, subtilis Ling. Radial shields long and narrow. Brazil.
6b.—Arms long and slender. A, Goési Ling. W. Indies.
aa.—Four or five arm-spines. Arms very long and slender. Radial shields long and narrow. Oral shields large, obovate. Adorals narrow. Outer oral papille very broad.
A. gracillima (Stimp.) Ling. S. Carolina.
AA.—Disk-scales with small, scattered spinules. Two small tentacle- scales. Radial shields in contact for half their length. Three arm-spines.
A. abnormis (Lym. ’78, as Ophiocnida). W. Indies.
* Among foreign species of this genus are the following: <A. squamata, Europe ; A. Torelli, Iceland; A. Pugetana, A. violacea, A. microdiscus, A. Pun- tarene, A. geminata, the last five from the west coast of America; A. Patagonica, Magellan Str.; A. Kochii (Lym.) and A. Corece (Duncan), East Asia. The follow- ing have four oral papille; A. impressa Lj., E. Indies; A. depressa Lj. and A. hastata Lj., from 8. Africa.
A. FE. Verrili—North American. Ophiuroidea. 313
Il.—Amphiodia Verrill, 1899a. (See p. 316.)*
Oral papille three, rarely four, subequal, or the outer one is smallest, forming a regular series on the side of the jaw. Arm- spines three, rarely four. No distal oral tentacle-scale.
B.—Disk-scales naked.
¢.—Two tentacle-scales. d.—Radial shields rather wide, in contact at least distally.
A. Riisei (Ltk.) Ver. Oral shield elongate ; adorals small, trigonal.
A. atra (Stimp.) Ver. Oral shield pelecoidal ; adorals lunate, narrow. (Sometimes has four oral papille and four arm-spines.)
A. planispina (V. Mart.) Ver. Oral shield ovate, broadest prox- imally ; adorals narrow, lunate. Brazil.
ce.—One tentacle-scale. Radial shields long and narrow, largely in contact.
é.—Disk with scales on the under side ; on the upper side larger scales form a rosette. Arm-plates separated above and below. Oral shield obovate, smallest proximally. Adoral shield large, trigonal.
A. pulchella (Lym. ’69) Ver. Florida.
ee.—Disk without scales below; no rosette above. Oral shield “spade-shape,” with a distal lobe. Adoral shield broad triangular.
A. repens (Lym. 775) Ver. Florida.
BB.—Some of the disk-scales, near the margin or beneath, bear small spinules, or granules, or both.t
A. Lutkeni (Ljng.) Ver. West Indies.
* Amphiodia is represented among extralimital species by a large series. Some are as follows. From west coast of America, five: A. Barbare (Lym.), A. grisea (Lj.), A. urtica (Lym.), A. occidentalis (Lym.), A. Chilensis (M. & Tr.), A. Orstedii (Ltk.), A. antarctica (Ljng.), Magellan Str. ; A. jissa (Ltk.), Amoor ; from the Indo-Pacific A. ochroleuca (Brock), A. olivacea (Brock), A. impressa (Ljng.), A. Andree (Ltk.), A. levis (Lym.); from South Africa, A. gibbosa (Ljng.), A. integra (Ling.)
+ Amphipholis Lutkeni Ljng. and Ophiocnida Loveni (Ljng., Lym.) would, perhaps, go here, but they are so closely related to the type of Ophioenida that I have referred to them under that genus (see p. 316).
314 A. E. Verrill—North American Ophiuroidea.
Amphioplus, gen. noy. (See p. 306.) Table of the species of Amphioplus from the West Indian region. C.—Tentacle-scales present.
h.—Two tentacle-scales.
7.—Oral papille four to six, in a series; one or two are distal ont tentacle-scales. Arm-spines usually three, sometimes four.
j.—Dorsal and ventral arm-plates, at base of arms, in contact. Radial shields narrow, separated, or barely touching. Arm- spines three,
A, tumida (Lym. ’78) Ver. Disk swollen ; radial shields linear. W. L., 321 fath. * 4. abdita Verrill, 72. Radial shields lunate, parallel, middle arm- spine stouter, flattened, obtuse. Four oral papille. Long I. Sound. *A. macilenta Ver. Five oral papille. Spines all slender. East Coast U. 8. A. nereis (Lym. ’83) Ver. Oral papille five, thick, unequal ;
genital scale with a row of papille. W.L, 148 fath. A, Agassizii Ver., p. 315. Oral papille six, slender, the two dis- tal ones larger. W. LL, 424 fath.
jj.—Dorsal arm-plates scarcely joined. Radial shields narrow and in contact distally. Five small, bead-like oral papille.
A. cuneata (Lym. ’78) Ver. Three slender arm-spines. W. Indies.
ti.—Four (varying sometimes to three) oral papillae. Arm- spines, three or four. Radial shields a little separated dis- tally, divergent. Oral shield pelecoidal, acute proximally. Disk-scales do not form a rosette.
A, duplicata (Lym. ’75) Ver. First under arm-plate often double; adoral shield narrow. W. Indies.
hh.—One tentacle-scale. Radial shields widely separated. Five unequal oral papillee. A. Stearnsi (Ives) Ver. W. Indies. CC.—No tentacle-scale. Four oral papilla. Four arm-spines. Radial shields touch distally. A. Verrillii (Lym. ’79) Ver. Radial shields rather large, diver- gent. Disk-scales form a rosette. W. Atlantic, 2650 fath.
Many extralimital species of Amphioplus have been described. Among them are the following :
rl
A. E. Verrill—North American Ophiuroidea. 315
A, canescens (Lym.) V., Pacific, 600 fath.; A. glauca (Lym.) V., Pacific, 345-420 fath.; A. cernua (Lym.) V., Pacific, 2300 fath.; A. patula (Lym.) V., Antarctic, 1975 fath.; A. dalea (Lym.) V., S. Atlantic, 2650 fath.; A. /evis (Lym.) V., Philippines.
Amphioplus Agassizii Ver., sp. nov. Amphiura, sp., Lyman, Bull. Mus. Comp. Zool., vol. x, p. 253, pl. v, figs. 64-66, 1883.
Disk covered with minute scales, of nearly uniform size, not form- ing a central rosette. Radial shields narrow, separated by several rows of small scales. Oral shield obovate or pear-shape, evenly rounded distally, longer than broad, sides a little ineurved. Adoral plate long, narrow, three-lobed, not meeting proximally. Oral papille six; of these the four inner ones are small, conical; the two outer are larger and broader and attached to the adoral plate.
Arm-spines three, slender, tapered. Tentacle-scales two, rounded. Under arm-plates are wider than long, broadly in contact and trun- cate at both ends. Upper arm-plates are broadly triangular, short, barely in contact.
This species, which was well figured by Mr. Lyman, but not named, is allied to A. nereis, but the latter has only five oral papille, of which four are stouter and blunter, while the outer one is minute ; its oral shield is rounder; its under arm-plates are barely in contact, and have an inner angle; its arm-spines are larger and its disk-scales are also rather larger.
West Indies, 116 fath., Blake Exp.
Ophiocnida Lyman. Subdivisions.
That this genus, as recognized by Mr. Lyman in his later works, is a heterogeneous group has been noticed by more than one writer. Mr. Lyman, himself, intimated as much in the Voyage of the Chal- lenger. According to his view no difference exists between this genus and Amphiura except that Ophiocnida has spines or grains on the disk. But some of the species have only a few granules, while at least one species that he referred to Amphiura (A. Lutkent) also has some small spinules on the disk, so that this distinction seems to be of little real value, taken by itself.* But as Mr. Lyman
* The same holds good in other cases. Thus Ophiacantha in some species has only granules, and O. levipellis often has naked scales. (See p. 343.) In Ophio- thrix similar variations are found.
316 A. E. Verrill—North American Ophiuroidea.
included in Amphiura four groups that differ in their mouth-parts so widely that I have been led to separate them as genera, he natur- ally admitted the same variations in the mouth-parts of Ophiocnida. In fact we find in this group, as he finally left it, four divisions cor- responding to the four divisions of Amphiura iu structure of the mouth. They should be separated, therefore, if those of Amphiura are to be separated.
When originally constituted the genus included only two species. These are much alike and agree in mouth-parts. They have three subequal, true oral papillae, arranged as in the division of Am- phiura that I have called Amphiodia, to which they are in every way closely allied. It is, in fact, rather doubtful whether these two groups might not be united into a natural genus. In that case the variations in the covering of the disk might be considered as of merely sectional value.
But if we restrict Ophiocnida to the species having the characters of the types, they form a natural and easily recognizable group, which it is weil, so far as known at present, to keep distinct. Mr. Ljungman gave the name Ophioecnidella to this typical group.
The second group, of which 0. Putnami may be taken as the type, agrees with typical Amphiura in its mouth-parts, having but a single true oral-papilla, placed at the tip’of the jaw, on each side, and one or two pairs of oral tentacle-scales at the distal corner. I have been inclined to consider this as a subgenus of Amphiwra, for which I have proposed above (p. 307) the name Amphioenida. It is, at any rate, very closely related to Amphiura. A study of its in- ternal skeletal plates may hereafter show distinctions of more evi- dent generic value.
The third group includes, so far as I know, only O. abnormis Lym. This agrees so completely in its mouth-parts, spines, ete. with typical Amphipholis, that I do not hesitate to unite it with that genus, considering the sparingly spinulose disk as merely of sectional value. (See p. 313.)
Another group, Amphilimna, having O. olivacea as its type, has more numerous oral papillae and arm-spines, and a generally robust structure quite unlike the typical forms. Although corresponding with Amphioplus in the number of oral papille, this group seems to have special characters worthy of generic rank.
The following synopsis will give the principal characters of the three more important divisions discussed above, and of most of the described species :
A. FE. Verrill—North American Ophiuroidea. 317
Ophiocnida Lym., 1865 (restr.). Type O. hispida Lym., 1865. Ophiocnida Lym., Il. Catal. Mus. Comp. Zool., i, p. 138, 1865; pars, Voy. Challenger, p. 152, 1882.
Ophiocnidella Liung., Ofv. Kongl. Vet. Akad. Férhandl., vi, 1871, p. 649 (Type O. scabriuscula).
Oral papiile three, subequal, arranged in a series along the jaw- margin. Disk-scales distinct, bearing spinules or granules. Arm- spines three to five. Radial shields divergent. -Two tentacle-scales ; rarely one.
A.—Disk with numerous acute spinules.
a.—Arr-spines three, rarely four,
O. hispida (LeC.) Lym. 65. Spines three. Disk with many slender sharp spinules. Panama.
O. scabriuscula (Ltk.) Lym. ’65. Spines three. Disk spinose. Disk-seales thick. Oral papille blunt, nearly equal. |W. Indies.
O. echinata Lym. ’74. Spines four. Disk very spinose. Adoral
shields trigonal. Radial shields narrow. E. Indies. O. sexradia Duncan. Six rays; four arm-spines ; one tentacle- scale. K. Indies.
aa.—Arm-spines five or six.
O. scabra Lym. ’79. Lowest spine thick and rough. Proximal oral papilla apical and bead-like. Adoral shield lunate. Off Bahia, 1275 fath.
AA.—Disk-scales partly bare; partly with granules or very short spinules, or both. b.—Disk-scales large, in a rosette, mostly naked; some marginal and submarginal bear granules. Three arm-spines.
O. filogranea Lym. ’75. Radial shields wide, divergent. Adorals lunate. Two tentacle-scales. Florida.
bb.—Disk-scales smaller, many naked; some marginal and sub- marginal bear grains or small conical spinules.
O. Loveni (Ling.). Disk-scales in a rosette; some at margin, bear spinules; others below, bear granules. Outer oral papilla flat. Radial shields touch distally. Rarely four arm-spines. Brazil.
O. Lutkeni (Ling. 71). A few submarginal and marginal scales bear small spinules. Three arm-spines. Dorsal arm-plates wide, usually broken into two or more parts. W. Indies.
Trans. Conn. AcapD., Vou. X. OctToBER, 1899. 22
318 A, E. Verrill—North American Ophiuroidea.
Amphiocnida, gen. nov.=Ophiocnida (pars) Lym.
Disk-seales bear small acute spinules. Apical oral papilla small. Distal oral papilla (oral tentacle-scale) acute, spiniform. Middle of jaw-margin naked. Arm-spines five to ten. Typical species have no tentacle-scale.
A. Puinami (Lym.). Arm-spines nine to ten, stout, upper one clavate. Radial shields separate. No tentacle-scale. Hong Kong.
A, pilosa (Lym.). Arm-spines five to six, slender, tapered. Adoral shields trilobed. Radial shields, little separate. No tentacle-
scale. Bass Straits. A, alboviridis (Brock). Arm-spines five to six. No tentacle-
scales, K. Indies. A. brachiata (Mont.). Arm-spines seven to ten, flattened, one
with an apical cross-piece. Europe.
Amphilimna Verrill. Type, A. olivacea. Amphilimna Verrill, Ophiur. Bahama Exp., v, p. 30, 1899.
Oral papillze four or five in a series. Tooth papille two to four. Arm-spines six to ten, of moderate length. Tentacle-scales usually two, spiniform, one each side of the tentacle-pore. Disk swollen dorsally, with a notch over the base of each arm, and covered with spinules. Radial shields parallel, largely in contact. This genus includes, besides the type, only A. Caribea Ljung.
Amphilimna olivacea Ver.
Ophiocnida olivacea Lyman, Bull. Mus. Comp. Zool., i, 10, p. 340, 1869; Tl. Cat. Mus. Comp. Zool., vi, pl. i, figs. 7, 8; Bull. Mus. Comp. Zool., v, 9, p. 227; op. cit., x, p. 258. Verrill, Amer. Jour. Sci., vol. xxiii, p. 219; Ann. Rep. U. S. Fish Com., vol. x, p. 661; op. cit., vol. xi, p. 549. Lyman, Report Voy. Challenger, Zool., Ophiuroidea, v, p. 156, 1882.
Amphilimna olivacea Ver., Ophiur. Bahama Exped., v, p. 30, 1899.
Puate XLII, Fieurss 1, la.
Arm-spines nine or ten ; oral papillee four or five.
Taken by the U. 8. Fish Commission at numerous stations off the east coast of the United States, from off Martha’s Vineyard to Cape * Hatteras, in 63 to 192 fathoms, and by the “ Blake ” from off Rhode Island to the West Indies, in 40 to 126 fathoms. Off Key West, Florida, 75 to 80 fath. (Bahama Exp.).
A. FE. Verrili— North American Ophiuroidea. 319
A. Caribea (Ljung.) Ver.
Arm-spines six, rough. Oral papille four, the two distal ones squamiform.
It is possible that this species, from the West Indies, 300 to 400 fath., is the young of A. olivacea. In that case the latter name would become a synonym.
Family, OPHIACANTHIDZ Ver.
Ophiacanthine (sub-family of Amphiuride) Ljungman, 1866; Liitken, 1869. Ophiacanthide Verrill, Ophiur. Bahama Exped., Nat. Hist. Bulletin, v, p. 34, 1899.
The family is characterized by the prominent and highly devel- oped side arm-plates, usually meeting above and below, and by the numerous, usually long, and more or less rough spines, which stand out nearly at right angles to the arm. The spines may be solid or hollow, glassy or opaque, terete or flat.
The oral papillz are usually rather numerous and form a continu- ous row along the sides of the jaws, but the outer ones may be of larger size or different in form from the others, or clustered, and in such cases they are really the distal oral tentacle-scales. There may be only a single apical tooth-papilla, or there may be two or three, and sometimes there is a large cluster. The first under arm-plate is usually concave or somewhat bilobed within the mouth-slit, and usually bears two vertical flat processes, which sometimes become movable, like oral papillz.
In some cases the outer oral tentacle-pore is exposed to view on the outer margin of the jaw, and then it has one, or sometimes several, special oral scales or papille by its outer side, or partly surrounding it. Some of its scales may be attached to the adoral plate, or even to the first under arm-plate. This plate is usually concave or somewhat bilobed, and usually bears two inner, lateral, scale-like processes, which are sometimes movable and papilliform like oral papille.
There is generally a single median acute tooth-papilla at the tip of the jaw, but there may be two or three, and in some cases (Ophiocamax, Ophiomitra, Ophiotrema) there may be a cluster of several spiniform tooth-papille. These were counted as oral papille by Mr. Lyman, but when they stand onthe dental plate they should be considered as true tooth-papille.
320 A. E. Verrill—North American Ophiuroidea.
The teeth are stout, flattened, obtuse; they vary from three to eight in number.
The internal structure of the mouth-parts and arms is much like that of some of the Amphiuride. The “ jaw-plate” or dental plate is generally separate from the jaws, and the three parts of the peris- tomial plates are generally distinct, but they are united in Ophioca- Max.
The genera Ophioplax and Ophiolebes are, in several characters, more or less intermediate between the two groups, both externally and internally.
This family, as here understood, includes the following genera : Ophiacantha, Ophiomitra, Ophiotrema, Ophiocamax, Ophiolebes, Ophiothamnus, Ophiocopa, Ophiochiton, Ophiotoma, and probably Ophioblenna. To these I have recently added several others, enumerated below, separated from Ophiacantha, Ophiomitra, and Ophiopsila.
The first six of those named above have the disk covered with scales bearing spinules or thorny processes, or sometimes granules. Ophiochiton and Ophiocopa have naked or nearly naked scales. Ophioblenna and Ophiotoma are covered with naked skin. The radial shields may be large or small, concealed or exposed.
Ophiacantha Mill. and Troschel, 1842. (sens. ext.)
The species of this genus, taken in the extended sense, are very numerous in all seas and are difficult to determine. They are abundant in deep water in northern latitudes, as well as in tropical seas. ‘Ten or eleven species are known off the coasts of New Eng- land and Newfoundland. Several of them occur only at great depths. About twenty species, including two described as new in this article, are known from the West Indian fauna.
This genus is very remarkable for the great variations in the armature of the disk. Some species have only rounded granules ; others well-formed tapered spines; others short, thorny stumps ; others small bifid or trifid spinules or crotchets; while many species have mixtures of two or more of these sorts.
It is probable that these structures have been developed as protec- tive organs, in accordance with the ordinary laws of Natural Selection, and that they are, therefore, directly correlated with the habits of the various species. But the habits of many species are not yet known. I have found several species clinging to gorgonian
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A. FE. Verrill—North American Ophiuroidea. 321
corals, or lodged among their branches; others have occurred on hydroids; certain species, like O. fraterna and O. bidentata, often occur in vast numbers in the dredge where the bottom is composed of broken shells, covered with hydroids, sponges, crinoids, etc., among which they evidently find shelter; some species, as 0. gracilis and O. pentacrinus, cling closely to crinoids.
It seems, therefore, that most of the species live more or less ex- posed to the attacks of fishes and other active enemies, against which a covering of sharp spines would afford some protection. But as fishes avoid coral-animals and hydroids, on account of their stinging powers, it might be expected that those species living among the branches of such organisms would require less protec- tion by spines than those that merely conceal themselves, more or less, among the debris of the sea-bottom. A more careful study of the habits of the shallow-water species may determine, hereafter, whether such differences in habits have determined the evolution of the spines of the disk.
As for the long arm-spines, characteristic of most of the species of this and allied genera, they appear to have been developed in nearly all genera that habitually live exposed,* while those genera that live buried in the mud or sand, like Ophioglypha, Ophiomusium, Amphiura, Amphipholis, etc., or securely hidden in crevices or under stones, generally have short arm-spines.
Some of the species of Ophiacanthaare brilliantly phosphorescent when first caught. I have myself observed this to be the case with O. bidentata, O. fraterna, and others. It may, very likely, be a peculiarity of the deep-water species, if not of all the others.
Owing to the difficulties in the way of the ready identification of the species, I have prepared the following analytical tables, which ought to aid materially in locating any of them, if the specimens be full grown, or nearly so. The young often differ considerably from the adults in the number and roughness of the arm-spines, armature of the disk, ete. The number of arm-spines counted is that of the largest groups, near the base of the arms; farther out the number rapidly decreases. The number of oral papillse often varies with age, and also individually, in many species, especially in those in which they are numerous and clustered. The number of tentacle-scales,
* The genus Ophiothrix is notable for the high development of itsspines. The species usually live more or less exposed, clinging to sponges, gorgonians, etc., which they often closely imitate in color, but some species live in the internal cavities of sponges.
322 A, EF. Verrill—North American Ophiuroidea.
or the number of joints that have two pairs varies in some species according to age, the number of these parts and of the spines in- creasing in the older specimens. In very large specimens of small species there is often a tendency to develop extra oral-papille and tooth-papilli, either above or below the regular series. The precise number of species cannot be considered as constant in any species, and must always be understood to vary within more or less definite limits. This character has been used in some of the analytical tables only because of its easy observation. The degree to which the larger basal rows of spines approximate dorsally is of more impor- tance, though not invariable, and the character of the serrulations or thorns on the spines is of considerable value, though slightly variable, even in adult specimens. The spines are always rougher in the young specimens.
Moreover, in using the analytical tables, it must be remembered that some of the species have been described only from a single specimen* and that the amount of variation is still unknown, in cer- tain deep-sea species, which have not yet been studied from the later and larger collections.
It is also to be noted that in the case of deep-sea species, especially those obtained by the “tangles,” many of the delicate parts are liable to be broken or torn off, and in the case of tentacle-scales and oral papille they may leave no traces. When such parts are reproduced they may not appear in the same number or form as at first.
Such accidents may account for many cases where the different arms or different jaws of a single specimen present variations in their appendages,t as wellas for specimens in which all the arms
*Mr. Lyman’s custom was to describe all his new species from a single type specimen. Had his health remained unimpaired he would, doubtless, have re- vised more fully the large collections from the later Blake Expeditions.
+In a large lot of typical Ophiacantha bidentata one abnormal specimen shows curious variations in the mouth-parts, which may be due to the repair of dam- ages. ‘The number of regular oral papille on the different jaws varies from three to five. Onsome jaws there isa rudimentary, wart-like, distal one ; in others it is as large as the next; on one jaw’ there is an extra, slender, clavate papilla, back of the first, on the lower face of the jaw; the distal papilla is thick, blunt, clavate, and usually somewhat triquetral ; one jaw has two papille grown together for half their length; one has an extra papilla above the inner one, and of the same form, The tooth-papille vary in form and size, and from one to three in number; one jaw has a terminal pair; and on one jaw some of the teeth are split into two. The first arm-plate has a vertical process on
a
A, E. Verrill—North American Ophiuroidea. 323
vary. These ophiurans are able not only to reproduce a whole set of arms, but the entire upper part of the disk itself may be lost and reproduced.
As for the species included in the following tables, I have person- ally studied nearly all of them, and the few that I have not seen are well figured by Mr. Lyman.
Dichotomous analytical table of the East Coast and West Indian species that have been referred to Ophiacantha (sens. ext.)
In this table I have arranged the species as nearly as possible in accordance with what seems to be their natural relations.
Those prefixed by an asterisk (*) are from the American coast north of Cape Hatteras. All others are from the West Indian fauna.
A.—Oral shields join the first side arm-plates. Adoral shields are entirely proximal to the oral shields.
B.—True Ophiacantha. Disk wholly, and radial shields mostly, covered with small crotchets, thorny stumps, or short spin- ules or granules, or with a mixture of these forms.
C.—Disk covered with small crotchets, or short thorny stumps, or short spinules, with no elongated spines nor granules.
d.—Arm-spines finely serrulated, or nearly smooth under a simple lens, usually long and tapered, hollow, not glassy.
€.—Opposite basal rows of arm-spines, in the adults, are closely approximate dorsally or nearly so.
f.—Oral papille form a simple row, the distal one being gener- ally the largest. —
* 0. bidentata(Retz.). Disk with short, thick, rough, obtuse stumps and crotchets. Distal oral papillae wider, truncate. Tentacle-scale single, obtuse.
* 0. aculeata Ver. Disk with slender, thorny, stumps. Distal oral papilla wide, flat, mucronate at the corner. Spines eight or nine, nearly smooth. Tentacle-scale lanceolate, acute. Arm-spines not always approximate dorsally.
each side, not movable. The large outer tentacle-pore is visible from below, when the distal papilla is removed. The first oral tentacle is far up in the distal part of the slit and has no papillae. The uppermost tooth is longer and more pointed than the rest. There may be two clavate tentacle-scales on the first joint.
324 A. E. Verrill—North American Ophiuroidea.
* O. fraterna Ver. When full grown the arm-spines usually closely approximate dorsally (see p. 321).
* O. abyssicola Sars. Spines six, short, nearly smooth. Disk coy- ered with fine crotchets.
* O. anomala Sars. Six arms. Oral shields narrow, or acute. Disk-spinules short, thick, conical or obtuse, roughly serrulate or thorny.
Sf—The distal oral papille, or oral tentacle-scales, are clustered or form a double row; all spiniform, Tentacle-scale spini- form,
* 0. enopla Ver. Arm-spines seven or eight, roughly serrulate. Disk covered with small, short, obtuse stumps, having several termi- nal thorns.
ce.—Basal rows of spines not very closely approximate dorsally. Oral papille in a simple row.
*O. fraterna Ver. Disk covered with very small thorny spinules and crotchets and some rough granules. Oral papille three, acute, spiniform. Arm-spines eight, serrulate. ‘Tentacle-scale small, flat, subacute, Arm-spines, in large specimens, are approximate dorsally.
O. cosmica Lym. Disk with coarse thorny stumps having several points at the end. Oral papille three, stout, conical. Tentacle pores large, with one large scale. Upper arm-plates slightly joined at base of arms.
dd.—Arm-spines decidedly thorny or prickly, and usually glassy, mostly long and slender. g-—Basal rows of spines approximate dorsally. Side arm-plates very prominent. Disk with small, slender crotchets or branched spinules.
O. aspera Lym. Arm-spines nine or ten, slender, very thorny. Disk covered with fine thorny crotchets and stumps, terminated by. two to six points. Tentacle scale single, flat, larger at the end, and thorny or lobed.
*O, millespina Ver, Arm-spines ten, long, roughly serrulate. Disk closely covered with small, thorny or branched spinules.
O. pentacrinus Ltk, Arm-spines six, upper ones very slender, not very thorny. Disk with fine crotchets. Tentacle-scale single, small, flat. Distal oral papilla flat.
i 4 ‘ r
a < s
A. E. Verrill— North American Ophiuroidea. 325
O. scutata Lym. Arm-spines eight to ten, long, decidedly thorny. Basal tentacle-pores with two flat scales; one, and spiniform, farther out. Three tooth-papille. Disk-scales covered with small, thorny crotchets. A pair of papillz on first under arm-plate.
gg-—Basal rows of spines not closely approximate dorsally. Disk with short thorny stumps.
O. stellata Lym. Arm-spines seven, very thorny. Three conical oral papillz. One tooth-papilla.
CC.—Disk entirely covered with tapered spinules or true spines, or having more or less of them mixed with granules or other structures, or else covered with granules only.
h.—Disk covered with spinules only, or else having spinules mixed with other structures, not granulated.
7.—Disk with spinules only or mainly.
j.—Dorsal rows of spines approximate dorsally.
*O. spectabilis Sars. Arm-spines six to eight, serrulate. Disk with tapered spines and some small conical stumps. Tooth-papille and distal oral papille clustered.
jj-—Basal rows of spines not approximate dorsally. k.—Arm-spines finely serrulate, not glassy. Tooth-papilla single. Oral papille in a simple row.
O. segesta Lym. Arm-spines tapered, nearly smooth. Disk-spines small, slender, smooth, mixed with few crotchets and thorny stumps. Tentacle-scale single, small, acute. Oral papille three, conical, all similar.
* O. crassidens Ver. Arm-spines short, stout. Oral papille and teeth large and thick, rough. Disk with small, acute, conical spin- ules.
kk.—Arm-spines thorny and glassy. Disk-spines slender, thorny, acute; several tooth-papille.
O. pectinula Ver. Outer edge of dorsal arm-plates with a row of small acute serrations. Several distal oral papille.
ii.—Disk bearing few tapered spines mixed with other structures. Rows of spines approximate dorsally. 1.—Disk covered with granules mixed with a few tapered spines. Arm-spines finely serrulate or nearly smooth.
326 A, EF. Verrill—North American Ophiuroidea.
O. vepratica Lym. Arm-spines eight, long, tapered. Oral papilla three, conical. 'Tentacle-scale single, large, conical or spiniform.
ii.—Disk-spines elongated, mixed with crotchets or thorny stumps. Arm-spines more or less finely serrulate.
O. varispina Ver. Arm-spines eight, serrulate, translucent. Disk with thorny stumps and few acute spines. ‘Tentacle-scale single, flat, subspatulate. Oral papille wide, flat, obtuse; distally there is often an extra marginal one.
hh.—Disk covered with small close granules alone. Basal rows of spines not approximate dorsally. Arm-spines serrulate ; under arm-plates short and broad, separated.
*O. granulifera Ver. Arm-spines nine, the upper ones long and slender, finely serrulate, lower ones short, rough. Oral papille all spiniform. Tentacle-scale lanceolate, two on first joints.
BB.—Radial shields largely uncovered. Disk-scales either partially naked and easily visible, but bearing more or less granules or spinules, or else entirely concealed.
m.—Disk-scales largely exposed.
n.— Ophialecea Ver., 1899a, pp. 38, 42. Dorsal arm-plates largely in contact. Arm-spines nearly smooth, the rows widely separated dorsally.
O. Nuttingii Ver. Arm-spines four, short, tapered. Disk-scales small, exposed, bearing small spinules. Radial shields partly exposed, narrow, separated. Tentacle-scales single, large.
nn.— Ophiomitrella Ver., 1899a, pp. 39, 48. Dorsal arm-plates separated by the side-plates. Arm-spines slender, thorny; the basal rows approximate dorsally.
O. levipellis (Lym.) Disk-scales small, sometimes entirely naked, sometimes with small scattered granules. Radial shields small, sep- arate, partly naked. Tentacle-scale single, small, acute. A pair of papille on the first under arm-plate.
mm.—Disk-scales mostly concealed, but radial shields naked.
0.— Ophiacanthella Ver., 1899a, p. 39. Basal rows of spines not approximate dorsally. Dorsal arm-plates largely in con- tact. Hadial shields long, mostly naked, in contact by their edges, Arm-spines nearly smooth. Three tooth-papille. Oral papillee four, conical, all similar.
A. EF. Verrill— North American Ophiuroidea. 327
O. Troscheli (Lym.) Arm-spines six, tapered. Disk-scales con- cealed, bearing granules and scattered spines. Tentacle-scale single, lanceolate.
00.— Ophioscalus Ver., 18994, pp. 39, 42. Dorsal arm-plates sepa- rated. Basal rows of spines closely approximate dorsally. Radial shields large, broad, naked, in contact for their whole length. Two or three tooth-papille. Arm-spines thorny and glassy.
O. echinulata Lym, Disk-scales small, nearly concealed by numer- ous slender, thorny spines.
AA,—The oral shield is separated from the side arm-plates by the distal lobe of the elongated adoral shields, which are there- fore, not entirely proximal to the oral shields.
D,.—Adoral shields narrow, trilobed, the narrow distal lobe separat- ing the oral shield from the side arm-plate. Disk-scales usually concealed by cuticle and spinules.
E.— Ophiopora Ver., 1899a, p. 43. No tentacle-scales, the pores are very large; spines small, usually smooth.
O. Bartletti (Lym.) Ver. One spiniform distal oral papilla by the side of the oral tentacle-pore. Disk covered with acute spinules.
EE.—One or two tentacle-scales. p.—Ophiolimna Verrill, 1899¢. Arm-spines seven or eight, nearly smooth, placed obliquely on the distal part of the plates, not strongly divaricate. Jaws more or less granu- lated. Disk-scales and radial shields concealed, bearing granules and spines.
*O, Bairdii (Lym.) Ver. Upper arm-plates separated. Rows of spines approximate dorsally. Tentacle-scale single.
O. mixta (Lym.) Ver. Upper arm-plates joined. Rows of spines wide apart dorsally. Two flat tentacle-scales.
pp.— Ophiopristis Ver.,1899a. Arm-spines serrulate, not obliquely placed. Strongly divaricate. Dorsal arm-plates separated. Tooth-papille usually three. g-—Spines partly flattened, serrulate on the edges. A row or cluster of several distal oral papillz at the large oral tenta- cle-pore. Two tentacle-scales on the basal joints.
328 A. E. Verrill—North American Ophiuroidea.
O. hirsuta (Lym.) Ver. Disk-spines slender, tapered, acute. Arm-spines five or six, strongly serrate on the edges. Three tooth- papille. Two flat tentacle-scales.
O. ensifera Ver. Disk-scales visible, bearing small conical spinules. Spines four, blunt, mostly flat. Two flat tentacle-scales on the basal joints.
O. cervicornis (Lym.) Ver. Disk with granules and small acute spinules. Tentacle-scales two, spiniform; pores very large, open. Arm-spines six, short, flat, serrate.
qq.— Ophiotreta Ver., 1899a, p. 40. Only one or two, rarely three, oral tentacle papille, which are flat. Two to four or more tooth-papille. Arm-spines terete or only a little flattened, slender, serrulate or nearly smooth.
O. lineolata (Lym.) Ver. Arm-spines six or seven, slender, nearly smooth. Tooth-papille three to five. Two unequal tentacle-scales on several basal joints. Disk evenly granulated, and with a few scattered spines. Jaws often bear granules.
O. sertata (Lym.) Ver. Tooth-papille two or three. Spines seven, finely serrulate, partly flattened.
DD.— Ophiothumnus Lym. Adoral shields large, wedge-shaped with the broad distal end separating the narrow ovate oral shield from the side arm-plate. Disk-scales exposed. Radial shields more or less naked, close together.
*Q, gracilis Ver. Arm-spines four or five, upper ones slender, lowest rough. Disk with truncate, thorny stumps. Tentacle-scale spiniform or palmate.
O. vicarius Lym. Disk-scales bear slender, tapered, acute spinules. Tentacle-scale small, conical.
O. exigua (Lym.)
Ophiacantha should be restricted and subdivided.
In this group the armature of the disk does not seem to be cor- related with other important characters ; neither does the number nor the length of the arm-spines, nor their solidity, or translucency, or hollowness, nor their degrees of roughness.
One of the characters that seems to be of much importance for the separation of the typical genus, from other allied generic groups,
A. EK. Verrili—North American Ophiuroidea. 329
hitherto confounded with it, is the nature of the adoral shields. In the typical group these are small and quite in front of the oral shields. In several other divisions they extend outward in a distal lobe that separates the oral shield from the side arm-plates, as in Ophiocopa, etc. (See group AA, p. 327, and group XIII, p. 340.)
Other characters of importance for the separation of groups of some value, are the presence of several tooth-papille at the apex of the jaw (groups B, C, G, K, pp. 330-333); the presence of a large sub- marginal oral tentacle-pore, with special papille around it, in a row or cluster (see group J, p. 333); the partial nakedness of the disk- scales and radial shields (group F, p. 332); the size and contiguity of the radial shields (group G, p. 332); the contiguity of the dorsal arm- plates (group G, p. 332); absence of tentacle-scales and the large size of part or all of the pores (group H, p. 333, and group EK, p. 332).
Some of these characters, even those of most importance, have not been referred to in many of the published descriptions, nor repre- sented in the figures. Therefore many of the species cannot, at present, be definitely classified. Mr. Lyman’s figures, in the Voyage of the Challenger, are generally very accurate, but even some of these fail to show certain details of structure needful for accurate classification of the species of this genus.
Ophiomitra Lyman (typical group) differs but little from some sections of Ophiacantha. It has the tooth-papille and distal oral papille numerous and clustered, as in section C; the distal oral ten- tacle-pore is large and partly exposed, as in section J. The radial shields are large and nearly naked and the disk scales are visible and spinose. Several species referred to Ophiacantha by Lyman also have naked disk-scales and radial shields (groups F and B, aa).
Subdivisions of Ophiacantha.
From the preceding remarks and table, it will be plain that several genera and subgenera* may be separated from the old genus Ophia- cantha with characters that appear to be of as great morphological value as those that characterize, for instance, Ophiomitra or Ophio- chiton.
* Most of these subdivisions were proposed in the Report on the Ophiuroidea of the Bahama Exped., Nat. Hist. Bull., Univ. Iowa, v, 1899. (Designated as 1899qa in this article.)
330 A, EF. Verrill—North American Ophiuroidea.
Series I. Ophiacantha (restricted). Types, O. setosa and O. bidentata. Group A.—Typical Ophiacantha.
Oral papille form a simple row. One median tooth-papilla at the tip of the jaw. No special oral tentacle-scales at the distal angles of the mouth-slits, though the outer papilla, which serves as a ten- tacle-scale, may be wider than the rest. Oral tentacle-pore not exposed outside of the jaw-margin. Disk-scales more or less ob- scured by integument and bearing spinules, thorny stumps, crotchets, or granules. Radial shields rather narrow, separated more or less, mostly concealed by cuticle. Arm-spines usually long and slender, unequal, more or less rough, often glassy or translucent, often hol- low. Dorsal arm-plates usually all separated by the side arm-plates; sometimes, on a few basal joints, they are slightly in contact.
To this section a large majority of all the described species belong.
Group B.— Ophientodia Ver., 18994, p. 41.
Two, three or four tooth-papillz clustered at the tip of the jaws. Otherwise nearly as in section A. Distal oral papillz not clustered.
The published figures of several species shows two paired papilla, directed centrally, at the tip of the jaws. They may not always stand on the dental plate’and in such cases should be counted as oral papille, but in some cases they have been determined as true tooth- papille. Probably in this section there may be a central tooth- papilla that has been overlooked in some species, by reason of its position, high up on the jaw, or its smaller size. In some cases it may have been accidentally lost. But in some specimens either two or three papille occur on different jaws. Therefore, I consider the presence of three tooth-papillz as the usual character of this division. The species need revision as to the tooth-papille. (See also group VII, page 338.)
a.—Radial shields rather small, narrow, mostly concealed.
O. scutata Lym, Three tooth-papille; eight to ten thorny arm- spines. Radial shields long and narrow, sometimes naked.
O. cuspidata Lym. Three tooth-papille.
O. pectinula Ver. Three or four tooth-papille. Dorsal arm-plates pectinate on the outer edge.
Pes
= et, oe, eS
A. E. Verrill—North American Ophiuroidea. 331
_aa.— Ophioscalus Ver., 1899a, p. 42. Radial shields large, wide, closely joined, naked. Disk-scales covered with rough spinules. Arm-spines approximate dorsally.
O. echinulatus Lym. Arm-spines ten, very thorny and glassy. Tentacle-scales spiniform, two on the first joint. Two or three tooth- papille.
Group C.— Ophiectodia Ver., 1899a, p. 42.
Outer oral papille (oral tentacle-scales) several, forming a cluster or a double row, some often standing on the lower face of the jaw or adoral shield. ‘Tooth-papillze one to three, or more. The oral papille are clustered nearly as in typical Ophiomitra.
O. enopla Ver. 'Tooth-papillz, one or two.
O. rosea Lym. Tooth-papille clustered, three or more.
O. spectabilis Sars. Tooth-papille three or four, in a cluster.
Series IT.
Group D.— Ophialewa Ver., 1899a, pp. 38,42. Types, O. Nut- tingti (Ver.) and O. tuberculosa (Lym.). (See p. 326.)
The dorsal arm-plates are broadly in contact, at least on many of the proximal joints. Disk-scales bear spinules or granules. Radial shields separate, sometimes more or less exposed distally, sometimes covered. Arm-spines rather short, few, nearly smooth, the rows not approximate dorsally. Oral papille nearly as in typical Ophia- cantha (group A). —
O. Nuttingii Ver., 1899a, p. 46. Arm-spines four, short, Oral- shield very large, ovate. Disk-scales more or less exposed, bearing conical spinules.
O. rufescens Keehl. Off the Azores, 845 meters. Ventral plates contiguous. Arm-spines six, finely serrulate. ‘Two large elongated tentacle-scales. Oral papill six or seven, outer one largest. Disk- scales covered with fine roundish granules. Distal end of radial shields naked.
O. tuberculosa Lym.,’98. EE. Indies. Disk and radial shields covered with cuticle and granules. Arm-spines four. One tooth- papilla. Three oral papille, the distal one broad and notched. Oral shield not large, transverse. Tentacle-scale single, small.
332 A, FE. Verrili— North American Ophiuroidea.
Group E.— Ophientrema, sub-gen. nov. Type, 0. scolopendrica (Lym.).
Tentacle-pores and scales on one, or a few, basal joints and larger than usual,* farther out decreasing rapidly to a small or rudimen- tary size. Disk-scales concealed by granules. Radial shields some- times partly exposed. Spines numerous, nearly smooth. Mouth- parts as in typical Ophiacantha.
O. granulosa (Lym.). Radial shields largely exposed, broad, in contact distally. Arm-spines ten, slender, the rows nearly approximate dorsally. Tentacle-pores of the first joint large, with one flat scale; of others small, with a narrow scale. Dorsal arm-plates all separate. Pacific.
O. scolopendrica (Lym.’83). Radial shields nearly concealed, close together. Arm-spines seven, unequal, the rows not approximate dorsally. Tentacle-pores large on four joints, with a small scale, rudimentary or lacking distally. Dorsal arm-plates joined on a few basal joints. European.
sroup F.— Ophiomitrella Ver., 1899a, p. 39. Type, O. levipellis (Lym.).
Disk-scales visible, bearing granules or spinules. Radial shields partly naked, not large, wide apart. Arm-spines slender, thorny or serrulate ; the rows approximate dorsally in the type. One tooth- papilla. In the type-species a pair of special, distal, oral tentacle- papille, on the first under arm-plate,t directed into the mouth-slit. Adoral shields wide. Otherwise the mouth parts are nearly as in typical Ophiacantha.
O. levipellis (Lym.,’83). Arm-spines eight, slender, thorny. Disk- scales naked or partly granulated. Upper arm-plates separated.
Group G.— Ophiacanthella Ver., 1899a, p. 39. Type, O. Troscheli (Lym.). Radial shields naked, long, parallel, in contact by their edges. Dorsal arm-plates largely joined. Three tooth-papille. Arm-spines nearly smooth.
* Several species that have been referred to Ophiomitra also have this charac- ter. (See Ophiomitra, section AA, p. 351.)
| The two papilliform appendages of the first under arm-plate are here sup- posed to be movable, but with the published figures and descriptions it is not always possible to distinguish them from the solid, immovable, crest-like lobes which are present on these plates in the same position in many species, including O. bidentata, Among extralimital species, these papillae are found in some species, such as O. serrata Lym., that have the disk-scales and radial shields
concealed.
ros.
A, E. Verrili— North American Ophiuroidea. 333
Seriss III.
In the following groups the oral shield is separated from the side arm-plates by the adoral shields.
Group H.— Ophiopora Ver., 18994. Type, O. Bartletti (Lym.).
Tentacle-pores all large and open. No tentacle-scales.
Group I.— Ophiolimna Ver., 1899a. Type, O. Bairdii (Lym.).
Spine-crest of the side arm-plates distally situated and oblique. Spines nearly smooth. Disk granulose and spinulose. Jaws more or less granulose.
Group J.— Ophiopristis Ver., 1899a. Type, O. hirsuta (Lym.). A row of distal oral papille alongside of the large outer ora
tentacle-pore. Arm-spines partly flattened with serrulate edges. (See p. 347.)
Group K.— Ophiotreta Ver., 1899a, p. 40. Type, O. lineolata (Lym.). One or two flat, distal oral papille by the side of the large oral tentacle-pore. Two or three tooth-papilla. Spines mostly terete, but sometimes flattened and with serrulate edges. (See p. 328.)
Group L.—Amphipsila Ver., 18994, p.55. Type, A. maculata Ver.
Oral papille form a simple row. Two or three tooth-papille in the marginal series, Disk-scales and radial shields naked, small. Tentacle-scale slender, spiniform or palmate. Arm-spines serrulate, flattened, hollow. (See p. 348.)
Ophiocopa Lym. also belongs in this series.
Ophiacantha, sens. ext.
Artificial groups of species from the West Indian region and from the East Coast of North America, arranged accord- ing to various special characters.
All the species appear in groups I and II. In these two groups all the northern species are indicated by an asterisk prefixed. ‘These groups are not intended as natural sections of the genus, though they
Trans. Conn. Acap., Vou. X. OcToBER, 1899.
23
334 A, EF. Verrill—North American Ophiuroidea.
may be so in some cases, but merely as aides for the comparison of the species. ‘They may be considered as morphological tables. The genus in these XIII groups is taken as in Mr, Lyman’s works.
For the natural subdivisions, see pages 329-333.
I: Arm-spines long, thorny or prickly, more or less glassy.
a.—Rows of spines approximate dorsally on first or second joint beyond disk. 6.—Radial shields covered ; disk spinulose.
O. aspera Lym.,’78. Arm-spines ten, slender, very thorny.
O. pentacrinus Liitk. Spines nine or ten, long, very slender, slightly thorny.
O. pectinula Ver., sp. nov. Spines ten or eleven, very slender and glassy. Tooth-papille twotofour. Acluster of oral tentacle-papille.
*Q. millespina Ver.,’79. Spines ten. Disk spinules slender, with three or four long sharp branches.
*O, gracilis Ver.,t 85. Spines four to six, short, except the upper basals; lower ones thorny. Disk-scales naked, with hour-glass shaped, thorny stumps.
*O. varispina Ver., ’85. Spines eight, little rough, glassy.
6b.—Radial shields largely exposed.
O. levipellis Lym., 83. Spines seven or eight, little flat; naked disk-scales.
O. echinulata Lym., 78. Spines nine or ten, long ; disk spinose. See under Ophiomitra.
aa.—Basal rows of spines not closely approximated dorsally.
O. stellata Lrym.,’75. Arm-spines seven, very thorny. O. scutata Lym., ’78. Spines nine or ten, slightly thorny. *O. granulifera Ver., ’85. Spines eight or nine, part of them slightly thorny. ; IY:
Arm-spines, in the adult, not distinctly thorny, but often finely serrulate on the edges, especially the lower ones; mostly rather opaque, but often translucent in alcohol ; usually hollow.
+This singular species, on reexamination, proves to belong to Ophiothamnus. Its long, wedge-shaped oral shields are widely separated from the arm-plates by the broad adoral shields,
A. EF. Verrill— North American Ophiuroidea. 335
a.—Spines slender, tapered, terete, or but little flattened; often nearly smooth, or only microscopically serrulate; rougher when young.
b.—Basal rows of spines approximate dorsally on the first or second joint.
e.—One odd tooth-papilla.
O. vepratica Lym. Arm-spines eight, long and tapered.
O. segesta Lym., ’78. Disk-spines slender, mixed with thorny stumps.
*O. Bairdii Lym., ’83. Disk with granules and some tapered spines. Jaws granulated more or less.
* 0. bidentata (Retz.). Spines somewhat rough or serrulate, espec- ially when young. Disk-spinules are small, thick, rough, obtuse stumps.
* 0. abyssicola Sars. Disk-spinules minute.
*O. aculeata Ver.,’85. Spines eight or nine, tapered, upper ones nearly smooth. Disk-spinules with three to five sharp points.
*0O. anomala Sars.+ Spines eight or nine, all terete, tapered, finely serrulate; six arms.
*O. enopla Ver.,’85. Spines four or five, serrulate; outer oral papillz clustered.
ce.—Two or three tooth-papille. Distal oral papille or oral tentacle-scales clustered.
*O. spectabilis Sars. Disk-spinules large, tapered, acute.
6b.—Basal rows of spines not very closely approximate dorsally. d.—Dorsal arm-plates, at base of arms, separated by side plates, or only slightly in contact.
*O. fraterna Ver.,’85. Disk covered with minute spinules having three to five sharp points.
*O. crassidens Ver., 85. Disk with small tapered spines. Oral papille very stout.
O. Bartletti Lym., ’83. (Ophiopora Ver., p. 345.) No tentacle- scales.
dd.—Dorsal arm-plates, at base of arms, distinctly joined. e.— Disk and radial shields covered with spinules, or mixed spines and granules,
+ This species is viviparous. One specimen from off Nova Scotia has several six-armed young clinging about the mouth and genital slits.
336 A. E. Verrili—North American Ophiuroidea.
O. cosmica Lym.,’78. Arm-plates only a little joined. O. lineolata Lym., ’83. Arm-plates broadly joined. Disk with grains and some spines. .
ee.—Jaws also more or less granulated.
O. mixta (Lym.,’78). Disk with grains and spines. ( Ophiolimna V., p- 845.)
eee.—Disk granulated ; radial shields partly naked. J.—Radial shields joined. (Ophiacanthella V., p. 344.)
O. Troscheli Lym., ’78. Jf.—Radial shields separated. (Ophialewa V., p. 331.)
O. Nuttingii Ver., 1899a, p. 46.
aa.—Spines partly distinctly flattened and serrulate on the edges ; the rows not approximate dorsally on the basal joints. Two or three tooth-papille. (Ophiopristis V., p. 347.)
O. hirsuta Lym.,’75. Spines four or five, slender, part flat. Disk with long, very slender spines.
O. ensifera Ver., 1899a, p. 47. Spines four, stout, mostly flat. Disk with small, conical spinules..
O. cervicornis Lym.,’83. Spines five, mostly slender, acute.
O. sertata Lym. Spines seven, translucent. Tentacle-scales two.
LIT:
Radial shields more or less exposed.
A,.—Radial shields partly or wholly in contact. a.—Radial shields rather wide, angular.
O. echinulata. Inner edges of radial shields wholly in contact. (See Ophioscalus, p. 331, also p. 342.)
aa.—Radial shields narrow and long.
O. Troscheli. Shields naked and largely in contact. (See Ophia- canthella, pp. 332, 344.)
AA,—Radial shields separated, not large. b.—Never entirely concealed.
O. levipellis. Small, wide apart. (See Ophiomitrella, pp. 343, 352.)
bb.—Sometimes nearly or quite concealed.
A, EF. Verrill—North American Ophiuroidea. 337
O. sertata. Small, pear-seed-shaped.
O. scutata. Long and narrow.
O. ensifera. Small, narrow, usually largely covered. (See Ophiopristis, p. 347.)
O. Nuttingii. Crescent-shaped, narrow. (See Ophialcca, p. 331.)
Several other species are apt to have the distal end of the radial shields more or less exposed. In many cases the covering is probably accidentally rubbed off.
IV.
Dorsal arm-plates, on basal part of arms, in contact, not separated by the side-plates.
a.—Dorsal arm-plates rather narrow.
O. lineolata. Ventral plates slightly separated. (See Ophio- treta, pp. 333, 347.)
aa.—Dorsal arm-plates broad. 6.—Dorsal plates extensively joined. (See Ophiacanthella, p. 344.)
O, Nuttingii. Ventral plates in contact. O. Troscheli. Ventral plates separated.
6b.—Dorsal plates little joined.
O. cosmica. Ventral plates separated. O. mixta. (See p. 346.)
Me
Tentacle-pores all unusually large.
O. Bartletti. No tentacle-scales. (See Ophiopora, p. 345.)
O. cervicornis. Two spiniform tentacle-scales. (See Ophio- pristis, pp. 333, 347.)
VI.
A cluster of three or more tooth-papillz at the tip of the jaw.* O. lineolata (three or four tooth-papillz). O. scutata (three or four, often only two visible below).
* Several extralimital species belong to this group, such as O. Valenciennesi Lym., with three tooth-papillz and one oral tentacle-scale ; O. cuspidata Lym. ; O. marsupialis Lym.; O. rosea Lym.; the last has a cluster of tooth-papille and also several oral scales, (See p. 338, note, and p. 348.)
338 A, E. Verrill—North American Ophiuroidea.
O. sertata (three, only two visible below).
O. Troscheli (three tooth-papille).
O. spectabilis (three to five tooth-papille ; a cluster of distal oral papille,
O. pectinula Ver. (two or three tooth papille; three or more distal oral tentacle-papille).
Vil.
A pair of tooth-papille (or apparent tooth-papille) close together, at the tip of the jaw ; no odd median one visible from below. (In some species the odd papilla may be concealed by the pair below it, when it is actually present, but published figures and descriptions are not definite enough to determine this in many cases; in some cases it may have been accidentally broken from the type-specimen. It certainly seems to be the normal condition, in group a, to have only two.)
a.—A distal cluster or row of special oral tentacle-scales. See Ophiopristis, p. 347.
O. cervicornis.
O. ensifera.
O. hirsuta.
aa.—Only one or two distal oral tentacle-scales.
O. Bartletti. No tentacle-scale on arms. O. echinulata. Two spiniform tentacle-scales on arms.
WAL:
A cluster of oral papille or oral scales near the outer corner of the mouth-slits, at the outer oral tentacle-pore, or else one or two special oral scales by the side of the tentacle-pore, which is on, or nearly outside of, the margin of the mouth-slit. (Published figures are often inaccurate as to this character.)
a.—Several distal oral tentacle-scales or papille to each pore.
O. cervicornis. About four spiniform distal papille in a row. O. pectinula Ver. Three to four distal papille. *0O. enopla. Four to six distal papill in a cluster.
+ Several extralimital species belong to this group ; among them are: O. rosea Lym. (in subsection a); O. marsupialis Lym. ; and O. Valenciennesit Lym. (in aa.)
ge) OP:
A, EF. Verrili— North American Ophiuroidea. 339
O. hirsuta. 'Two to four papille in a row. O. ensifera. Four to five papille in a curved irregular row. *Q, spectabilis. Three or four in an irregular group.
aa.—One or two special distal oral scales or papillae, usually attached to the adoral plate ; oral tentacle-pore large.
O, Bartletti. One, spiniform, distal oral papilla.
O. lineolata. Two flat papille.
O. sertata. Two flat papillee.
O. levipellis. One flat papilla, attached to first arm-plate.
IX.
A pair of small, apparently movable, oral papille attached to the proximal end of the first under arm-plate, which is emarginate.+ In most species of the genus there are, in this place, two flat, usually fixed, processes or crests. It is generally impossible to tell, from published figures, the character of these parts. They are generally badly represented,
*0O. anomala.,
O. levipellis. (See group VIII, aa.)
X. Oral papillz unusually large and stout. * 0. crassidens. Three or four thick and rough papille, the distal ones smaller, * 0. varispina. Three or four broad, flat, obtuse papille.
we.
Tentacle-scales of peculiar or unusual forms, or spiniform.
a.—Tentacle-scales elongated ; flat, spatulate, or lobate distally.
O. aspera. End of tentacle-scale branched or thorny. *O. varispina. End spatulate.
aa.—Tentacle-scales elongated, slender or spiniform.
O. echinulata. Scales dagger-shaped ; two pairs on first joint. O. cervicornis. 'Two, spiniform, slender. O. Troscheli. One, long, acute.
+ Of extralimital species, O. serrata Lym., O. cornuta Lym., and 0. Valen- ciennesi Lym. belong to this group.
340 A. E. Verrill—North American Ophiuroidea.
O. vepratica. One, large, conical.
O. segesta. One, small, acute.
*O. gracilis. One, slender, palmate, distal ones acute. O. pectinula V. Spiniform, two or three on first joint.
XII.
Tentacle-scales wanting. Adoral shield asin XIII. ( Ophiopora V.) ,
O. Bartletti (see p. 345). Outer oral tentacle-pore large, with a conical papilla.
XIII.
Adoral shields long, usually trilobed; the distal lobe separates the oral shield from the side arm-plate. In all typical species of the genus the oral shield and adoral shield both join the side arm- plate.
A.—One or two tentacle-scales. B.—Disk-scales mostly concealed. a.—Jaws not granulated, or only slightly so. ( Ophiopristis V.) 6.—A row or series of special distal oral papille. O. hirsuta. (See p. 336.) O. ensifera. (See p. 336.) O. cervicornis. (See p. 347.)
bb.—One or two distal oral papille or scales. (Ophiotreta V.)t+ O. sertata. (See p. 348.) O. lineolata. (See p. 348.) AA.—Jaws granulated. (Ophiolimna V.) *O. Bairdii. (See p. 346.) O. mixta (Lym., as Ophiocheta). See p. 346. BB.—Disk-seales entirely exposed. Radial shields more or less naked. Adoral shields broad distally. ( Ophiothamnus.) *O. gracilis Ver. Disk-scales bear hour-glass-shaped spinules with a terminal group of points. AA.—No tentacle-scales, Tentacle-pores all very large and open. ( Ophiopora V.) O. Bartletti. (See p. 345.)
| Among extralimital species that belong to this group, are O. placentigera (Lym.) and O, Valenciennesi (Lym.).
<A ia tel
ost
A. E. Verrill—North American Ophiuroidea. 341
Ophiacantha scutata Lyman.
Ophiacantha scutata Lym., Bull. Mus. Comp. Zool., vol. v, p. 229, pl. i, figs. 1-3, 1878; op. cit., vol. x, p. 261, 1883 (variations).
Specimens sent to me by Mr. Lyman differ somewhat from his figures and descriptions.
The oral papille may be either three or four on different jaws of the same specimen; they are rather stout, spiniform, the inner largest, and all appear to be on the buccal plate, but the jaw-plates and adoral shields are so closely united together that the sutures are mostly invisible. There is often an extra outer papilla of small size, which is situated at the union of the buccal and adoral plates, out- side the oral tentacle-pore, which is large, but situated inside the mouth-slit. Tooth-papille may be from two to four on different jaws of a large specimen. Usually there is a stout median one with a pair of smaller ones just above it, invisible from below, and another small median one outside. The last is often lacking, and the upper pair may be replaced by a single one, which is, perhaps, absent in small specimens. The oral shield is more nearly transverse- elliptical than figured, with a more obtuse inner angle. The madre- poric shield is longer than the others and more rhombic, thickened, with a median concavity. The adoral shields of a large specimen are smaller than figured, narrow and tapered proximally, and the ends do not meet medially, but in small specimens they are nearly as figured and meet medially. The first under arm-plate is small, rounded, emarginate on the inside, with a small vertical crest at each side, directed inward.
The under arm-plates of the larger specimen are unlike the figure; they are narrower and longer, with the distal end projecting and strongly convex; the proximal end is very obtusely angulated or subtruncate; a little farther out on the arm they become more oblong, with the outer end more projecting and the inner end trun- cate and scarcely narrowed. They are slightly separated.
Tentacle-scales, on two or three of the basal joints, are flat, erect, lanceolate, and cuspidate ; occasionally, in the larger examples, there are two on the first joint. Beyond the fifth or sixth joint they become slender, acute, spiniform. The first two or three pairs of tentacles are decidedly larger than those beyond.
Arm-spines, in the largest rows, are ten, shaped about as figured, with numerous small, sharp prickles on all sides. The basal rows
342 A, FE. Verrill—North American Ophiuroidea.
approximate dorsally, but not so closely as in some other species. Upper arm-plates nearly as figured, transverse lozenge-shape, with the distal edge convexly curved at first, but becoming prominent and angulated farther out.
The whole upper surface of the disk, including the radial shields, is thickly covered with small, short, thorny spinules or stumps, ter- minated by three to five or more short, sharp points. Diameter of disk of largest specimen, 14"™; of smallest, 7”.
Off Barbadoes, 200 fathoms, Blake Exped.
Ophiacantha (Ophiectodia) pectinula, sp. nov.
Disk-scales small, entirely hidden by cuticle and bearing crowded, very slender, elongated spinules, thorny on the sides and at the tip. Radial shields small, near together, entirely hidden, by cuticle and spinules.
Two to four clustered tooth-papille. Oral papille numerous, nearly equal, compressed, spiniform, smaller than the tooth-papille ; four or five form a regular lateral row ; five or six more distal ones form a cluster or two rows, and serve as oral tentacle-scales. Oral shield broadly pelecoidal, wider than long, with a slightly convex distal lobe. Adoral shields about as large as the oral, oblong-lunate, meeting within.
Tentacle-scales long, acute, spiniform, thorny, two at the basal pores. Arm-spines long, very slender, thorny, very acute ; some of the rows nearly approximate dorsally at base of arms, where there are nine or ten in arow. Dorsal arm-plates small, quadrant-shape, the sides nearly straight, the outer end convex with a marginal row of minute, sharp denticles. Under arm-plates small, widely separated ; the inner end forms an obtuse angle ; the outer end is convex, prom- inent, side arm-plates prominent, meeting above and below.
Diameter of disk, 8™™; length of arms, broken at tips, about Le
West Indies, Blake Exped., 1883.
This was sent to me by Mr. Lyman as 0. echinulata, with which it does not agree. The type of the latter has broad naked radial shields, in close contact, and the oral papille are much fewer and are figured as forming a simple row. The spines and tentacle-scales are similar, though the basal rows of spines approximate more closely dorsally.
a a a
A, E, Verrill—North American Ophiuroidea. 348
Ophiomitrella Ver., 1899a, p. 39. (See p. 336, and p. 352.) Ophiomitrella levipellis (Lym.) Ver.
Ophiacantha levipellis Lym., Bull. Mus, Comp. Zool., vol. x, p. 259, pl. vi, figs. 82-84, 1883. Ophiomitrella levipellis Verrill, Ophiur. Bahama Exped., v, p. 39, 1899.
About twenty specimens of this species were sent to me by Mr. Lyman. They show considerable variation among themselves, and all differ more or less from his figures and description.
The disk is strongly five-lobed, owing to a deep incurvature of the interradial areas. The upper side is closely covered with small thin scales, which are usually smooth and nearly destitute of gran- ules, but in some examples there are a few scattered, low, verruci- form grains, especially near the margins; in others the grains are thinly scattered over nearly all the surface ; in some cases part of the grains are conical. The scales themselves vary somewhat in size and distinctness.
The radial shields appear to be long and narrow and nearly par- allel ; a narrow ridge, in dry specimens, often runs inward nearly to the center from each shield ; only the ends of the shields are com- monly exposed ; this naked part varies in form and extent, but is usually long, narrow, wedge-shaped, widest distally, and the ends often project somewhat beyond the edge of the disk over the base of the arm and may bear a few marginal granules. The ends of the shields are sometimes near together, being separated by a space less than half their breadth; in other specimens they are separated more than their breadth.
Oral papillze vary in number, even on the different jaws of the same specimen, from three to five; most frequently there are three in the regular series, with a smaller and much shorter distal one, just at the distal end of the adoral shield and above the large pore of the outer oral tentacle. In many cases this outer papilla develops to full size, like the next one, which is stout, erect, obtuse, larger than those that follow it; the latter are usually compressed vertically, sub- acute ; the inner one is a little longer and more conical. Attached to each inner corner of the first arm-plate there is a small vertically flattened scale or papilla that appears to be movable ; it guards the oral tentacle on the inside and is sometimes wanting. It corresponds to a similar process which in several other species seems immovable.
Tooth-papilla one, or perhaps none, for the odd papilla at the tip of the jaw agrees nearly in size and form with the true teeth. It
344 A. EF. Verrili—North American Ophiuroidea.
varies in form, however, even on the different jaws of the same specimen. It is usually ovoid, or obtusely lanceolate, or even obovate; sometimes it is acute or mucronate at tip, and then it differs a little more decidedly from the teeth. It stands on the tip of the dental plate.
The oral and adoral shields are thickened and prominent, shaped nearly as in the figure by Mr. Lyman. The oral shield is small and somewhat fan-shaped, or rather pelecoidal, for the inner lateral edges are strongly incurved. In one specimen the oral shields were unusually narrow and acutely angled proximally. ‘The adoral shields are relatively large, lunate, confined to the proximal side of the oral shield.
First under arm-plate is small, irregularly six-sided, strongly emarginate within. The second is much broader than long, curved distally, and obtusely angled proximally. Those following are still shorter, transversely narrow-elliptical, with a very obtuse proximal angle, or nearly truncate and broadly curved distally, often showing a slight median incurvature of the edge, which becomes more dis- tinct on those farther out. They are thick and widely separated by the side arm-plates, which lie in grooves. More distally they become more uearly square, with the inner end more angulated.
Tentacle-scale small, spiniform, subacute, rather rough, becoming more slender farther out. All the tentacle-pores are small.
Arm-spines about as figured, except that many of them are more thorny, especially those near the base of the arms and in the upper series, most of which have irregular sharp divergent thorns; farther out they are mostly minutely serrulate. They are not usually dis- tinetly flattened, as stated, but slender, terete, tapered, acute. The rows are closely approximate dorsally on the second and third joints, becoming separated farther out.
Upper arm-plates thick, swollen, widely separated, rather triangu- lar or quadrant-shaped, with an obtuse proximal angle and a broadly convex distal edge. On the middle of the proximal part there is a small, wart-like elevation.
Diameter of disk of those described above, 3 to 6™™,
Off St. Vincent, 88 and 124 fathoms. Blake Exped.
Ophiacanthella Verrill, 1899a, p. 89. Type, 0. Troscheli (Lym.) Ver.
Three terminal tooth-papille in a group. Radial shields long, narrow, largely in contact, more orless naked. Disk-scales obscured by cuticle, granulose or spinulose. No special oral tentacle-scales.
A. EF. Verrill—North American Ophiuroidea. 345
Oral papille all similar. Dorsal arm-plates largely in contact. Tentacle-scales one or two, all similar. Spines slender, finely serru- late or nearly smooth. 2
This genus is, in most respects, closely related to Ophiomitra. It differs in having the disk-scales mostly concealed by cuticle ; in hav- ing three apical tooth-papille, instead of one ; in having the dorsal arm-plates joined ; and in the smoothness of the spines.
It is separated from Ophiacantha especially by the naked and contiguous radial shields, and from the typical section of that genus by having three tooth-papillz and contiguous dorsal arm-plates.
Ophiopora Ver., 1899a, pp. 39, 43. Type, O. Bartletti (Lym.) Ver.
Adoral plates with two distal lobes, one of which embraces the lateral edge of the oral shield and separates it from the side arm- plates, as in Ophiopristis. No tentacle-scales ; tentacle-pores all very large and widely open. Two or three tooth-papille. Outer oral tentacle-pore is submarginal and furnished with one special, acute, papilla or oral tentacle-scale on the adoral shield. Disk-scales, above, and the radial shields are concealed by cuticle and spinules ; on the under side the scales are visible. Arm-spines few, nearly smooth. Dorsal arm-plates are separated by side plates.
This genus is closely allied to Ophiolimna, but differs in the large open tentacle-pores, without scales.
Ophiopora Bartletti (Lym.) Ver. Ophiacantha Bartletti Lym., Bulletin Mus. Comp. Zodl., vol. x, p. 256, pl. v,
figs. 738-75, 1883. Ophiopora Bartletti Ver., Ophiur. Bahama Exp., Bull., v, p. 39, 1899.
This is the only known species of this group. The disk is covered by slender acute spinules. The four arm-spines are small, tapered, - rather short, nearly smooth. The side arm-plates are not prominent.
West Indies, 291 fathoms, Blake Exped.
Ophiolimna Ver., 1899a, pp. 40, 44. Type O. Bairdii (Lym.) Ver.
Adoral shields trilobed ; one distal lobe extends back between the oral shield and first side arm-plate, so that the oral shield is detached from the arm. The jaws may bear more or less granules. Disk-scales and radial shields covered with granules, or spinules, or
both.
346 A. £. Verrill—North American Ophiuroidea.
Side arm-plates prominent, with the oblique, spine-bearing crest near the distal margin, so that the spines are directed more or less distally, especially on the distal half of the arm. One or two tooth- papille. Several simple oral papille in a regular row, the outer ones broader. One or two tentacle-scales. Arm-spines tapered, nearly smooth, rather short.
This genus agrees with Ophiacantha and Ophiomitra in its mouth- parts, but differs from both in the more oblique position of the rows of arm-spines and in the separation of the oral shields from the side arm-plates. In the last character it agrees with Ophiopristis, Ophio- pora, ete. :
The type, O. Bairdii (Lym., 1883,)* was taken by the Blake, off the east coast of the United States, in 1242 and 1394 fathoms, and by the United States Fish Commission Steamer Albatross in 1390 fathoms, in the same region.
This species has seven or eight smooth, acute spines, of moderate length ; the rows closely approximate dorsally ; the disk bears small acute granules and a few short spines ; jaws partly naked, but with some granules; outer oral papilla (oral tentacle-scale) broad and flat; one tooth-papilla; one small tentacle-scale; oral shield broadly obovate; dorsal arm-plates scarcely joined on basal joints. The spine-crests are distally placed on the side arm-plates, and the spines are mostly directed distally, or often lie nearly parallel to the arms.
Ophiolimna mixta (Lym.) Verrill.
Ophiocheta ? mixta Lyman, Bulletin Mus. Comp. Zool., vol. v, p. 222, pl. ii, figs. 40-42, 1878; Voyage Challenger, Oph., vol. v, p. 110, pl. xxxix, figs. 15-17, anatomy, 1882.
Ophiolinna mixta Ver., Ophiur. Bahama Exped., Bull., v, p. 40, 1899.
This species has two large flat tentacle-scales ; seven smooth arm- spines ; two tooth-papille; six oral papillae; jaws granulated ; dor- sal and ventral arm-plates not separated by the side arm-plates on the basal joints ; disk crowdedly covered with granules mixed with some slender spines.
The internal structure, as figured by Lyman, is much like that of Ophiacantha, The radial shields, seen from within, are broad, three-cornered, separated. It does not appear to be closely allied to Ophiocheta.
West Indies, in 160 to 576 fathoms, Blake Exped.
* Bulletin Mus. Comp. Zodl., vol. x, p. 256, pl. v, figs. 70-72 (as Ophiacantha).
A, FE. Verrili— North American Ophiuroidea. B47
Ophiopristis Ver., 1899a, pp. 39, 44,47. Type, O. hirsuta (Lym.) Ver.
Adoral plates elongated, three-lobed; the distal end is two-lobed; one distal lobe joins the first under arm-plate; the other joins the first side arm-plate and separates it from the oral shields, so that the oral shield is quite detached from the side arm-plates, as in Ophio- copa. Disk-scales bear spinules or granules. Radial shields are separated, mostly concealed by cuticle. Arm-spines rather long; in the typical group, mostly flattened and with regularly serrulate edges. The-rows are not approximate dorsally. Tentacle-scales one or two.
Tooth-papille two to four. Oral papillze numerous ; two or more of these are special oral tentacle-papille or scales, guarding the large outer oral tentacle-pore, which is on or near the margin of the jaw and is conspicuous. First under arm-plate is concave and has a flat process on each inner corner.
Synoptical table of Ophiopristis and Ophiotreta.
A.—Typical Ophiopristis. A row of several small conical or slen- der, distal, oral tentacle-papille outside the large pore. Arm-spines partly flat with serrulate edges. Two or three tooth-papille. Dorsal arm-plates separated, but rows of spines not approximate.
O. hirsuta Lym. (See p. 336.)
O. ensifera Ver. (See p. 336, Pl. xxi, fig. 4.)
O. cervicornis Lym. A regular row of four or five slender oral tentacle-papille. Two tooth-papille. Two spiniform tentacle-scales; the pores very large and open. Disk and radial shields covered with fine granules and small acute spinules; some spinules on upper arm- plates. Six short flat spines serrate on the edges.
AA.— Ophiotreta Verrill, 18994, p. 40. (See p. 333.) Spines terete or only little flattened. One or two, rarely three, distal : oral papille or scales at the large oral tentacle-pore. Two to five or more tooth-papille.
a.—Dorsal arm-plates joined. Rows of spines not approximate dorsally. Spines nearly smooth. 'Tooth-papille three to five. Distai oral tentacle-scales flat, blunt, two or three. Two tentacle scales on several basal joints ; one is flat, the other spiniform.
348 A. E. Verrill—North American Ophiuroidea.
O. lineolata (Lym.). Disk and radial shields closely covered with granules and spines. Jaws bear granules. Arm-spines seven.
aa.—Dorsal arm-plates separate. Rows of spines not closely approximate. Two or three tooth-papille. Spines nearly smooth or finely serrulate or thorny; some a little flattened. —
O. sertata (lym.). Arm-spines seven. Tentacle-scale large, flat. Two flat, distal, oral scales. Disk covered with granules and tapered spines. Radial shields sometimes naked, ovate, separate, but usually concealed.
O. Valenciennesi (Lym.). Tooth-papille three; oral tentacle-pore large, marginal, with a large round distal scale ; a pair of small papille on first under arm-plate ; two large tentacle-scales. Spines four, with blunt thorny tips. Disk granulated.
To this group may also be referred O. placentigera (Lym.), off Fiji Is., 1350 fath. It has three tooth-papille ; a large, broad, oral tentacle-scale; granulated disk; six smooth spines; one tentacle- scale.
Amphipsila Verrill, 1899a, p. 55. Type, A. maculata Ver. Puate XLII. Ficures 5, da.
Disk rounded, covered with thin, naked scales, above and below. Radial shields narrow, separated, naked. Arm-plates distinct, above and below. Arm-spines of moderate length, numerous (five to twelve), serrulate. Oral shields clearly visible, at least when dry. A simple row of oral papillae. Only two or three conical apical papille, in a row; these may be considered as tooth-papillx, but there is no distinct cluster of inner tooth-papille, below the teeth, as in Ophiopsila. 'Tentacle-scale spiniform.
I have separated this genus from Ophiopsila, as understood by Lyman, for he included in the latter A. fulva (Lym.), which is closely allied to our type-species.
In true Ophiopsila (type, O. aranea), to which O. Riiset of the West Indies also belongs, there is a cluster of many special tooth- papille, within the mouth, below the teeth, as in Ophiocoma, and the disk is covered with thick cuticle, nearly or quite concealing the scales. It appears to belong to the family Ophiocomide, while our genus seems to be closely related to Ophiacantha, with which it agrees in its mouth-parts and spines. It differs from typical Ophia- cantha it its naked disk-scales and radial shields, in having the upper arm-plates joined, and in the distal prolongation of the adoral plates, much as in Ophiopristis and Ophiolimna, though less distinct, ow- ing to its narrowness.
A, FE. Verrili— North American Ophiuroidea. 349
Ophiomitra Lyman.
Bulletin Mus. Comp. Zodél., vol. i, p. 325, 1869; Voyage Challenger, v, pp. 202-209, 1882, pl. xlv, figs. 4-6, (anatomy). Verrill (restricted), Oph. Bahama Exped., Bull., v, p. 57, 1899.
This genus is very closely allied to Ophiacantha. The only special distinctions given by Lyman are the larger size and naked- ness of the radial shields and the naked or nearly naked scales of the disk.
Mr, Lyman also described the disk of the type-species as rounded and cap-like,—a character due, perhaps, to immaturity, for in large specimens of that species the interradial margins are incurved or emarginate.
When adult the type-species (0. valida Lym.)* has numerous spiniform, clustered oral papille and tooth-papille. The distal oral tentacle-pore is large and sub-marginal, partly sheathed by proximal processes from the concave first under arm-plate and inner side of the jaw. The adoral shields are very broad, but wholly proximal to the small oral shields. The basal tentacle-pores are larger and fur- nished with two prominent tentacle-scales. The large, broad radial shields are largely in contact. The disk-scales are not large, of nearly uniform size, without specialized marginal ones, and bear coarse, short, clavate, thorny stumps. The arm-spines are numerous, somewhat thorny and glassy. The dorsal arm-plates are slightly separated by the side-plates.
Most of the species subsequently described by Mr. Lyman and others differ much from the type, in several characters,
They nearly all have a single odd tooth-papilla and a simple row of oral papille, as in typical Ophiacantha. The interradial mar- ginal scales are usually large and specialized. The radial shields are often entirely separate and in some cases not particularly large.
In fact, they have little in common with the type, except the partial nakedness of the radial shields and disk-scales,—characters also found in species of Ophiacantha.t Therefore it seems necessary to subdivide the genus,
* The specimens originally described and figured by Lyman were all immature, and had not developed the true character of the mouth-parts.
+One species (O. Normani) referred to this genus by Mr. Lyman does not agree with it even in these characters, for the separated radial shields are no larger and no more exposed than in several species of Ophiacantha, and its disk- seales are granulated. In its arm-spines, which are smooth and only four in
Trans. Conn. AcaD., VoL. X. OcroBerR, 1899. 24
350 A, E. Verrill—North American Ophiuroidea.
Ophiotrema of Koehler seems to be very closely related to typical Ophiomitra.
Like the latter, it has clustered tooth-papille and oral papille, with a large, conspicuous distal oral tentacle-pore and special papillee around it. The tentaclé-pores are large and surrounded by several small acute spinules. Disk-scales are small, visible, bearing acute rough spinules. Radial shields are small, naked, separate, divergent. Arm-spines five, serrulate.
O. Alberti Keehl., 96, the type, is from off the Azores.
Certain species referred to Ophiomitra by Keebler appear to belong to Ophiomitrella. (See p. 352.)
Synoptical table of the species that have been referred to Ophiomitra (sens. ext.) and Ophiomitrella,
Group A.—Typical Ophiomitra. |
Tooth-papille several, clustered. Oral papille numerous, clus- tered distally. Distal oral tentacle-pore large. Xadial shields large, naked, usually joined. Disk-scales visible, all of moderate size, bearing stumps or spinules. Arm-spines thorny and usually glassy. ‘Iwo or three tentacle-scales on the basal joints, in the type. Adoral shields broad, proximal to the oral shields. Interradial mar- gins of disk somewhat incurved, sometimes convex, not deeply notched.
a.—Radial shields largely in contact. 6.—Disk-scales bear short clavate or capitate stumps. Rows of arm-spines not approximate dorsally.
O. valida Lym., 769. Arm-spines nine or ten, solid, not very long, roughly serrulate or thorny. bb.—Disk-scales bear longer and shorter, tapered, thorny spines. Rows of spines approximate dorsally. O. ornata Ver,, 1899a.. This Vol., pl. xii, fig. 3. Arm-spines
long, slender, thorny.
ad.—Radial shields divergent, in contact only distally. Arm- plates separate above and below.
number, it differs from all the species of Ophiomitra. Therefore I have referred it to Ophiacantha, with which it agrees in its mouth parts. (See p. 329.)
Another species (O. exigua Lym.) I refer to Ophiothamnus. (See p. 328.) This, also, has smooth spines, and the small oral shield is separated from the side arm- plates by the large adorals.
= OS ieee
A, EF. Verrill—North American Ophiuroidea. 351
*O. spinea Ver. Oral papille not very numerous distally. Disk- scales small, with small conical spinules. Arm-spines eight or nine, rough, hollow, the rows not approximate dorsally. One or two tentacle-scales.
AA.— Ophioplinthaca, gen. nov. Type, O. dipsacos Lym. One odd tooth-papilla, oral papille in a nearly simple series, not clustered distally. Interradial margins of disk notched or deeply emarginate. Marginal and submarginal disk-scales large and specialized. Arm-spines thorny and glassy, usually hollow. xadial shields large and broad, naked or nearly so. First tentacle-pore decidedly larger, with two or more scales. Oral shield joins the first side arm-plates.
B.—Radial shields in contact along most of their length. Rows of spines not approximate dorsally.
e.—Upper arm-plates distinctly separated by the side plates.
O. dipsacos (lym.). Arm-spines six, hollow, very thorny, the two upper ones very long. First tentacle-pore with two scales.
ec.—Upper arm-plates on basal joints slightly joined, or barely separate. Two or more distal oral tentacle-scales, similar to other oral papille.
O. incisa (Lym.). Arm-spines five or six, flattened, thorny on edges. Three to six tentacle-scales at first pore, one or two farther out. Disk-scales with few, nearly smooth, conical spinules.
BB.—Radial shields not in contact, or only slightly so distally. Spines glassy and thorny, the rows not approximate dorsally.
d.—Radial shields in contact at the distal ends, very large and wide.
e.—Upper arm-plates separated.
O. carduus (Lym.). Spines six, very thorny. Disk-scales with thorny stumps, marginal scales very large. Tentacle-scale one, lobate. .
ee.—Upper arm-plates, on basal joints, not separated by side plates. First tentacle-pore larger, with two or three scales.
A flat oral tentacle-scale. O. plicata (Lym.). Radial shields joined distally in young; sepa- rate in adult. Disk-secales with small, conical spinules. Arm-spines
352 A. EF. Verrili—North American Ophiuroidea.
five or six, short, very thorny. Distal edge of under arm-plates bent downward.
dd.—Radial shields well apart, with intervening rows of scales.
f.—Upper arm-plates not separated by the side plates. Two tenta- cle-scales on first joint.
O. Sarsii (Lym.). Radial shields rather small, far apart. Disk- scales with small conical spinules. Arm-spines seven or eight, very thorny. 'Tentacle-scale lobate. A stout oral tentacle-scale.
j.—Upper arm-plates separated by the side plates. Disk-scales coarse; marginal ones larger.
O. chelys (Lym.) Radial shields narrow, sunken. Disk with small conical spinules. Arm-spines six, hollow, very thorny. One tentacle-scale.
AAA.— Ophiomitrella Ver., 18994, p. 39. (See p. 343.) Radial shields small, wide apart, naked. Disk-scales all nearly alike, not very large, outlines easily visible. One tooth-papilla. Oral shields join the side arm-plates. Interradial margins of the disk not deeply emarginate and without large scales. Disk bearing scattered granules or stumps.
g.—Arwm-spines serrulate, the rows not approximate dorsally.
O. globulifera (Kehler, ’95). Arm-spines five. Disk-granules glassy, spherical. Europe, 1700 meters.
O. cordifera (Keebler, ’96)._ Arm-spines six or seven. Disk with small capitate granules. Azores, 1143 meters. q
gg.—Arm-spines thorny and glassy, the rows approximate dorsally. A special outer oral tentacle-scale on the first under arm- plate.
O. levipellis (lym., as Ophiacantha). Seven or eight spines. Disk-scales naked or sparsely granulated ; all small. Disk pentag- onal. (See also page 343.)
O. cornuta (Lym., as Ophiacantha). Spines eight. Disk rounded, bearing small thorny stumps; a larger scale between the radial shields. (See p. 339, note.)
AAAA.—Radial shields very large and in contact, covering most of the disk. Oral shields small, triquetral, not joining the side arm-plates; adorals large and broad. Arm-spines smooth or nearly so, the rows not approximate dorsally.
A. FE. Verrilli—North American Ophiuroidea. 353
O. ewigua (Lym.). Disk-scales few, coarse, with few thorny stumps. Arm-spines six, rather short, tapered; oral papille three, the outer one on the adoral shield. 'Tentacle-scale one.
The last named ‘species should, I think, be referred to Ophio- thamnus, with which it agrees well in all external characters. (See p. 350.)
O. Normani Lym., omitted from the table, is to be referred to Ophiacantha (typical group) as already stated (p. 349, note). It has small and rather widely separated radial shields and four smooth arm-spines.
Ophiomitra valida Lyman.
Ophiomitra valida Lyman, Bull. Mus. Comp. Zool., i, 10, p. 325, 1869; op. cit., x, p. 264, 1883; Lyman, Il]. Cat. Mus. Comp. Zool., vi, pl. ii, figs. 4-6; Lyman, Report Voy. Challenger, Zool. Ophiuroidea, v, p. 209, pl. xl, figs. 4-6, 1882.
Ophiomitra cervicornis (young) Lyman, Ill. Cat. Mus. Comp. Zool., viii, pt. ii, p. 14, pl. ii, figs. 19, 20, 1875; Bull. Mus. Comp. Zool., vol. v, part 9, p. 231.
Ophiomitra valida Verrill, Ophiur. Bahama Exped., Bull., v, p. 58, 1899.
Several large specimens of this species, sent by Mr. Lyman, differ considerably from his figures and description, which were made from immature specimens.
The most important differences are found in the mouth-parts. The largest of our specimens have very numerous oral papillae and tooth-papillz, crowded together in clusters, very much as in Ophio- camax. 'The tooth-papille often consist of a row of three, on or below the margin, and of two or three pairs above these, next the teeth, but frequently they are so crowded that no such regular arrangement can be made out. There may be as many as nine or ten on one jaw-apex. The distal oral papillze form crowded groups of four to seven, or more, or they may stand in two rows, so as to cover or conceal most of the width of the jaw. They are all rather stout, spiniform, subacute. The distal oral tentacle-pore is large, marginal, exposed, nearly surrounded by sheath-like processes of the jaw and first under arm-plate, which is small and deeply concave. The oral tentacle is large, with a thickened basal part into which the distal part can be retracted.
Tentacle-scales two or three on the basal joints ; farther out the pores and scales decrease in size rapidly. On the basal joints the inner scale is large, flat, lanceolate, erect, hollowed out on the side next the tentacle; the other is narrow, subacute. The under arm-
354 A, E. Verrili— North American Ophiuroidea.
plates are smaller on the basal joints than farther out. They are broader than long, with a concave emargination on the distal edge.
The dorsal arm-plates are rather quadrant-shaped, with a broad lobe on the distal edge, and with prominent lateral angles. Some of the basal arm-plates are slightly in contact. Arm-spines ten, rather stout, roughly serrulate, blunt, the rows not approximate dorsally. The radial shields are larger, irregularly triangular, more or less encroached upon by the disk-scales and granules. The disk- scales are not large, all nearly equal, sparsely covered with short, coarse, rough, capitate stumps. The interradial margins have a small notch in dried specimens, but not larger scales.
Oral shields small, pelecoidal, with an acute inner angle, and a prominent convex outer end. Adorals about as large as the orals, wide, lunate, the surface finely granulous in appearance. In younger specimens the distal oral papille form one irregular row of about three or four around the pore.
Common throughout the West Indies in 10 to 1105 fathoms (Blake Expedition).
OPHIOCAMAX Lym. Type, O. vitrea Lym.
Ophiocamax Lym., Bull. Mus. Comp. Zool., vol. v, p. 156, 1878; Voy. Chal- lenger, v, p. 209, 1882.
This genus is closely allied to typical Ophiomitra, Like the latter it has numerous tooth-papille in an apical cluster, and a cluster of distal oral papillee, even more numerous than in Ophiomitra.
In the type-species there is also a special small distal plate (pro- cess of under arm-plate?) which bears two or three small papille directed proximally and serving as part of the papille for the large oral tentacle-pore.
The basal tentacle-pores have three or four elongated, erect tenta- cle-scales forming a sheath for the tentacles. Radial shields are wide and in contact. Disk-scales, which are usually visible, bear thorny spinules, but in the type species they are scarcely visible and closely spinulose in the adult.
The adoral shields are large and broad, situated in front of the oral shields.
Synoptical table of the species of Ophiocamaz.
A.—Typical. Dorsal arm-plates, at base of arm, not separated by the side-plates. Rows of spines not approximate dorsally.
rye
A, EF. Verrill— North American Ophiuroidea. 355
O. vitrea Lym. Disk closely covered with small, acute spinules; the scales nearly concealed. Tooth- and oral papille very numerous, slender, acute. Nine thorny arm-spines. Tentacle-scales large, obtuse.
- O. hystrix Lym. Disk-scales visible, irregular, bearing few, short, conical, rough spinules. Radial shields in contact distally. Arm- spines eight, slender, the.upper ones very long.
AA.—Dorsal arm-plates all separated. Rows of spines not approxi- mate dorsally.
O. fasciculata Lym.,’83. Arm-spines six, rather short, flattened, serrulate on the edges. Disk-scales plainly visible, bearing few small, tapered, acute spinules. Radial shields not very large, broad, wholly in contact. Dorsal arm-plates widely separated.
O. austera Ver., Ophiur. Bahama Exped., p. 60, pl. vi, figs. 1, la, pl. vii, fig. 2. Arm-spines seven, slender, the upper ones very long and very thorny, scarcely flattened. Disk-scales visible, bearing longer and shorter, rough, acute spinules. NRadial shields large, triangular, extensively joined. Dorsal arm-plates nearly in contact on the basal joints. Four lanceolate tentacle-scales. A cluster of about six acute, distal, oral papillae, pointing inward on each side, part of them arising from the lateral lobes of the deeply bilobed first under arm-plate. (See this vol., Plate xin, figure 2.)
OPHIOCHONDRIN &, subfam. noy.
This group differs from typical Ophiacanthidz chiefly in having the internal arm-plates so modified that the arms can be coiled in a vertical plane, like the Astrophytons, etc. The arm-spines are short, subequal, not very rough. The disk-scales may be thickly covered with cuticle and granules, or they may be naked. ‘The oral papille are few, ina simple row. The thick cuticle sometimes covers the mouth shields and lower side of the arms.
The modifications of the ambulacral ossicles fits these species more perfectly for clinging closely to gorgonians, etc., by coiling the arms closely around the branches. But this power is also common to various species of Ophiacantha, in a lesser degree.
Ophiochondrella Ver., gen. nov. Type, O. sgywamosus (Lym.).
This differs from true Ophiochondrus in having the disk covered with naked scales, above and below; in having the under arm-plates
356 A. FE. Verrili— North American Ophiuroidea.
covered and concealed by thick cuticle ; in having the under arm- plates in contact; and in having two tentacle-scales.
The arm-spines are short, nearly equal. Radial shields naked, ovate, and separate.
Ophiochondrella squamosus (Lym.). Ophiochondrus squamosus Lym., Bull. Mus. Comp. Zool., x, p. 275, pl. vii, figs. 108-110, 1883. The disk-scales are thick, swollen, irregular. Arm-spines eight, tapered. Oral papille three, small, spaced. Tentacle-scales minute, rounded. West Indies, 250 fathoms, Blake Exped.
Ophiochondrus Lyman. Bulletin Mus. Comp. Zool., i, p. 328, 1869; Voy. Challenger, p. 247, 1882. Type, O. convolutus Lyman.
The characters given to this genus by Mr. Lyman should be modi- fied by adding that there are two or three nearly vertical plates at the base of the arm, supporting the ends of the radial shields, so that the edge of the disk is considerably raised above the arm, making a sharp angle with it. The radial shields are still more strongly sup- ported by an elongated genital plate, running up each side of the genital slits and joining the radial shields.
Ophiochondrus crassispinus Lyman. Ophiochondrus crassispinus Lyman, Bulletin Mus. Comp. Zool., vol. x, p. 275, 1883.
Several specimens from the Blake Exp., Station 232, 88 fath., off St. Vincent, were sent to the Yale Museum by Mr. Lyman under the name of O. convolutus, under which they are also evidently recorded in Mr. Lyman’s lists of 1883.
These, on careful study, appear to belong to O. crassispinus, as defined and figured by Mr. Lyman. The latter was described from a single specimen, from 229 fath., Blake Exp.
They have three acute, conical, oral papille, exclusive of the odd terminal one. The oral shield is small, thick, pear-shaped, with an acute proximal angle ; arm-spines six or seven, short, nearly equal. A very small acute tentacle-scale is usually present in the larger specimens. Upper arm-plates on proximal part of arm, except two basals, are nearly quadrant-shaped with the outer edge convex and the lateral angles acute; distally they become more nearly trian- gular, with the sides a little convex.
ce erg
A, FE, Verrili—North American Ophiuroidea. 357
The radial shields are large, elongated, separated by a band of small flat scales. Central part of disk and interradial spaces covered by very small, flat, naked scales. The arms are relatively stout, of moderate length, tapering rapidly.
0. convolutus L, is described as having the disk granulated ; oral papille four and squarish ; oral shield broader ; radial shields shorter and broader; besides other differences. Possibly the two forms may be only variations of one species, but none of my specimens are intermediate.
One of the specimens from Station 232 was clinging closely to a group of Zoanthoid corals (Hpizoanthus). The genus is evidently adapted to living clinging to gorgonians and similar organisms, for protection.
Family, OPHIOSCOLICID Ltk., 1869.
Ophiomyxide (pars) Ljung., 1866.
Ophioscolicine (sub-family) Ljung., 1871.
The upper side of the arms is covered with naked skin, beneath which the arm-bones can usually be seen. Under arm-plates and side arm-plates are present, though sometimes much degenerated. Arm-spines are moderately long, often rough or thorny, two to six in number. Tentacle-scale often wanting, but three or four are present in Ophiambix. Oral papille are usually numerous and form a continuous row, but sometimes they are few, and rarely lacking (in Ophiobyrsa). Tooth-papille usually lacking, rarely present as irreg- ular spiniform papille. Teeth simple, spiniform.
Disk covered with a soft skin, which may contain minute scales and may bear granules or spinules. Radial shields small or rudi- mentary, sometimes lacking. Oral and adoral shields normal.
Internally the arm-bones of Ophioscolex are deeply grooved ven- trally and dorsally and cut away laterally at the ends; the mouth- frames are reduced and simple, but the peristomial plates are large and in three pieces.
In some of the other genera the arm-bones are more rudimentary. In Ophiogeron and Ophiosciasma they are entirely separated, along the median plane, in two elongated parts, curved towards each other. This is an embryonic character, illustrating the relatively low development of the skeleton in this family.
This family, as now known, seems to be more nearly allied to Ophiacanthide than to Ophiomyxide, with which it was formerly united.
358 A, FE. Verrill—North American Ophiuroidea.
Externally there are no tangible characters to distinguish this family from certain of the Ophiacanthide, except the lack of upper arm-plates, which are always present in the latter. But the internal structure, so far as studied, is peculiar.
The family, as here understood, includes the following described genera: Ophioscolex, Ophiosciasma, Ophiogeron, Ophiobyrsa, and Ophiambiz.
Certain species that have been referred to some of these genera do not agree in structure with the typical species, and therefore I have established two new genera for their reception :
Ophiobyrsella, gen. nov. Type, Ophiobyrsa serpens Lyman.
Astrogeron, gen. nov. Type, Ophiogeron supinus Lyman.
Ophioscolex fragilis Ver., sp. nov.
Five slender arms. Oral shield small, narrow, pear-shaped, with an acute proximal angle. Adoral shields narrow, oblong, the inner end acute and touching. Oral papille six or seven, forming a row which is not regular in the middle ; the two outer ones are larger than the others, tapered, acute; the four inner ones, which do not lie in just the same line, are small, slender, acute. Lower arm-plates hour-glass-shaped, narrow, longer than wide, truncated at the ends and closely joined, and apparently soldered with the side plates. Arm-spines three, slender, tapered, acute, nearly equal, about as long as a joint. Tentacle-pores large. No tentacle-scale. The disk is destroyed in my specimens.
Diameter of the disk-scar, 10™™; length of arms, 22™™.
Off Barbadoes, Station 2938. Blake Exp., 82 fathoms.
Ophiobyrsella Ver., gen. nov. Type, O. serpens (Lym.).
Disk pentagonal, covered entirely by naked skin, which hides the oral and adoral shields and extends out over the upper and under sides of the arms and spines. Small spinules are situated over the region of the radial shields and along the margins of the disk, or over the whole disk. No tentacle-scales ; tentacles large. Arm- spines three to five,—three in the type; rough, glassy. About five spiniform teeth. Oral papille form a regular lateral row, besides two or three tooth-papille at the tip of the jaw.
This genus is very near Ophioscolex in external characters.
True Ophiobyrsa (type, O. rudis) differs in having only one oral papilla, no teeth, and only a few spiniform tooth-papille ; these parts being very much reduced.
A, E. Verrill—North American Ophiuroidea. 359
Ophiobyrsella serpens (Lyman) is from the West Indies, in 69 fathoms.
Another species, O. hystricis (Lyman), with five slender and rather long spines, was dredged off the Shetland Islands, in 345 fathoms. (Bulletin Mus. Comp. Zool., vol. x, p. 272, Pl. viti, figs. 120-122, 1883.)
Astrogeron Ver., gen. nov. Type, Ophiogeron supinus Lym.
Disk and arms covered by a naked skin containing minute scales ; beneath the skin are small rounded radial shields and very short genital plates and scales. ‘Teeth small, spiniform; a cluster of spini- form tooth-papille at the end of the jaw, and a row of oral papille on the edge. Oral and adoral shields normal. No tentacle-scale. About two slender, glassy arm-spines covered by skin. Arm-bones divided longitudinally. The typical species of Ophiogeron has no oral papille, the jaw-plates being naked except for a few small spini- form teeth at the tip, but, otherwise, it agrees pretty closely with this. .
Astrogeron supinus (lirym.) is from the West Indies, in 200 to 464 fathoms.*
Mr. Lyman has described several very remarkable genera, allied in some respects to Ophioscolecidw, but presenting such peculiar structures that it does not seem reasonable to refer them to any of the described families. They should be considered as types of two distinct families, if the differences that separate families in other cases are taken as our criteria, In fact, they present greater diver- sities than can be found elsewhere in the entire group of regular Ophiuroids. Therefore I propose to classify them as follows :
Ophiomycetide, fam. nov.
Sub-family, Ophiomycetine, nov. Type, Ophiomyces Lyman. Sub-family, Ophiotholine, nov. Type, Ophiotholia Lyman.
Ophiohelide, fam. nov. Type, Ophiohelus Lyman.
These groups will be described on subsequent pages.
Family, OPHIOMYCETIDZ, nov.
Disk swollen, covered with scales, which may be either naked or spinulose. No radial shields. Teeth few. Two, three, or more apical tooth-papille. Oral papille numerous, the outer ones large,
* Bulletin Mus. Comp. Zool., vol. x, p. 270, Pl. vii, figs. 108 to 106, 1883.
360 A. EF. Verrill—North American Ophiuroidea.
flat, foliate or spatulate, recurved, in two or more divergent rows or clusters, partly on the adoral plates. The arms can be turned up vertically above the disk.
’
Sub-family, Ophiomycetine, noy.
Disk with small but distinct scales, usually spinulose. No grapel- shaped spinules on the arms. Arm-spines of the basal joints, and sometimes the lower ones on some of the joints beyond the disk, are flattened or spatulate ; others are long and slender. Tentacle-scales of basal joints flat, often multiple. Upper arm-plates small, sepa- rated. Side plates large, meeting above and below. Oral shield small, Adoral shield long, carrying many of the spatulate oral scales,
Internal mouth-frames slender; genital scales and plates broad, flat, and curved up over the base of the arm. Arm-bones well developed, but peculiar in form, and without a distal condyle. The arm can be turned up vertically above the disk.
Only four or five species are known. Two, O. mirabilis Lym. and O. frutectosus Lym., are West Indian. 0. spathifer Lym. is from off Japan, in 565 fathoms, and O. grandis Lym. is from the South Atlantic, in 1000 fathoms.
Sub-family, Ophiotholinge, nov.
No visible radial shields. Disk-scales delicate, spinulose. Arm- spines present on all joints, slender, about three ; associated with the spines, beyond several basal joints, are clusters of grapel-shaped spinules,* like those of Ophiohelus. A simple row of flat oral papille and tooth-papille surrounds the proximal ends of the jaws. Distally the oral papille and oral scales are very numerous, in several divergent rows, recurved, broad, flat, foliate or spatulate, much as in Ophiomycetes.
Several spiniform teeth. Tentacle-scales, on the basal joints, two or more, flat; on other joints, spiniform. 'The side arm-plates meet broadly below. Under arm-plates covered by cuticle.
This group is closely allied to Ophiomycetinw, from which it differs mainly in having grapel-shaped spinules on the distal joints and in the more simple arm-spines.
*Myr. Lyman describes these as ‘‘ pedicellariz,” but they are totally different from all forms of true pedicellaria. They seem to me strictly homologous with the curved hooks and hooklets of Astronyx and allied genera, though much more complex in structure. ;
A, EF. Verrili—North American Ophiuroidea. 361
Only a single species is known: Ophiotholia supplicans Lym., taken off Juan Fernandez, in 1825 fathoms, Challenger Expedition.
Family, OPHIOHELIDA, nov.
No radial shields. Disk-scales very thin or rudimentary. Arm- bones divided into right and left plates. Oral papillae spiniform, few, in asimple row. Teeth spiniform. Distal joints of arms bear rows of peculiar grapel-like or “parasol-shaped” spinules, in place of true spines. Only one genus is known.
Ophiohelus umbella Lym. Mem. Boston Soc. Nat. Hist., 1880, pl. i, figs. 5-10 and 16.
This, the type-species, was taken eff Barbadoes, in 82 fathoms.
O. pellucidus Lym. Op. cit., pl. i, figs. 11-15, 1880.
This species was taken off the Fiji Islands, in 1350 fathoms, Chal- lenger Expedition.
Family, OPHIOMYXIDZE Lijung. (restr.), 1866.
Ophioscolecide (pars) Lutk., 1869.
Ophiomyxinc (sub-family) Ljung., 1871.
Ophiomyxide Carus, Faune Medit., p. 96, 1884. Verrill, Oph. Bahama Exped., p. 65, 1899.
Disk and arms covered with thick cuticle, and usually with only a row of marginal disk-scales, and a few scattered ones imbedded in the cuticle, but visible only when dried. Radial shields small, usually with a proximal series of small supplementary scales.
Teeth and oral papille stout, flat, with the end serrated. No tooth-papille. True tentacle-scales generally absent. Under arm- plates small. Side arm-plates sub-ventral, bearing several rough divergent spines. Upper arm-plates rudimentary or lacking ; when present, composed of small pieces. Two large, triangular, peristo- mial plates on each mouth-angle.
Arm-bones peculiar, belonging to the modified ‘“hour-glass- shaped” type, with well-formed condyles on both ends.
Ophiomyxa and Ophiodera are the only genera described. The second genus has the following characters :
362 A, EF. Verrill—North American Ophiuroidea.
Ophiodera Verrill, Oph. Bahama Exped., p. 67, 1899. Type, O. serpentaria (Lym.).
Marginal disk-scales are rudimentary and concealed by thick cuticle ; the disk-scales proximal to the radial shields are lacking. No upper arm-plates. Side arm-plates may be soldered to the under arm-plates. They are not continued upward by a row of small plates. Three or four arm-spines enclosed in cuticle; the inner one is smaller and may serve as a tentacle-scale; it is sometimes forked distally. Teeth and tooth-papille serrate, nearly as in Ophiomyxa, but with finer denticles.
Ophiodera Stimpsoni Verrill.
? Ophioscolex Stimpsoni Lyman, Illust. Cat. Mus. Comp. Zool., viii, p. 28, pl. i, figs. 11-15, 1875. Ophiodera stimpsoni Ver., Oph. Bahama Exped., p. 67, pl. ii, figs. 4, 4a, 1899.
Pirate XLII. Ficurss 2, 2a, 2b, 2c.
Arms very long and slender. Disk five-lobed, the lobes extending out a little on the base of the arms. Teeth three or four; upper one stout, spiniform, the others thicker, subtruncate.
Whole upper surface of disk and arms and lower surface of disk are covered with thin naked cuticle, wrinkled when dry, containing imbedded, scattered, microscopic scales on the disk, and a row of irregular small, marginal scales. Sometimes there are a few small, irregular granules along the margin and on the under side of the disk, and also on the bases of the arms. Radial shields very small or rudimentary, concealed by cuticle.
Diameter of disk 7™; length of arms about 45™™.
West Indies, 60 to 240 fathoms.
Oral papille about five, partly slender, subspiniform, rough at tip, irregularly crowded in a row, nearly equal in length, but some are flattened and obtuse at tip. Sometimes there is also a somewhat stouter tooth-papilla. Within mouth-slits, on each side, there are two (sometimes only one) slender papillae between the two oral ten- tacle-pores,
Genital slits wide and open near the oral shields, but narrow dis- tally and not extending to the edge of the disk, bordered by narrow, naked scales.
Tentacle-pores are small and round. In some specimens there is a small, slender, spiniform tentacle-scale, which is often deeply forked, or even double, and in alcohol is covered with a sheath of cuticle ; it stands nearly in line with the other spines, beside the
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A, FE. Verrill—North American Ophiuroidea. 363
tentacle-pore. It is often reduced to a minute spinule, and is fre- quently absent.
Arm-spines three or four, divaricate, small, nearly equal, sharp, roughly serrulate and glassy, more or less covered by cuticle when in alcohol.
The internal arm-plates show as transversely rhombic plates sepa- rated by wider intervals.
Family, HEMIEURYALIDZ Ver., Oph. Bahama Exped., p. 70, 1899.
In this family are included several genera of true Ophiure, which very much resemble, in form and habits, the simple-armed Euryale or Astrophytons. Like the latter, they coil their arms closely around the branches of gorgonian corals on which they dwell.
The disk is pentagonal and covered with thick plates or tubercles, which may be conical. The radial shields are large and prominent.
Upper arm-plates may be entire and accompanied by supplemen- tary plates, or they may be replaced by a mosaic of small plates. They are thick or tubercular.
Under arm-plates well formed. Side-plates separated by extra plates. Oral and adoral shields normal. Spines few, short and stumpy. A row of oral papille. Teeth, but no definite cluster of tooth-papille.
Genital pores small, situated near together at the outer end of the oral shield. Arm-bones have special forms approaching those of the Astrophytons. Mouth-frames strongly ossified.
The genera belonging to this family are Hemieuryale, Ophioplus, and Sigsbeia.
Hemieuryale pustulata Von Martens.
Hemieuryale pustulata Von Mart., Monatsb. Konig. Akad. Berl., p. 484, 1867; Ljung., Dr. Goes., Oph., Ofv. Kong. Akad., p. 617; Lyman, Ann. Sci. Nat., xvi, Art. 4, p. 5; Bull. Mus. Comp. Zool., iii, 10, p. 268, pl. v, figs. 8-11; op. cit., x, p. 277; Report Voy. Challenger, Zool. Ophiuroidea, v, p. 249, pl. xliii, fig. 7-10 (anatomy), 1882.
Ophiura cuspidifera (?) Lamk., Hist. Anim. s. Vert., iii, p. 226, 2d ed., 1840 ; Encyclop. Meth., pl. exxii, figs. 5-8.
Disk small, thick, swollen, pentagonal, with a swelling opposite the base of each arm when dried; whole surface, except radial shields, covered with larger and smaller thick scales and verruce. The central primary scale is round and rough like the radial shields, but not swollen, Five primary rounded radial scales, which are
364 A. E. Verrill—North American Ophiuroidea.
larger than the rest, are strongly convex or pustular, and often white. Five somewhat smaller convex interradials may usually be distinguished by their size ; the other plates and scales are of vari- ous sizes, the larger ones convex or somewhat verruciform, while the small ones are nearly flat. A radial band of small scales extends continuously between the radial shields and out over the upper side of the arms, becoming flat, angular, and closely crowded on the arms, so as to form a fine mosaic.
Radial shields long-ovate, widely separated; the surface is evenly covered with fine hemispherical elevations; the side arm-plates are ornamented in a similar manner. Along each side of the arms, above, there is a row of elevated, verruciform or almost hemispheri- cal plates, alternating with the side arm-plates ; part of these are usually pure white, alternating with others that are deep brown. Under arm-plates well developed, trapezoidal, with acute outer angles, and with a rather deep median emargination or notch in the distal edge, except on six or seven basal ones; close to the base they are larger than broad, but farther out they become broader than long. Arm-spines generally two, small, short, nearly equal, obtuse, becoming longer and more slender distally. Tentacle-scale rather large, ovate, obtuse. Oral shield large, often white, somewhat ‘“‘spade-shaped ” with the outer end and sides, evenly rounded, the inner edges concave, the median and inner lateral angles acute; they vary considerably in different specimens. Adoral shields swollen, somewhat crescent-shaped or pear-seed-shaped, with the acute inner ends touching. Oral papille about five, angular, crowded in a close series, the outer ones larger. Genital openings small, like an angu- lar pore, between two angular plates, at the outer edge of the oral shields.
Color reddish or yellowish brown, spotted and blotched with clear white in various ways, so as to closely imitate the color and appear- ance of certain species of Gorgonella to which it habitually clings. Usually there are rows of white verruciform plates along the arms on the upper side, resembling in size and color the verruce of the coral, which has a brown ground-color, like that of the Heméeuryale. Part of the tubercular plates of the disk, part of the oral shields, part of the spines, and part of the side arm-plates are also usually white.
Common in water of moderate depth throughout the West Indies wherever the Gorgonella lives. Specimens from off Barbadoes have been in the Yale Museum many years. ‘Taken by the Blake Expedi- tion in 74 to 180 fathoms.
A. E. Verrill— North American Ophiuroidea. 365
Ophioplus Verrill, Ophiur. Bahama Exped., p. 70, 1899.
Type, Hemiewryale tuberculosa Lyman.
Disk small, pentagonal, thick, covered with small, thickened or tubercular scales. Radial shields large, naked, separated. Oral shields and adoral shields well developed and naked. Oral papille in regular series. No tooth-papille. Under arm-plates rather large. Upper arm-plates entire, swollen and well formed, separated by a transverse row of small, tubercle-like plates. Side arm-plates prom- inent, separated above by a supplementary lateral plate. Arm- spines short, two or three in a row. Tentacle-scale single. A pair of small, round genital pores under the outer end of the oral shields.
This genus differs decidedly from Hemieuryale, to which it is allied, in having distinct and well formed dorsal arm-plates. It is also closely allied to Sigsbeia. In fact, it stands between these two genera in several characters.
Ophioplus tuberculosus (Lym.) Ver. Hemieuryale tuberculosa Lyman, Bull. Mus. Comp. Zool., vol. x, p. 276, pl. vili, figs. 120-127, 1883. Ophiomusium (?) Nutting, Narrative, p. 78.
Ophioplus tuberculosus Ver., Ophiur. Bahama Exped., p. 71, pl. i, figures 1-1b, 1899 (description).
Puate XLII. Ficures 6-6d.
Color deep brown, variously spotted with whitish, imitating the colors of Gorgonella to which it clings.
Usually many of the more prominent verruciform plates of the upper side of the arms and disk are white; under arm-plates dark brown.
Taken by the Blake Expedition in 96 and 115 fathoms ; Bahama Expedition, Station 15 and 16, off Havana, 200 fathoms.
Sigsbeia murrhina Lyman.
Sigsbeia murrhina Lyman, Bull. Mus. Comp. Zool., v, 9, p. 284, 1878, pl. ili, figs. 55, 58; op. cit., x, p. 277; Lyman, Report Voy. Challenger, Zool., Ophiuroidea, v, p. 250, pl. xliii, figs. 4-6, 1882, anatomy ; Three Cruises of the Blake, ii, p. 114, fig. 399, 1888. Nutting, Narrative, p. 79. Verrill, Oph. Bahama Exp., pp. 72, 73, pl. ii, figs. 1, la, 1899 (Young, description.)
Puate XLII. Ficure 7.
This species clings to gorgonians, which it imitates by the form of its arms and the tuberculated surface of the disk, and probably also in color, when living. Our figure represents a young specimen, which differs considerably from the adult.
Trans. Conn. ACAD., Vou. X. OcroBER, 1899. 25
366 A, FE. Verrili— North American Ophiuroidea.
Family, OPHIOBRACHIONTIDZ Verrill, nov.
Disk entirely covered with acute spinules and cuticle, without radial shields or ribs. Arms long, slender, serpentine, so covered with cuticle that the plates are hidden. Under arm-plates are pres- ent. Side arm-plates not prominent, bearing rows of small double hooks on all the joints, but no spines. Upper arm-plates rudimen- tary or lacking. Tooth-papille spiniform, in an apical cluster; a few similar mouth-papille. |
The only species, Ophiobrachion uncinatus Lym., is from off Cuba, in 250 fathoms.
Order Il, EURYALZE Miill. and Trosch., 1842.
Euryalide Gray, Synop. Brit. Mus., p. 63, 1840.
Astrophytonide Norman, Ann. and Mag. Nat. Hist., xv, p. 104, 1865.
Phytastra Heckel, Gen. Morph., ii, p. 67, 1866.
Astrophytide Lyman, Ljungman, and others.
Euryale Mill. and Troschel, Syst. Aster., p. 85, 1842. Ljung., Oph. Viv., p. 334, 1867. Carus, Fauna Medit., p. 97, 1884. Verrill, Ophiur. Bahama Exped., p. 73, 1899.
Cladophiure Bell, Proc. Zool. Soe. London, p. 180, 1892; Catal. Brit. Echinod., p. 26, 1892.
Euryalida of several authors.
Family, EURYALIDZE (pars) Gray, 1840, restricted. Astrophytide (pars) Lyman, and many other authors.
Arms more or less dichotomous. Disk covered with cuticle and granules, and having ten strong radial ridges.
Teeth strong and large, in a single vertical row, as in Ophiure. Tooth-papille, few or none. Oral papillae minute, papilliform, or lacking. Adoral shields and jaw-plates large, well formed. Oral shields rudimentary.
Under arm-plates simple; they extend the whole length of the arm. Side arm-plates small, appressed proximally, but prominent or erect distally, where they bear double claw-like hooks and thorny spinules ; toward the base of the arms they bear few, small, rough, simple spines or tentacle-scales. ‘Two rows of small plates, extend- ing up from the side-plates, form transverse ridges, around the arms. Large spines, along the upper side of the arm, are borne by some of these plates. Dorsal arm-plates are represented only by small detached pieces.
~ Stars ahaha et *
A. E. Verrili—North American Ophiuroidea. 367
In having a regular row of teeth and simple normal under arm- plates this group resembles ordinary Ophiure and differs widely from the Gorgonocephalide.
Subfamily, Euryalinee, nov.
Arms wide at base, many times dichotomous, with short inter- nodes, bearing two dorsal rows of spines. Disk large, with ten granulated radial ridges. Radial shields long, narrow, composed of only one piece, covered, like the rest of the disk, with cuticle. Inter- brachial areas, below, covered with strong united plates.
The type, Euryale aspera Lam., is from the East Indies and China Sea.
Subfamily, Trichastrinse Ljung., 1872.
Disk relatively small, but thick, with ten stout radial ribs. Arms angular, stout and high at base, divided only distally into a small number of forks. Tentacle-scales short and stout, about three in a row.
Oral shield well developed.
The type, Trichaster palmiferus (Lam.), is from the East Indies.
The arms have two dorsal rows of short, stout, obtuse, conical spines ; they generally occur on alternate ridges.
The teeth are very large and thick, with truncate ends. Usually there is a single, large, conical tooth-papilla at the apex of the jaw. Oral papillz small, papilliform, in two or more rows. The disk and arms, above and below, are rather coarsely granulated.
Family, GORGONOCEPHALID Verrill.
Gorgonocephaline (pars) Ljung., 1867. Bell, Catal. Brit. Hchinod., p. 27, 1892.
Gorgonocephalide Vervrill (restr.), Ophiur. Bahama Exped., Bull. Univ. of Towa, v, p. 83, 1899.
Arms divided dichotomously into numerous branches. Disk swollen, with ten prominent radial ribs, covered with cuticle, which may bear granules or scattered spinules, or it may be more or less naked. Radial shields, each composed of several united plates.
The entire surface -of the arms and disk above and below is covered with cuticle which is usually granulated, so that the plates are hidden,
Under arm-plates mostly rudimentary, consisting of two or more small pieces, sometimes absent. Side arm-plates are united below
368 A, E. Verrili— North American Ophiuroidea.
and cover most of the under side of the arms. They bear a row of few, small, rough spines or spiniform tentacle-scales, which are usually hook-like distally. Two or more rows of small plates run up from each of the side plates and form transverse ridges around the arms, covered with granules; these usually bear rows of small glassy hooks. The dorsal arm-plates are rudimentary or wanting.
Teeth and tooth-papille numerous, spiniform. Oral papillae, when present, small, conical or papilliform, Adoral shields well-developed, but usually concealed by cuticle, sometimes broken into several plates. Oral shields rudimentary or wanting. Sometimes there are five small madreporic plates, but usually only one.
The three generic names: Gorgonocephalus Leach, 1815; Huryale Lamarck, 1816 ; and Astrophyton Agassiz, 1835, were, as originally used and intended, exact synonyms. As now employed, they only date back to Lyman’s paper on the Challenger Ophiuroidea, 1878.
That he rightly divided these forms into three distinct genera cannot be doubted, and he doubtless had the right to apply the three names, as he did, to the respective groups, though it might, perhaps, have saved some confusion of nomenclature if he had given new names to two of the genera.
It is certainly useless to go back to Linck, 1733, as the prior authority for Astrophyton, for he was not a binomial writer.
For the same reason it is useless to try to restore the ancient pseudospecific names given by Linck and even by Seba (e. g. costo- sum), when later and determinable specific names have been given by binomial writers.
Gorgonocephalus Leach (Zool. Miscell., 1815) is the oldest of the three names under the binomial system, Leach gave a short diagnosis of the genus, and stated that he separated it from Ophiwra on account of its branched arms. He mentions no special type, but refers to the fact that most writers, following Linné, had referred all the species to “ Asterias caput-medus.”
As the latter was primarily based on a species of northern Europe, Lyman’s selection of the northern genus to bear this generic name was fully justified.
As for the other two names, since they were synonyms he could have applied each of them to either of the remaining groups with equal propriety, for each name had been used for all the known species.
There certainly is no good reason why Mr. Lyman’s usage should not be followed, so far as these genera are concerned.
A, E. Verrill—North American Ophiuroidea. 369
This family, as here defined, includes only two described genera : Gorgonocephalus and Astrophyton. To these should be added a third, to include Huryale verrucosum Lam., which differs much from both the others. For this I propose the name Astrocladus.
Astrocladus Verrill, gen. noy.
Resembles Gorgonocephalus in form, but differs in having no pavement of plates on the margins and interradial areas ; in the absence of under arm-plates ; in having no minute hooks on the arm- ridges ; in having no spines or tentacle-scales on the basal points ; in having the side arm-plates more degenerate, and not covering all of the under side of the arms, leaving spaces of naked cuticle between them. The arms have very numerous forks.
The type, A. verrucosus (Lam.), has rather large, rounded or verruciform tubercles, arranged in two irregular rows along the upper side of the arms and on the radial ridges of the disk.
There are usually three short, stout, obtuse tentacle-scales, thorny at the tips.
The tooth-papille and teeth are very numerous, elongated, spini- form. The oral papille form two or more rows; the larger ones are cylindrical or spiniform, the smaller ones conical.
The whole surface of the arms and disk, above and below, is covered with fine and close granules. The annulations of the arms are not very prominent.
The forkings of the arms are very numerous, with short inter- nodes. The arms are stout at the base.
The adoral and oral shields are represented by a group of irregu- lar plates. The interradial areas below are covered with thin gran- ulated cuticle, without plates.
It has been recorded from the Cape of Good Hope, ete. A speci- men in the Yale Museum is labeled as from Japan, but this locality may possibly be erroneous.
Family, ASTROCHELIDZ: Verrill.
Astrochelide Verrill, Ophiur. Bahama Exped., p. 79, 1899.
Arms simple or with a few distal forks, granulated, and also annulated with raised ridges. Disk with five or ten radial ridges, its surface granulated or spinulose.
The genital openings are short, situated toward the margin of the disk, or not close to the inner angles.
Under arm-plates rudimentary or lacking. Side arm-plates cover most of the under surface, but are hidden by cuticle and granules.
370 A. E. Verrili— North American Ophiuroidea.
They bear a short row of small rough spines or tentacle-scales ; above them are double vertical rows of small plates, forming raised ridges and bearing granules and also rows of minute glassy hooks, on the sides and top of the arms. These sometimes extend on the radial ridges of the disk.
Teeth and tooth-papillze numerous, spiniform ; the latter form an apical cluster. Oral papille similar in form, sometimes lacking. The teeth may form double vertical rows.
This family includes Asétrochele, Astrogomphus, Astroporpa, and Astrotoma all with simple arms, and Astroenida with the arms forked near the ends.
Family, ASTROSCHEMIDZ: Verrill. Astroschemide Verrill, Ophiur. Bahama Exped., v, p. 76, 1899.
Arms simple, long, slender, coiled. Disk five-lobed, with ten radial ribs ; naked or granulated. Radial shields narrow, usually elongated. Under arm-plates small. Upper arm-plates poorly developed, often wanting, sometimes represented by two or more pieces, covered by naked skin or granulated. Side arm-plates rela- tively large, covering a large part of the lower side of the arm, and usually bearing two elongated spines or tentacle-scales.
Teeth are large, stout, several in a vertical row. Oral papille are small or wanting.
Oral and adoral plates, regularly formed, but covered by cuticle. Genital slits short, situated near the outer margin of the disk.
Internal mouth-frames strong, well developed, but without wing- like processes.
This family includes Astroschema, Astrocreas, and Ophiocreas.
Family, ASTRONYCIDZ Verrill. Astronycina (pars) Ljung., Oph. Viv., 1867. Bell, Cat. Brit. Echin., p. 27, 1892.
Astronycide Verrill, Ophiur. Bahama Exped., Bull. Univ. Iowa, v, p. 74, 1899.
Arms undivided, long, slender, coiled, not annulated nor granu- lated. Disk with ten narrow radial ridges formed by long narrow radial shields, covered with thin, smooth scales or naked skin.
Upper and under arm-plates rudimentary or absent. Side arm- plates cover most of the lower side of the arm and project laterally, bearing two, three, or more spines or tentacle-scales, which may be either simple or hook-like. The genital slits are short, near together in a depression near the oral shields.
SCE
ee
ee ea) ee tas
A. F.. Verrili— North American Ophiuroidea., 371
Teeth stout, well formed, in a single row. 'Tooth-papille one or two, conical, sometimes absent. Oral papille small, like conical granules, placed above the margins of the jaw. Oral and adoral plates regularly formed.
Astronyx was the only described genus of this family, till recently, when I added to it a new genus, Astrodia (type, A. tenuispina Ver.), from deep water off the U. S. coast.
Astronyx Lymani Verrill.
Astronyx Loveni Lym., Bull. Mus. Comp. Zool., vol. x, p. 282, pl. viii, figs. 136-138, young (non Mill. and Troschel).
Astronyx Lymani Verrill, Ophiur. Bahama Exped., v, p. 74, pl. viii, figs. 4-4e, 1899.
Puate XLII. Ficurrs 6-6ce.
Arms five, long, slender, coiled. Disk pentagonal with incurved margins, and ten high, long radial shields, which are widely sepa- rated, curved outward in the middle and somewhat sinuous distally, the outer end a little clavate or knobbed ; the edge is serrulate with small scales. The radial shields and disk are covered with a thin, smooth skin which extends out on the arms, above and below. Inter- brachial region below, in the dry specimen, concave or sunken, with the two short but wide genital openings close together, near the inner angles.
Astrodia Verrill. Type, Astronyx (2) tenuispina Ver. Astrodia Verrill, Ophiur. Bahama Exped., p. 74, 1899.
Disk small; arms very long, slender, much coiled. Upper and under surfaces of the disk and arms covered with thin, delicate, closely imbricated scales, without granules.
Under arm-plates not distinct, except on one or two basal joints.
Arm-spines three, except on a few basal joints, rather long, tapered, simple, thorny at the tip, but not becoming hooked, even on the distal part of the arms.
Teeth stout, obtuse, the lowest not differing much from the rest. No tooth-papille. A row of small granule-like oral papille. Oral and adoral shields well developed.
The type-species lives clinging, by its coiled arms, to a species of slender, pennatulid coral (Scleroptilum gracile V.), in 1362 to 2033 fathoms, off the United States East Coast. (See Amer. Journ. Sci., vol. xxviii, p. 219, 1885; and Annual Rep. U.S. Fish. Com. for 1883, p. 550 (as Hemieuryale tenuispina).
372 A, E, Verrill— North American Ophiuroidea.
Parr IL—A Faunat Caratocgur oF THE KNOWN SPECIES OF OPHIUROIDEA FROM THE West INDIAN REGION.
In the following article the West Indian Zodgeographical region is taken, in its broadest sense, as extending from South Florida and the Bermudas to Yucatan, and to Pernambuco, Brazil.
The extreme geographical range is here given only for those that are known. to extend beyond the West Indian region. When no special localities are given the Carribean Sea and West Indies are to be understood. Those species marked with an asterisk extend north- ward on our coast.
The synonymy of most of the species is given by Mr. Lyman in the Voyage of the Challenger, vol. vy, Ophiuroidea. But dates have been given to indicate the works where the species were first pub- lished. Most of these are given on pages 301 and 383. A fuller bibliography will be given with Part III.
I have added references to the excellent woodcuts in the “ Three Cruises of the Blake,” vol. ii, by Mr. Alexander Agassiz (Bull. Mus. Comp. Zool., vol. xv, 1888); and also to Part I of this series; and to the report on the Ophiuroidea of the Bahama Expedition (1899@), when the species or genera are there described or figured.
Order I. OPHIURZE Mill. and Trosch., 1842. (See this volume, p. 303).
Family, PECTINURIDZ Ver., 1899a, p. 4. (See this volume, p. 303.)
OpuiuraA Lam., 1801.
brevicauda Lym., ’65. 0-35 fath. guttata Lym., 65. 1-10 fath. brevispina Say, 725. 1-122 fath.
Bermudas and Florida to Brazil. *var. olivacea. Florida to Cape Cod. Shallow water. * Holmesii Lym., ’60. 8. Carolina. cinerea Lym., 65. 0-115 fath.
Florida to Abrolhos Reefs, Brazil. pallida Verrill, 18994, p. 7, pl. 11, fig. 3. 110-200 fath. rubicunda Lym., ’65. Littoral. Shallow water.
~
squamosissima Lym., ’65. Littoral?. Shallow water.
bh ‘
A. EF. Verrill—North American Ophiuroidea. 3 appressa Say, °25. 0-20 fath. Bermudas and Florida to Bahia, Brazil. elaps Lym.,’65. Nar. Blake Exped., p. 111, fig. 394. 120-300 fath. OpHioPeza Peters, 1851. Yoldii Ltk., 56. Shallow water. Petersi Lym., ’78. 8-177 fath. Prctrnura Forbes, 1842. angulata Lym., ’83. 88-248 fath. tessellata Lym., ’83. 451 fath. lacertosa Lym., ’83. 159 fath. OpHiop PALE Ljng., 1871. Goesiana Ljn., 71. Lym., in A. Ag., Nar. Blake Exped., pal by figs 303. 38-250 fath. Family, OPHIOLEPIDIDZ Ljung., 1867. OpnioLeris M. and Tr., 1840. paucispina Mill. and Tr., ’42. 3-4 fath. Florida and West Indies to Rio de Janeiro. *elegans Ltk., 59. Charleston, 8. C.; West Indies. 84-30 fath. Opuniozona Lym., 1865. impressa Lym., 65. 69-300 fath. nivea Lym., 75; Nar. Blake Exped., p. 110, fig. 390. 50-424 fath. var. compta Verrill, 1899a, p. 9, pl. m1, fig. 2.
This Vol., p. 303, pl. xiu, figs. 1, la. 110-200 fath. insularia Lym., ’78. 310-315 fath. marmorea Lym., 83. 114-250 fath. clypeata Lym., ’83. 88-157 fath. tessellata Lym., ’78. 60-300 fath. Antillarum Lym., 778. 94-508 fath. dubia Lym., ’78. 539 fath.
OpHioceRAMIS Lym., 1865. Januarii Lym., ’65. 35-100 fath.
West Indies to Rio de Janeiro, Brazil, and Patagonia. albida Lym., 775. 19-100 fath.
West Indies to Rio La Plata. OpHIoTHYREUS Ljng., 1871. Goési Ljn., ’71. 80-300 fath. OpuHiERNus Lym., 1878. adspersus Lym., ’83. 159-1030 fath.
3
Gulf of Bengal and off the Malabar Coast, ~
490-1997 fath. (Keehler).
374 A, FE. Verrill— North American Ophiuroidea.
Opntoeityrua Lym., 1860.
Sasciculata Lym., ’83. 288 fath. abyssorum Lym., ’83. 1097 fath. scutata Lym., 83. 95 fath. tenera Lym., ’83. 124 fath. acervata Lym., 69. 60-350 fath. falcifera Lym., ’69. 200-576 fath. *lepida Lym., West Ind., 425-1242 fath.
East Atlantic, 425-900 fath. Off East Coast U.S., 813-2574 fath.
Ljungmani Lym. Off Bahia, 350 fath. variabilis (?) Lym. (Var. ?) 175-955 fath. ? irrorata Lym., ’78; 783. 1058-1097 fath.
Type, Cape Good Hope, 1900 fath.; Australia, 410 fath. ? convexa Lym., 78; 783. 114-270 fath.
Type, Pacific, 2050-2300 fath.; W. Africa, 2350 fath. OpuiocTEN Lutk., 54.
depressum Lym., 779. 315 fath. Opuiomusium Lym., ’69. eburneum Lym., ’69. 92-500 fath.
var. elegans Verrill, 1899a, p. 12, pl. m1, figs. 1, la. 110-260 fath. *Lymani Wy. Thom., ’73. North Atlantic, 288-2369 fath. Off coast of Europe; off Kast Coast U. States, south to S. Carolina, common ; off Tristan d’ Acunha, 1100
fath.; Pacific, 565-1825 fath.
serratum Lym., 778. 124-1097 fath. cancellatum Lym., ’78 (? var.). Bermuda, 435 fath.
Perhaps the same asthe next. Mr. Lyman’s type was from off Japan. stellatum Verrill, 18994, p. 14, pl. 1, figs. 3, 3a. 110-260 fath.
planum Lym.,’78; Nar. Blake Exped., p. 112, fig. 396. Mediterranean, 4020 meters; Gulf of Bengal, 1520-
1987 fath, (Kehler). W. Indies, 300-955 fath. acuferum Lym., 75. 27-1030 fath. archaster Wy. Thom., ’73. Off Brazil, 1900 fath.
sculptum Verrill, 18994, p. 106, pl. 1, fig. 2, pl. vim, fig. 2. 110-260 fath.
pulchellum Wy. Thom. 150-1675 fath. North of Cape Verde Is.; off Brazil; 5. Atlantic. validum Liung., ’71. 60-1568 fath.
North of Laquedives, 931 fath. (Kcehler). testudo Lym., ’78. 69-508 fath.
A, E. Verrill—North American Ophiuroidea. 375
Opurotipus Lym. Agassizii Lym.,’78; Nar. Blake Exped., p. 115, fig. 401.
100-118 fath. Oruiomastus Lym.
secundus Lym., °78; Nar. Blake Exped., p. 113, fig. 398.
60-1131 fath. OpxHicpHYLLuM Lym.
? petilum Lym.,’79; Nar. Blake Exped., p. 110, fig. 391.
Type, off Fiji Is., 600 fath. W. Indies, 542 fath. OPpHIOPREYN Lym. longispinus Lym., ’78. 60-625 fath.
Opnioconis Lutk., 1869. miliaria Lym.,’78; Nar. Blake Exped., p. 112, fig. 395. Ver. 1899a, p. 17. 163-450 fath.
Family, OPHIOTHRICHIDZ Lijung. Ver., This Vol., p. 304.
OpnHiotTHRix Miill. and Trosch.
*angulata Ayres, ’52. Ver., 1899a, p. 18 (descr.). 0-200 fath. Chesapeake Bay and Bermudas to Rio de Janeiro.
Girstedii Ltk., 56. Ver., 1899a, p. 20, colors, 0-18 fath. lineata Lym., ’60. 0-20 fath. pallida Liung.,’71. 180 fath. Suensonii Ltk., 56. Ver., 1899a, p. 21, colors. 0-262 fath. Bermudas and Florida to Brazil, Lat. 22° S.
Family, OPHIOCOMIDZ Ljung., 1867.
Opniocoma L. Agassiz, ’35.
echinata I.. Agassiz, °35. 0-18 fath. Bermudas and Florida to Brazil. Riisei Liitken, ’56, 0-210 fath. Bermudas and Florida to Brazil. pumila Ltk., 56. 0-100 fath.
Bermudas and Florida to Northern Brazil.
Opuiopsita Lutk. Riisei Ltk., 59. 0-200 fath. Bermudas and Florida to Northern Brazil.
: Family, AMPHIURIDZ Lijung., 67. Ver., This Vol., pp. 305-319.
Opuiactis Lutk. Mitlleri Ltk., ’56. 1-837 fath.
376 A. FE. Verrill—North American Ophiuroidea. f Bermudas and Florida to Abrolhos Islands. : var. guinqueradia Lym. 27-338 fath. % dispar Verrill, 1899a, p. 31, pl. vim, figs. 3-3e. 3-13 fath. 4 * Krebsit Ltk., °56. 1-20 fath. Bermudas and 8. Carolina to Rio de Janeiro. : loricata Lym., ’69. 10-110 fath. 4 Lymani Ljn., °71. 40 fath. . plana Lym., ’69. 10-140 fath. Ampuiura Forbes. Ver., This Vol., pp. 306, 307 (rest.). * Ottert Liung., ’71. 175-1467 fath. Off Portugal and off New England to West Indies. Palmeri Lym., ’82. 100 fath. incisa Lym., ’83. 583 fath. semiermis Lym., 69. 377-539 fath. Hlexuosa Liung. Var. ?, Lym., ’83. 262 fath. Florida to Southern Brazil. grandisquama Lym., ’69. 10-262 fath. lunaris Lym., 78. _ 424-955 fath. Stimpsont Ltk., °59. 10-69 fath.
West Indies to Cape Frio, Brazil.
The two following supposed ‘‘ varities” of antarctic species are recorded by Lyman, but have not been described nor figured, from the West Indian fauna. They were, therefore, omitted from the analytical table (pp. 508-311).
angularis Lym., 78. Var.? 476 fath., Antarctic Ocean. tomentosa Lym.,’78. Var.? 464 fath., Antarctic Ocean. AmpuHIPHoLis Liung., 66. Ver., This Vol., pp. 306, 311.
*tenuispina Ling., 64; 67. 60-487 fath.
Off U.S. East Coast, C. Hatteras to Cape Cod ; North
Europe to Iceland.
*tenera (Ltk.) Ling. 4-200 fath.
S. Carolina to Cape Frio, Brazil.
*gracillima (Stimp.) Ljng., ’67,’71. S. Carolina. West
Ind. Brazil? (as A. Januarii Ling.). Littoral.
* Goési Ling., ’71. 14-280 fath. Bermuda and Cape Hatteras to Antilles.
abnormis (Lym., ’78.) Ver. 101 fath.
Ampuiopia Ver., 1899a, p. 25. This Vol., pp. 306, 312.
repens (Liym., ’75) Ver. Florida, 14 fath. pulchella (Lym., 69) Ver. Florida, 18-39 fath.
*atra (Ltk., 59) Ver. S. Carolina. Littoral. Riisei (Ltk., 760) Ver. Shallow water.
West Indies to Southern Brazil. Littkeni (Ling., ’71) Ver. 10 fath.
A, E. Verrill—North American Ophiuroidea. 377
Ampuioptts Ver., 1899a, p. 25, 1899. This Vol., pp. 306, 314. tumida (Lym., ’78) Ver. 94-321 fath. nereis (Lym., ’83) Ver. 148 fath. Agassizii Ver. This Vol., p. 315, 116 fath. euneata (Lym., ’78) Ver. 159-370 fath. *duplicata (Lym., ’75) Ver. 73-1568 fath.
Mediterranean, 1385-2178 meters; off the Azores, 1300-1850 meters (Keehler).
Stearnsii (Ives) Ver. Shallow.
Verrillit (Lym., °79) Ver. 424-2650 fath. Ampuitimna Ver., 1899a, p. 30, 1899. This Vol., p. 318.
Caribea (Ljng., ’71) West Indies, 300-400 fath.
Perhaps identical with the next.
*olivacea (Lym.) Ver., This Vol., p. 318, pl. xu, figs. 1, la. West Indies, 40-126 fath. West Indies, northward to N. England, in 63-266 fath., beneath the Gulf Stream. Oputocnipa Lym. Ver., This Vol., p. 315 (restr.). scabriuscula Lym., 65.
Florida to Bahia, Brazil. Littoral.
Jilogranea Lym., 775. Shallow. Hemrpxouis Lym. *cordifera Lym., ’65. Littoral. Shallow water.
N. Carolina to Brazil. OPHIOPHRAGMUS Lym. * Wurdemani Lym., ’65. Beaufort, N. C. to West Indies. Littoral. Shallow water.
septus (Lutk., 59) Lym., *65. 47 fath. AmMPHILEPIS Ljung., ’67.
patens Lym., ’79. 2160 fath. OpHIONEMA Lutk., 69.
intricata Ltk., 69. : 180 fath. OPHIONEPHTHYS Lutk., ’69.
limicola Ltk., 69. Littoral ? Shallow water. OPpHIONEREIS Lutk., 59.
reticulata Lutken, 759. 0-94 fath.
‘Bermudas and Florida to Rio de Janeiro, Brazil. OpuiopLtax Lym.
Ljungmani Lym., 75. 80-250 fath. OpwiosticMa Lutk., ’56. isacanthum (Say) Lym., ’65. 0-122 fath.
a
378 A, E. Verrill—North American Ophiuroidea.
OpniocHyTrA Lym. tenuis Lym., ’83. 291-383 fath.
Family, OPHIACANTHIDEA Verrill, 1899a, p. 34. This Vol., p. 319.
Opuriacantsa Mill. and Trosch.,’42. Ver., This Vol., pp. 320-340
(restr. ) vepratica Lym., ’78. 291-600 fath. aspera Lym., 778. 73-400 fath. stellata Lym., ’75. 56-262 fath. pentacrinus Ltk., 69. 74-625 fath. segesta Lym., ’78. 1075 fath. cosmica Lym., ’78. 350-2225 fath. (Ophientodia) scutata Lym.,’78. Ver., This Vol., p. 341,
(deser.). 124-338 fath.
(Ophientodia) pectinula Ver., This Vol., p. 342 (descr.). Station 227, Blake Exped., 573 fath. ( Ophioscalus) echinulatus Lym., ’78. 205-955 fath. Opuiatcma Ver., 1899a, p. 42. This Vol., pp. 326, 331. Nuttingii Verrill, 1899a, p. 46, pl. 1, fig. 2; pl. vim, figs.
1: ile: 200 fath. OPHIACANTHELLA Ver., 18994, p. 39. This Vol., pp. 326, 344. Troscheli (Lym., ’78) Ver. 73-300 fath,
OpHIOMITRELLA Ver., 1899a, p. 43. This Vol., pp. 336, 352. levipellis (Lym.,’83) Ver. This Vol., p. 343 (deser.). 88-124 fath. Opuiouimna Ver., 18994, p. 44. This Vol., p. 345. mixta (Lym.,’78) Ver. This Vol., p. 346. 160-576 fath. Oputopora Ver., 1899a, p. 43. This Vol., p. 345. - Bartletti (Lym., ’83) Ver. This Vol., p. 345. Orutopristis Ver., 18994, p. 47. This Vol., p. 347.
hirsuta (Lym., 75) Ver. 82-955 fath. ensifera Ver., 1899a, p. 47, pl. 1v, figs. 1-1d. This Vol., pl.
XLIH, fig. 4. ? 110-160 fath. cervicornis (Lym., 83) Ver. 208-573 fath.
OpniorrEtaA Ver., 1899a, p. 40. This Vol., pp. 333, 347.
lineolata (Lym., ’83) Ver., 18994, p. 51. , 110-208 fath. sertata (Lym., ’69) Ver., 1899a, p. 54. This Vol., p. 348.
123-411 fath,
Ampuipsita Ver., 1899a, p. 55. This Vol., pp. 333, 348. maculata Ver., 1899a, p. 55, pl. 11, figs. 4, 4a. This Vol., p. 848; pl. xu, figs. 5, 5a. 200 fath. JSulva (Lym., ’78). 13-175 fath.
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—
er
A. E. Verrill— North American Ophiuroidea. 379
OpnHiomiTRA Lym., Ver. This Vol., pp. 349, 350; restr. valida Lym., ’69. Ver., This Vol., p. 353 descr.
10-1105 fath. ornata Ver., 1899a, p. 58, pl. v, figs. 1, la. This Vol., p.
350, pl. xin, fig. 3. 100-260 fath. OpuHiopiintTHaca Ver. This Vol., p. 351. dipsacos (Lym.,’78) Ver. This Vol., p. 351. 390 fath.
incisa (Lym., 83) Ver. This Vol., p. 351. 334-508 fath.
chelys (Lym., ’78) Ver., This Vol., p. 352. 1124-1530 fath. Opuiocamax Lym., Ver. This Vol., p. 354.
hystrix Lym., ’78 ; Nar. Blake Exped., p. 110, fig. 392.
Ver., This Vol., p. 395. 114-300 fath. austera Verrill, 18994, p. 60, pl. vt, figs. 1, 1@; pl. vu, fig. 2. This Vol., p. 355, pl. xii, fig. 2. 110-200 fath.
Jasciculata Lym.,’83. Ver. This Vol., p. 355. 180-250 fath. Off Andaman Islands, 130-250 fath. (Keehler). OpHioTHAMNUS Lym., ’69. vicarius Lym., ’69. 15-611 fath. exiguus (Lym.,’78) Ver. This Vol., p. 353. 84-400 fath. OpuHioLeBes Lym., ’78.
humilis Lym., ’69. 125-324 fath.
claviger (Ljng.) Lym. (Var. ?), 83. 524 fath. OPpHIOBLENNA Lutk.
Antillensis Ltk., 59. W. Indies. Shallow water. Opnioroma Lym.
coriacea Lym., 83. 1242 fath.
Subfamily, OPHIOCHONDRINZ Ver., This Vol., p. 355.
OpuiocHonprus Lym., 69. Ver., This Vol., p. 356 (restr.).
convolutus Lym., ’69. 80-400 fath.
erassispinus Lym.,’83. This Vol., p. 356. 229 fath.
gracilis Ver., 1899a, p. 64. 100-260 fath. OpHIOCHONDRELLA Ver., This Vol., p. 355.
squamosus (Lym) Ver. This Vol., p. 355. 88-250 fath.
Family, OPHIOSCOLICIDZ Ver. (See This Vol., p. 357.)
OpuHioscoLex Mill. and Trosch., 1842. *purpureus Dub. and Koren., ’44 (var. ?) West Indies, 110 fath. Northern Europe to Norway, 64-767 fath. Fragilis Ver.,’99. This Vol., p. 358. 82 fath. tropicus Lym., ’78. 390 fath.
380 A. FE. Verrill— North American Ophiuroidea.
OruroscrasMA Lym., 778.
granulata Lym., ’83. ‘ * 96-100 fath. AsTROGERON Ver., This Vol., p. 359.
supinus (Lym.) Ver. 200-464 fath. OpuioByRsELLA Ver., This Vol., p. 358.
serpens (Lym.) Ver. This Vol., p. 358. 69 fath. Opniosyrsa Lym.
Perrieri Lym., ’83. 288 fath.
Family, OPHIOMYCETIDZ Verrill, This Vol., p. 359. Subfamily, OPHIOMYCETIN A! Ver., This Vol., p. 360 (descr.).
Opruiomyces Lym., ’80. mirabilis Lym., ’68. 237-422 fath. Jrutectosus Lym., 69; Nar. Blake Exped., p. 111, fig. 397. 77-288 fath.
Family, OPHIOHELIDZ Ver., This Vol., p. 361.
OpnionHELus Lym., ’80. umbella Lym.,’80; Nar. Blake Exped., p. 116, figs. 202, 203. Barbadoes, 82 fath.
Family, OPHIOMY XID Ljng.; Ver., 1899a, p. 65; This Vol., p. 361.
Orniomyxa Miill. and Trosch., 42. Jlaccida Ltk., ’59. Ver., 1899a, p. 65, colors. 0-175 fath. Bermudas and Florida to Bahia and the Abrolhos Reefs, Brazil. tumida Lym., ’83, Ver., 1899a, p. 67, pl. 11, fig. 5; This
Vol., pl. xxi, figs. 3, 3a, 30. 13-300 fath. brevicauda Ver., 1899a, p. 66, pl. m1, fig. 3; This Vol., pl. xx, figs. 4-4d. 110-200 fath.
OpuiopERA Ver., 1899a, pp. 65, 67. This Vol., p. 362. Stimpsoni (Lym., 75) Ver., 1899a, p. 67, pl. 11, figs. 4, 4a; This Vol., p. 362, pl. xu1, figs. 2—2c. 60-240 fath.
Family, HEMIEURYALIDZ Ver., 1899a, p. 70; This Vol., p. 363.
HermIEuRYALE Von Mart., 67. pustulata Von Mart., 67. Ver., This Vol., p. 363. 74-180 fath.
A. E. Verrill—North American Ophiuroidea. 381
Oruioptus Ver., 1899a, p. 70; This Vol., p. 365. tuberculosus (Lym.) Ver., 1899a, p. 71, pl. 1, figs. 1-14; This Vol., p. 365, pl. xx111, figs. 6-6d. 96-200 fath.
StesBEera Lym., ’78.
murrhina Lym., 778; Nar. Blake Exped., p. 114, fig. 399. Ver., 1899a, p. 72, pl. 11, figs. 1, 1a, (young, descr.) ; Tins Vol pe3se65, pl. xu, fig. 7. 88-422 fath. Young, 3-200 fath.
Family, OPHIOBRACIONTIDZ: Ver., This Vol., p. 366.
OPHIOBRACHION Lym., ’83. uncinatus Lym., ’83. Off Cuba, 250 fath.
Order I1.—EURYALZ Mill. and Trosch., 1842.
(See This Vol., p. 366.)
Family, GORGONOCEPHALIDZ Ver., 1899a, p. 88 (restr.) ; This Vol., p. 367.
Astropuyton L. Agassiz, 735. *muricatum Ag. (costosum Lym.) Ver., 18994, p. 84 (descr.) Charleston, S. C., Ives, 1889, p. 178. 1-18 fath. Krebsii CErst. and Ltk., 56. 50-125 fath. Cecilia Ltk., 56; Blake Exped. Narrative, i, p. 310, fig. 388. 3-124 fath. GorGonocePHatus Leach, 1815. *aborescens Agassiz, 739. W. Indies? Mediterranean.
cacaoticus Lym., ’74. 20 fath.
mucronatus Lym., 69. Ver., 18994, p. 85 (descr.) 80-288 fath.
Family, ASTROCHELIDZ Ver., 1899a, p. 79; This Vol., p. 369.
Astrocnipa Lym., ’72. isidis Lym., ’72; Nar. Blake Exped., p. 115, fig. 400. Ver., 1899a, p. 83. 56-120 fath. Astrocompnuts Lym., ’69.
vallatus Lym., 69. Ver., 1899a, p. 80 (deser.). 80-337 fath.
rudis Ver., 18994, p. 82, pl. vu, figs. 1, 1a (descr.). 116-200 fath.
Trans. Conn. AcapD., Vou. X. OctoBER, 1899.
26
382 A, #. Verrili—_North American Ophiuroidea.
AsrroporpPa (Ersted and Lutk., 56. *annulata CErst. and Ltk., ’56. 20-163 fath.
_ Off Cape Hatteras and Chesapeake Bay, 48-167 fath. affinis Ltk., °59. 50 fath.
Family, ASTROSCHEMIDZE Ver., 1899a, p. 76; This Vol., p. 370.
AstroscHEMA (Ersted and Lutk., ’56.
oligactes Ltk., ’56. 69-288 fath. arenosum Lym., ’78. 124-805 fath. tenue Lym., 775. 88-124 fath. brachiatum Lym., ’79. 270-435 fath. leve Lym., ’75. 56-262 fath. suleatum Ljn., °71. 200-320 fath. intectum Lym., °78. 175-200 fath.
Nuttingii Ver., 18994, p. 77, pl. vu, figs. 3, 3a. 105-125 fath. OpniocREAs Lym., 69. lumbricus Lym., ’69. 60-580 fath. spinulosus Lym., ’83; Nar. Blake Exped., p. 110, fig. 389. 116-288 fath. ceedipus Lym., ’79 (Var. ?). 580 fath.
Family, ASTRONYCIDZE Ver., 1899a, p. 74; This Vol., p. 370.
Astronyx Miill. and Trosch., 742. Lymani Verrill, 1899a, p. 74 (descer.), pl. vi, figs. 4-4e; This Vol., p. 371 (descr.), pl. xLu, figs. 6-6ce. 200-980 fath.
A. E. Verrill— North American Ophiuroidea. 383
BIBLIOGRAPHY.
Principal Works relating to the West Indian Ophiuroidea.
Many general works, which contain West Indian species, have been omitted from this list, as well as several minor articles, not of special importance here, or else not quoted in the preceding articles. Additional Bibliography will be given with Part III.
1725.—Sloane, Hans. Voyage to Jamaica, ii, 272.
1801.—Lamarck, J. B. Systeme des Anim. sans vertébres.
1815.—Leach, W. E. Zodlogical Miscellany, II.
1816.—Lamarck, J. B. Histoire des Anim. sans vertébres, 1st ed.
1825.—Say, Thomas. Journal of the Academy of Natural Science at Philadel- phia, v, p. 145.
1830.—Bosc, S. A. G. Histoire Nat. des Vers, ii, Suites a Buffon.
1840.—Miiller, J., & Troschel, F. H. Gattungender Ophiuren. Wiegmann’s Archiy., vi, p. 326.
1842.—Miiller, J.. & Troschel, F. H. System der Asteriden.
1850.—Duchassaing, P. Animaux Radiaires des Antilles, p.4. (For a review of this paper, see Lyman, 1872.)
1851.—Ayres, W. O. Proceedings Boston Soc. Nat. Hist., iv, p. 133; iv,
i 1852. p. 249.
1852.—Stimpson, Wm. Two new species of Ophiolepis [Amphiura] from the southern coasts of the United States. Proc. Boston Soc. Nat. Hist., iv, p. 224.
1856.—Liitken, Chr. Fr. Oversigt over de Vestindiske Ophiurer. Naturhist. Foren. Vidensk. Meddelelser.
1859.—Litken, Chr. Fr. Additamenta ad Historiam Ophiuridarum. Pt. ITI. 92 pp., 5 plates. Kgl. Danske Videnskab. Selskabs Skifter, 5'° Rekke, Naturvidensk. og mathem. Afdeling, V.
1860.—Lyman, T. Descriptions of New Ophiurans. Proceed. Boston Soc. Nat. Hist., vii, Feb. and June, pp. 195, 252.
1862.—Dujardin et Hupé. Histoire Naturelle des Zodphytes. Echinoderms, Suites a Buffon.
1864.—Ljungman, Axel VY. Tilligg till kiinnedomen af Skandinaviens Ophiu- rider. Ofvers. af K.Vet.-Akad. Foérh. for 1863, No. 7, with one plate.
1865.—Lyman, Theod. Ophiuride and Astrophytide. Illustrated Catalogue Museum Comparative Zodlogy, I, 200 pages, 2 colored plates.
1866.—Ljungman, Axel V. Om niigra nya arter af Ophiurider. Ofversigt af Kongl. Vetenskaps—Akademiens Férhandlingar. No. 6, pp. 163-166.
1867.—Verrill, A. E. On the Geographical Distribution of the Echinoderms of the West Coast of America, and Comparison of the tropical Echinoderm Faunz of the East and West Coasts of America. Trans. Conn. Acad. Sciences, i, pp. 323-339.
1867.—Ljungman, Axel V. Ophiuroidea Viventia hue usque cognita eneumerat. Ofversigt af Kongl. Vetenskaps-Akademiens Férhandlingar for 1866, No.9.
384 A. E. Verrill—North American Ophiuroidea.
1867.—Martens, E. Von. Monatsbericht der Konig. Akad., Berlin, p. 345 and 481.
1868.—Verrill, A. E. Notice of the Corals and Echinoderms collected by Prof. C. F. Hartt at the Abrolhos Reefs, Province of Bahia, Brazil, 1867. Trans. Conn, Acad. Sciences, i, pp. 351-371, 1 pl.
1869a.—Liitken, Chr. Fr. Additamenta ad Historiam Ophiuridarum. Part ITT. Kgl. Danske Videnskab. Selskabs Skrifter, 8, Bd. ii, pp. 24-101.
1869b.—Litken, Chr. Fr. Synopsis generum Ophiuridarum verarum. (Forms part of the preceding work,pp. 87-100.)
1869.—Lyman, Theodore. Preliminary Report on Ophiuride and Astrophy- tid dredged in deep water between Cuba and the Florida Reefs, by L. F. de Pourtales. Bulletin Mus. Comp. Zo6l., Vol. i., No. 13.
1871.—Ljungman, Axel V. Forteck. ofver uti Vestindien af Dr. A. Goés samt under korvetten Josefinas Exped. i Atlantiska Oceanen samlade Ophiu- rider. Ofvers. Kong. Vetenskaps-Akad. Férhandlingar. No. 6, pp. 615- 658.
1871.—Lyman, Theodore. Illustrated Catalogue of the Museum of Compara- tive Zodlogy. No. VI. Supplement to the Ophiuride and Astrophy- tide. 17 pp., two plates. Memoirs Mus. Comp. Zoélogy, II.
1872.—Lyman, Theod. Note sur les Ophiurides et Euryales qui se trouvent dans les collections du Mus. d’Hist. Naturelle de Paris. Ann. Sci. Nat., xvi, 8 pp. (Contains a synonymical list of the species described by Duchassaing, 1850.)
1872.—Verrill, A. E. Radiata from the Coast of North Carolina. Brief. Cont. to Zoél., No. 22. Amer. Journ. Sci., vol. iii, p, 439.
1872.—Liitken, Chr. Fr. Ophiuridarum novarum vel minus cognitarum descrip- tiones nonnulle. Oversigt Kongl. Danske Vid. Selskabs Forhand., 84 pp., 2 pl., with Résumé in French.
1875.—Thomson, C. Wyville. The Depths of the Sea.
1874.—Lyman, Theod. Ophiuride and Astrophytide, Old and New, Bull. Mus. Comp. Zoél., iii, p. 221, 51 pp., 7 pl., part anatomical.
1875.—Lyman, Theod. Zodlogical Results of the Hassler Expedition, II. Ophiuride and Astrophytide, Ilustrated Catalogue Museum Compara- tive Zoélogy, viii, 34 pp., 5 plates. Mem. Mus. Comp. Zodl., iv.
1877.—Lyman, Theod. Mode of Forking among Astrophytons, Proc. Boston Soc. Nat. Hist., xix.
1877.—Thomson, C. Wyville. Voyage of the Challenger. The Atlantic.
1878a.—Lyman, Theod. Reports on the Operations of the U. 8S. Coast Survey Steamer ‘‘ Blake,” Ophiurans and Astrophytons. Bull. Mus. Comp. Zool., v, p. 217, 21 pp., 3 pl.
1878b.—Lyman, Theod. Ophiuride and Astrophytide of the Exploring Voyage of H. M. S. Challenger, under Prof. Sir Wyville Thomson. Part I. Bull. Mus. Comp. Zoél., v, pp. 65-168, x plates.
1879.—The same. Part II. Bull. Mus. Comp. Zodl., vi, pp. 17-838, pl. x-xix. F
1879.—Rathbun, Richard. A List of the Brazilian Echinoderms, with Notes on their Distribution, ete. Trans. Conn. Acad. Sci., v, p. 189.
eens ARNE A Sie ea Ss
a a
e r 4 | : :
A. #. Verrill—North American Ophiuroidea. 385
1880a.—Lyman, Theod. Anniversary Memoirs of the Boston Soc. of Natural History.
1880b.—Lyman, Theod. Preliminary List of the known genera and species of living Ophiuride and Astrophytidze. Cambridge, Mass., 45 pp., 4to.
1882.—Lyman, Theod. The Voyage of H. M.S. Challenger. Zodlogy, v, Report on the Ophiuride dredged during the years 1873-6. pp. 368, 48 plates, 4to. ‘
1883.—Lyman, Theod. Reports on the Results of Dredging, under the super- vision of Alexander Agassiz, in the Caribbean Sea in 1878-79, and along the Atlantic Coast of the United States during the Summer of 1880, by the U. S. Coast Survey Steamer ‘‘ Blake.” Report on the Ophiuride. Bull. Mus. Comp. Zo6l., x, p. 227, pp. 60, 8 pl.
1888.—Lyman, Theod., in Agassiz, Alexander. Three Cruises of the Blake, vol. ii, pp. 109-116, with cuts. Bull. Mus. Comp. Zoél., vol. xv.
1889.—Ives, J. E. Catalogue of the Asteroidea and Ophiuroidea in the Collec- tion of the Acad. Nat. Sciences of Philadelphia. Proc. Acad. Nat. Sci., Philad., p. 169.
1890.—Ives, J. E. Echinoderms from the Northern Coast of Yucatan, and the Harbor of Vera Cruz. Proc. Acad. Nat. Sci. Philad., pp. 317-340, 1 plate.
1891.—Ives, J. E. Echinoderms from the Bahama Islands. Proc. Acad. Nat. Sci. Philadelphia, p. 337.
1895.—Nutting, C. C. Narrative and Preliminary Report of Bahama Expedi- tion. Bulletin from the Laboratories of Nat. Hist. of the State Univ. of Iowa, vol. iii.
1896.—Kcehler, René. Note Prélim. sur les Ophiures rec. pend. les Camp. de VHirondelle. Mem. Soc. Zool. de France, ix, pp. 203-213.
1897.—Keehler, René. Echinodermes recueillis par ‘‘l’Investigator” dans VOcéan Indien. Ann. Sci. Nat., Zool. et Paléont., pp. 277-370, pl. v—ix.
1898.—Clark, H. L. Notes on the Echinoderms of Bermuda. Annals New York Acad. of Science, xi, pp. 407-413.
1899a.—Verrill, A. E. Report on the Ophiuroidea collected by the Bahama Expe- dition in 1893. Bull. Labor. Nat. Hist. of the State Univ. of Iowa, v, No. 7, pp. 1-86, 8 plates.
1899b.—Verrill, A. E. North American Ophiuroidea. Part I. Revision of cer- tain Families and Genera of West Indian Ophiurans. Trans. Conn. Acad. Sciences, x, part 2, pp. 301-371.
1899c.—Verrill, A. E. The same. Part If. A Faunal Catalogue of the known Species of West Indian Ophiurans. Trans. Conn. Acad. Sciences, x, part 2, pp. 372-386, pl. xlii, xliii.
386 A. E. Verrili— North American Ophiuroidea.
EXPLANATION OF THE PLATES. Puate XLII. Figure 1—Amphilimna olivacea (Lym.) Ver., p. 318. Under side of a part of
the disk and the proximal part of an arm. x7.
Figure la—The same. A row of spines from the middle of anarm, x 18.
Figure 2—Ophiodera Stimpsoni (Lym.) Ver., p. 862. Under side of a part of the disk and of aray. x7.
Figure 2a—The same. A row of spines. x 28.
_ Figure 2b—The same. Under side of the distal part of an arm. x 7.
Figure 2c-—The same. One of the teeth. x18.
Figure 83—Ophiomyxa tumida Lym., p. 380. Under side of a part of the disk andofanarm. x.
Figure 8a—The same. A row of spines from the middle of anarm. x12.
Figure 4—Ophiomyxa brevicauda Ver., p. 380. Under side of a part of the disk and ofanarm. x9.
Figure 4a—The same. One of the teeth. x 28.
Figure 5—Undetermined Ophiuran (Young?). Nearly vertical view of two joints from the middle of anarm. x 28.
Figure 5a—The same specimen. Side view of two joints from the distal parts ofanarm. x28.
Figure 6—Astronyx Lymani Ver., p. 377. Type. Under side of a part of the disk and of anarm. x7.
Figure 6a—The same specimen. Side view of three joints from the middle part ofanarm. x9.
Figure 6b--The same. Hook-shaped spines, more enlarged.
Figure 6c—The same. Hooks from the distal part of an arm.
Figure 7—Sigsbeia murrhina Lym. Young, p. 365. Under side of a part of the disk and of two arms. x6.
Puate XLII.
Figure 1—Ophiozona nivea Lym., var. compta Ver., p. 303. Type. Upper side of the disk and bases of the arms. x 4.
Figure la—The same specimen (297, Blake Exped.). Under side of the disk. x dls.
Figure 2—Ophiocamax austera Ver., p. 30d. Type. Under side of a part of the disk and of an arm; m, madreporic plate. x5.
Figure 83—Ophiomitra ornata Ver., p. 830. Type. Under side of a part of the disk and of an arm; m, madreporic plate. x 5.
Figure 4—Ophiopristis ensifera Ver., p. 347. Type. Under side of a part of the disk and of anarm. x6.
Figure 5—Amphipsila maculata Ver., p. 848. Under side of a part of the disk and base of anarm. x7.
Figure 54a—The same. <A row of spines from the middle of anarm. x 14.
Figure 6—Ophioplus tuberculosus Ver., p. 365. Dorsal side. x5.
Figure 6a—The same. Underside. x7.
Figure 66—The same. Profile view of a part of the disk and coiled arms. x7.
Figure 6c—The same. Side view of a part of the middle of an arm. x5,
Figure 6d—The same. Dorsal view of a more distal part of anarm. x95. 7
ths EAP ENS eae IELTS Oe rer ie Nina CA eee
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VIII.—Tue Hawaiuan Hepatic or THe TRIBE JUBULOIDEZ.— By ALEXANDER W. Evans.
A FEw Hepatice from the Hawaiian Islands were collected by Menzies in 1793. They consist of small specimens and of fragments picked from other plants and are for the most part in the herbarium of Sir William J. Hooker, now preserved in the collections of the Royal Gardens at Kew. In the early decades of the present century additional small collections were made by Beechey, by Gaudichaud, and by Meyen, during their voyages of exploration, and these, together with the Menzies plants, are the Hawaiian Hepatic referred to in the Synopsis Hepaticarum of 1844-47. Scarcely thirty species, most of them from the island of Hawaii, are mentioned in this volume, which gives us, therefore, little idea of the richness of the hepatic flora of the Islands.
About thirty years ago another small collection was made by the late Dr. William Hillebrand. His attention, of course, being chiefly devoted to the phanerogams and pteridophytes, to the knowledge of which he made important and well known contributions, the Hepatic which he gathered were somewhat fragmentary in char- acter and consisted mainly of large and conspicuous species, to which oceasionally smaller forms remained attached. The collection, never- theless, included a number of undescribed plants. Some of the specimens were sent for determination to Mr. C. F. Austin and others to Mr. William Mitten, and, as these two writers worked independently of each other, certain of the new species received two names apiece. The published accounts of Hillebrand’s plants appeared between 1869 and 1876. In 1872, the Swedish botanist, Dr. Johan Angstrém, published a list of the Hawaiian Hepatic collected twenty years previously by Professor N. J. Andersson, during the voyage around the world of the frigate Eugenie; and, in 1874, Austin published a list of the species collected by Messrs. Mann and Brigham in 1872. In both of these lists new species are described and the synonymy is complicated by giving new names to certain previously described species. Most of the plants in these three col- lections came from the island of Oahu.
The first thorough and systematic collection, however, was the one made in 1875 and 1876 by Mr. D. D. Baldwin, nearly all of whose specimens came from the island of Maui, Mr. Baldwin sent
388 A.W. Lvans— Hawaiian Hepatice of the Tribe Jubuloidee.
his plants to Professor D. C. Eaton, who forwarded sample-specimens of them to Mr. Austin for determination. Austin described several of the new species from this collection in 1879 and left several others in manuscript. In 1892, the present writer compiled a.provi- sional list of the species from the Islands, based upon Baldwin’s collection and depending largely on the determinations made by Austin.
In 1896 Herr Stephani described two of Gottsche’s manuscript species from specimens collected by Didrichsen, and in the follow- ing year, published a revised list of the Hawaiian Hepatic, adding several new species from the collections made by Mr. A. A. Heller in 1895, and several others from specimens in the herbaria of Drs. Askenasy and Spruce. He also ascribed the various Lejeuneee to their respective genera, as these are at present understood.
During the summers of 1897, 798 and ’99, large and important collections were made by Mr. C. M. Cooke, Jr., mainly on the islands of Oahu and Kauai. These collections have brought to light several new and interesting species and have added much to our knowledge regarding the distribution of forms previously known. Mr. Cooke’s specimens have furnished the material for most of the descriptions and illustrations in the present paper.
Even now, however, our knowledge of the Hawaiian hepatics is doubtless far from complete: the collections of recent years have been made almost entirely on the islands of Oahu, Maui and Kauai, and we know little more about the species growing on the large island of Hawaii than was known fifty years ago. From the island of Molokai also, which seems favorable for these plants, only three species have been reported. It is quite possible that a systematic collection on Hawaii and a careful search on the other islands, par- ticularly for the minuter forms, would more than double the number of species which we now know.
The present paper includes a part only of the leafy Hepatice, the Jubuloidee of Schiffner. In its preparation, I have been able, through the kindness of correspondents and the curators of herbaria, to examine the original specimens of nearly all of the Hawaiian species. ‘Those who have given me the most assistance and to whom I would express my grateful acknowledgments, are the following: Mr. W. H. Pearson, Professor A. G. Nathorst, Mr. William Mitten, Herr F, Stephani, Professor Victor Schiffner, Mr. W. B. Hemsley, Dr. P. Hennings, Professor L. M. Underwood and Mr. A. Gepp.
—s»-
A. W. Hvans— Hawaiian Hepatice of the Tribe Jubuloidee. 389
The tribe Jubuloidew, as defined by Schiffner,* is the equivalent of the subtribe Jubulex of the Synopsis Hepaticarnm} and of the tribe Jubulez of Spruce.{ It is without doubt the most natural assem- blage of forms among the leafy Hepatice; it isin fact so natural that Spruce did not hesitate to place it in contrast to his tribe Jungermanniex, which included all the remaining acrogynous Jun- germanniacee and the anacrogynous Jungermanniacexe or Metzgeri- acex§ as well. Schiffner, on the other hand, looks upon the group as one of the minor divisions under the acrogynous Jungermanniacer, equal in systematic value to the Ptilidioidex, the Scapanioidex, or any of the other five tribes which he recognizes. The morphological papers of Leitgeb|| would, of course, prevent a wide acceptance of Spruce’s view, and the disposition made by Schiffner is more in accord- ance with the views of most recent hepaticologists and apparently with the facts. A reason for the unusual division advocated by Spruce is perhaps to be found in the extraordinary development of the Jubuloidee in the tropics. Among the Hepatice of equatorial South America, to whose collection and study Spruce devoted many years of his life, more than half of the species which he found belonged to this group; in the Hawaiian Islands, only about a quarter of the known species are Jubuloidez, but it is probable that the higher proportion will be reached both here and in other tropical countries, when their hepatic floras shall have been more thoroughly investigated.
Although so natural for a hepatic group, the characters of the Jubuloidez, particularly those drawn from the gametophyte, are somewhat difficult to define. This is partly because the sexual plant exhibits considerable variation within the group, and partly because several of its most striking and constant peculiarities recur in other genera, sometimes widely removed from the Jubuloidex. In the first place the gametophyte is very variable in size ; from the smaller species of Cololejeunea, which are often only a few millimeters long, we may pass by all gradations to the larger Frullanie, some of which form drooping tufts a half meter in length. The more essen- tial characters drawn from the vegetative organs are likewise just as variable, although they show certain peculiarities which are fairly
* Engler & Prantl, Natiirl. Pflanzenfam. i*, 116. 1893.
+ L. c. 283. 1845.
+ Hep. Amaz. et And. in Trans. & Proc. Bot. Soc. Edin, xv. 1. 1885. $ Underwood, Bot. Gazette, xix, 356. 1894.
| Untersuch. tiber die Lebermoose, 1874-1881.
390 A.W. Hvans— Hawaiian Hepatice of the Tribe Jubuloidee,
constant throughout the group. The leaves are complicate-bilobed,* the antical lobes being the larger and incubous in their arrangement, while the smaller postical lobes or “lobules” are usually wholly or partially inflated and serve as water-sacs. Except in the two genera Cololejeunea and Metzgeriopsis, underleaves are always present; usually there is only one underleaf developed for each pair of side- leaves, but in the genera Diplasiolejeunea and Colurolejeunea there is an underleaf for every side-leaf, a peculiarity found nowhere else among the Hepatic. The branches, sexual as well as vegetative, are invariably lateral. The female inflorescence, which is of course always terminal, is sometimes borne on the main stem or on a prin- cipal branch, sometimes on a short, special branch. The number of archegonia is always small, almost never exceeding four. The peri- cheetial bracts, like the leaves, are complicate-bilobed, but they are usually larger and their lobulesf are never inflated; the bracteoles{ likewise are larger than ordinary underleaves. A perianth is always present and is entirely free from the bracts; it is of the hypogonian- thous type, but its keels are not always distinct; in the upper part, it is abruptly contracted into a more or less distinct beak with a small opening, and it becomes lacerated when the capsule is extruded. The calyptra is free. The antheridia are borne, usually in pairs, in the axils of inflated, complicate-bilobed, perigonial bracts, whose lobes are subequal in size; these bracts are imbricated, often very densely so, and occur in clusters of from two to many pairs, sometimes in the course of an ordinary branch, sometimes on a short specialized branch. The corresponding bracteoles are usually smaller than ordinary underleaves and are often absent from the upper part or even from the whole extent of the antheridial spike. The rhizoids of the Jubuloidex are sometimes abundant and sometimes very scanty. They are borne in clusters, each cluster arising from the lower surface of an underleaf, close to the base. In the genus Colo- lejeunea, the clusters of rhizoids are found on the postical surface of the axis, in the position where underleaves would naturally be expected.
The sporophyte, although so much simpler than the gametophyte, affords important and constant characters. The stalk, instead of
* A single exception is found in the monotypic genus Myriocolea Spruce, of South America, in which no lobule is developed.
+ In certain genera the lobules are small and indistinct and are sometimes entirely obsolete.
| The bracteoles are absent in the genera without underleaves; they seem to be absent also in most species of Colurolejeunea.
A.W. Evans— Hawaiian Hepatice of the Tribe Jubuloidew. 391
being of the same diameter throughout, as in most hepatics, broadens out above into a disc of the same color as the capsule-wall and several cells thick in the middle part.* On the circumference of this dise the four valves of the capsule are inserted, and it appears, therefore, as if the capsule were not split to the base. The valves themselves are usually two-cells thick and each bears on its inner surface close to the apex a cluster of truncate unispiral elaters. The cells of the valves do not show the peculiar band-like thickenings usually found in the cells of the inner layer of the capsule-wall, but are merely irregularly thickened.
The Jubuloidez fall naturally into two well-marked subtribes: the Frullaniez and the Lejeuneer.
The Frullaniee are almost never of a bright green color but are tinged with brown or red, sometimes so deeply so as to be nearly black. The leaf-lobes are ovate to orbicular “in shape, more or less convex, and usually with entire margins. The lobules are in the form of inflated hood-shaped or club-shaped sacs, attached to the postical margin of the lobe close to the base. The opening of the sac is usually near the point of attachment and is directed backwards; in a few species, however, the lobule is reflexed and the opening is consequently directed forwards. Sometimes only a part of the lobule is inflated, and sometimes, particularly in moist localities, the lobule appears as a narrow lanceolate lamina and is not inflated at all. A third part of the leaf, the stylus, is usually distinct in this subtribe; it is situated on the inner side of the lobule and is inserted, partly at least, on the stem. The stylus is usually in the form of a minute, subulate process, and may be looked upon as a part of the postical lobe, or as an appendage to it. Underleaves are always present and are almost invariably bifid.
A branch in the Frullaniexe develops from the whole of the pos- tical half of a segmentt and replaces the lobule of the leaf beneath whose lobe it is situated.{ Corresponding with this method of branching, the first leaf of a branch is an underleaf; the second, a side-leaf turned toward the apex of the main axis; the third, a side-
* Cf. Spruce, Hep. Amaz. et And. 5. 1885.
+ This type of branching is found in many genera of Hepatice ; it is particu- larly clear in Porella, Lepidozia, and others with incubous leaves, but is to be made out also in various genera with succubous leaves.
¢ An exception to this rule is found in the innovations and sometimes in the antheridial branches of Jubula, which conform to the type of branching described for the next subtribe.
392 A.W. Evans—Hawaiian Hepatic of the Tribe Jubuloiden.
leaf turned away from the apex; the fourth, a second underleaf, and so on.
The Lejeuneex are pale or bright green in color and are some- times tinged with brown or black, but never with red. The leaf- lobes vary from orbicular to lanceolate in shape, and their margins show all gradations from entire to laciniate-dentate. With rare exceptions, the lobule is attached to the lobe by a broad fold and to the axis by a long, almost longitudinal line of insertion. The lobule is strongly convex when seen from the postical surface, and its free margin is either involute or appressed to the stem, so as to form with the lobe a fairly tight water-sac. In moist situations this becomes unnecessary, and the lobule is usually more or less reduced, some- times so much so as to be hardly distinguishable. Underleaves, when present, are sometimes undivided and sometimes bifid.
A branch in the Lejeunee springs from the basiscopic part of the postical half of a segment,* and the lobule of the corresponding leaf is normally developed. The branch remains close to this leaf and is apparently borne just behind it.t Corresponding with this method of branching, the first leaf of a branch is an underleaf; the second, a side-leaf turned away from the apex of the main axis; the third, a side-leaf turned toward the apex, and so on. According to Leit- geb, the first three leaves of a branch do not appear as such, but remain united as a sheath, enclosing the apical region of the branch, which may or may not develop farther.
A peculiar type of branch is the “ innovation,” which is found in nearly every genus of the Lejeuneeex. These innovations are borne just behind one or both of the perichetial bracts, which are here reduced to a single pair. Sometimes an innovation is small and
b
simple, but it is more frequently as large as the axis bearing it, whose own growth has been stopped by the formation of the arche- gonium. In such a case, the innovation itself, while still short, often gives rise to a second flower and a new innovation. This mode of growth is sometimes repeated several times, the result being a complicated and characteristic flower-cluster. In case an innovation is developed behind only one of the bracts, it is not unusual to have the bracts and perianth pushed to one side, where they seem to assume a lateral position, the innovation apparently
* An exception is found in the genus Stictolejeunea, where the branches are borne as in the Frullaniew. Cf. Sprace, Hep. Amaz. et And. 307.
| This type of branching is found also in the genera Radula and Scapania. Of. Leitgeb, Unters. tiber die Leberm., ii, 29.
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A.W. Evans— Hawaiian Hepatice of the Tribe Jubuloidew. 393
being a continuation of the main axis. This condition may be readily distinguished from that in which a flower is really borne on a short lateral branch by the order of the leaves on the apparent axis beyond the flower : in case this is the true axis, the first leaf will be on the side next the flower ; in case it is an innovation, the first leaf will be on the side away from the flower. Very striking examples of each condition may be found in the genus Cololejeunea.
Subtribe I. FRULLANIE A.
The Frullaniez include only two genera, Frudlania and Jubula, of which only the most important characters will be enumerated here.
In Frullania, both antical and postical lobes are attached to the axis by very short, almost transverse lines of insertion, the attach- ment in the case of the postical lobe being limited to the base of the stylus. The leaf-lobe, beneath which a branch is situated, is similar to the others in position and is attached to the main axis rather than to the branch. The first underleaf of the branch seems to function as the lobule of this lobe, and their lines of insertion come very close together. This underleaf is similar to other underleaves,* but is often distorted in position; and sometimes one of its divisions, the one next the lobe, is inflated like an ordinary lobule. Between this underleaf and the lobe, we occasionally find a small leafy structure, now appearing as a minute, subulate process, now as a small, more or less rudimentary sac. According to Leitgebt this interposed body is derived from the first postical segment of the branch, which normally gives rise to the underleaf alone; it may be looked upon, therefore, as a supplementary part of this underleaf. Properly speaking, innovations are never present in Frullania,; that is, no branches are ever developed just behind the perichetial bracts. It is not unusual, however, to find branching a little farther back from the perianth, and such a branch, which arises in the usual way, may apparently continue the axis, as in the Lejeuneex. The stalk of the capsule is more complicated than in Jubula or the following subtribe, and is usually built up of four concentric layers of cells.
* According to Spruce (Hep. Amaz. et And. 5), this underleaf is truly the modified lobule, a view which is also expressed by the writer in his own work on the North American Frullanice (Trans. Conn. Acad. x, 3). The explanation just given is that of Leitgeb (Untersuch. iiber die Lebermoose, ii, 22), and, being based on embryological studies, is probably more nearly correct. It must be acknowledged, however, that the interposed body (described in the text) some- what obscures Leitgeb’s explanation.
+ Unters. itber die Lebermoose, ii, 25. 1875.
394 A.W. Evans—Hawaiian Hepatice of the Tribe Jubuloidec.
In the genus Jubula, which is intermediate in some respects between Frullania and the Lejeuneex, the plants are distinctly green and are never tinged with red. The leaf-lobe is attached to the axis by a long oblique line of insertion, the lobule is distant from the axis and is therefore not attached to it at all, and the stylus, which is reduced to a single cell, becomes obsolete very early : it appears, therefore, as if no part of the leaf, except the lobe itself, were attached to the axis. The lobes are often dentate or ciliate. The lobe beneath which a branch is situated is more oblique than the others and is attached partly to the axis and partly to the branch. The first underleaf of a branch is usually a simple lanceolate lamina : it is much pushed out of position, being attached partly to the axis and partly to the branch, and its line of insertion does not meet that of the lobe, which is here obviously without a lobule. Innovations, like those of the Lejeuneez, are present ; in typical cases, there are two innovations for each inflorescence, though it is not unusual to find only one of them developed. In Jubula Hutchinsie,* the male branches apparently arise in the same way as those of the following subtribe, being borne behind small but otherwise normal and lobu- late leaves: this peculiarity, however, is not constant for the genus and is not found in the Hawaiian species. The stalk of the capsule is formed of only two concentric layers of cells, as in the Lejeuneee.
1. FRULLANIA Raddi.
In the last published list of Hawaiian Hepatice,t ten species of Frullania are enumerated. An eleventh species, 7’ Oahuensis, first collected by Meyen, is not included here. Although published in 1843, this species was omitted from the Synopsis Hepaticarum, per- haps through an oversight, and has not since appeared in hepatico- logical literature. Several of these eleven species cannot be main- tained. Three of them, / arietina, Fi Kunzei, and / squarrosa, are listed on incorrect determinations, and three of the others, F. explicata, F. oceanica, and F. Helleri, are synonyms. Ang- strom’s 4, Sandvieensis, on the other hand, is a mixed species and was based on two perfectly distinct plants. The six species which I have been able to distinguish fall naturally into three of Spruce’s subgenera and may be identified as follows :
* Cf. Leitgeb, Unters. tiber die Leberm. ii, 37. 1875. } Stephani, Hepaticae sandvicenses. Bull. de l’Herb. Boissier, v, 842. 1897.
#.
A, W. Hvans—Hawaiian Hepatice of the Tribe Jubuloidew. 395
Key to the Species.
Lobule inflated in upper part only, the lower forming a more or less distinct, plane expansion ; perianth with two distinct postical keels; inflorescence autoicous.
FE (Chonanthelia) Aongstroemii.
Lobule inflated throughout, attached to the lobe by a very short, almost transverse keel.
Lobule galeate (about as broad as long); perianth with one or two postical keels ; inflorescence dioicous. Perianth slightly roughened on the keels, otherwise smooth;
leaves not squarrose. F. (Trachycolea) Oahuensis. Perianth distinctly tuberculate ; leaves more or less squar- rose. FE? (Trachycolea) Sandvicensis.
Lobule clavate (longer than broad); perianth with a single posti- cal keel; inflorescence autoicous.
Lobule distant from axis and parallel; lobes sharp-pointed; underleaves contiguous or imbricated; branches not microphyllous. EF. (Diastoloba) apiculata.
Lobule close to axis and parallel; lobes rounded at apex ; underleaves distant or contiguous but not imbricated (except near the end of a stem or branch); branches not microphyllous. EF. (Diastoloba) Meyeniana.
Lobule distant from axis and widely spreading ; lobes blunt or apiculate at apex; underleaves imbricated; some of the branches microphyllous and of short, limited growth.
F. (Diastoloba) hypoleuca.
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I. Subgenus CHoONANTHELIA Spruce. 1. Frullania Aongstroemii sp. nov.
Frullania Sandvicensis Angsty. Ofversigt af Kongl. Vetensk. Akad. Forhand. xxix, Hift 4, 28. 1872 (in part). Plate XLIV., figs. 1-11.
Autoicous: plants growing in wide depressed tufts, greenish, tinged with yellow or brown: stems irregularly pinnately branched : leaves imbricated, not squarrose, the lobe ovate, arching over the stem, but scarcely or not at all cordate at base, decurved at the rounded apex, entire, connected with the lobule by a long fold par- allel with the stem; lobule galeate above, and forming a plane triangular expansion below ; hood extending to the middle of the
396 A.W. Bvans— Hawaiian Hepatice of the Tribe Jubuloidee.
lobule or beyond, compressed at the mouth and inflated in upper gnd outer parts; stylus minute, subulate: underleaves contiguous or subimbricated, small for the subgenus, orbicular, plane or somewhat revolute on sides, scarcely or not at all auriculate at base, bifid about one third with subacute lobes and sinus, margins entire or sinuate- unidentate on the sides: leaf-cells rather thin-walled, but with con- spicuous trigones and occasional intermediate thickenings: @ inflo-_ rescence borne on a principal branch; bracts in about three pairs, unequally bifid, the lobe ovate (or narrowly ovate on the innermost bracts), obtuse to subacute, entire or sinuate on the margins, lobule ovate or ovate-lanceolate, acute or acuminate, subentire but bearing a distinct lobe-like tooth or stylus on the inner edge near the base; bracteole shortly connate with bracts on one or both sides, ovate, bifid about one third with narrow, acute or acuminate lobes and narrow sinus, subentire or unidentate on sides; perianth about half exserted, obovate, gradually narrowed into a short, broad beak, strongly two-keeled postically and with a broad shallow furrow antically: é bracts in two or three pairs, occupying a short subglobose spike below the perianth.
Stems 0.177" in diameter, lobes of leaves 1x0.8™™, lobules 0.5x0.3™™", underleaves 0.4x0.35™™, leaf-cells at edge of lobe 14p, in the middle, 154, and at the base, 20u in diameter, bract I, lobe 1.85x0.85™", lobule 1.85 x 0.75™™, bracteole 1.35 x0.7™", bract II, lobe’ 1:45 x 0.75™™, lobule: *1;25 x 0/6", bracteole;, Tl, lishatseze perianth 2.5 x 1.2™™. :
On rocks and trees. Oahu: Nuuanu (Heller); Luakaha (Cooke).
Frullania Aongstroemii, in most of the specimens at my disposal, grows in company with what I have called / Sandvicensis; and it is quite evident that Angstrém’s original Frullania Sandvicensis, as described by its author, is a mixture of these two species. His sterile type-material, kindly sent me by Professor Nathorst, agrees closely with his description (so far as leaves and underleaves are concerned), and it agrees also with the numerous specimens of the emended /! Sandvicensis, which I have been able to examine. In his description of the perianth, however, he states :—“perianthium obovatum dorso subconcavum canaliculatum spe leviter bicari- natum, ventre bicarinatum,” showing clearly that he did not have the perianth of a Zrachycolea before him, but that of a Chonanthelia. The lower bracteoles, moreover, are described as bipartite, which certainly does not apply to the bracteoles of Sandvicensis, although it is not quite accurate for those of 4! Aongstroemti. The
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oa
Or Ny Te ee ee ee ee
:
A, W. HKvans— Hawaiian Hepatice of the Tribe Subuloidew. 397
two species, although belonging to different subgenera, bear a certain superficial resemblance to each other. In certain cases the resemblance is still more marked from the fact that the lobule of / Aongstroemii is sometimes inflated throughout nearly its whole extent, losing thereby its characteristic Chonanthelia-form, and closely approaching the lobule of / Sandvicensis. These facts, together with the great rarity of the perianths in & Sandvicensis, might easily account for the two having been confused. Even with- out their very characteristic perianths, however, there are good points of distinction between the species. The strongly cordate base of the lobe in F&F: Sandvicensis, the broad, emarginate and cordate underleaves, and the very conspicuous thickenings of the leaf-cells are quite unlike what we tind in 4? Aongstroemii.
In Frullania arietina Tayl., of tropical and subtropical America, F. Aongstroemii finds a close ally. It differs from this species in its autoicous, not paroicous inflorescence, in the entire margins of its bracts, which are less highly connate with the bracteoles, and in the different shape of its lobules. It will be remembered that Austin* has already reported #. arietina from the Hawaiian Islands as growing with his Dendroceros Clintoni. Mixed with the specimens of this last species from Mr. Pearson’s collection, I find a few sterile stems of 7. Sandvicensis but no other /rullania, so that it is prob- able that Austin’s determination was incorrect. Another species of tropical America, /! gibbosa Nees, resembles /? Aongstroemii in its autoicous inflorescence and entire bracts, but it differs in its squar- rose, densely imbricated leaves, with much larger lobule and large disc-like stylus, in its broader underleaves, cordate at the base and less deeply bifid at the apex, and in its more pointed perianths.
II. Subgenus TracnycoLea Spruce. 2. Frullania Oahuensis Hampe.
Frullania Oahuensis Hampe; G. L. & N. Nova Acta Acad. Leop.-Car. xix, suppl. 1, 471. 1843. Plate XLIV., figs. 12-19.
Dioicous: plants closely appressed to substratum, scattered or forming loose thin mats, reddish- or brownish-green, sometimes almost black: stems irregularly pinnately branched: leaves imbri- cated, the lobe ovate, somewhat convex, arching over the stem and slightly cordate at base, rounded at the apex, entire; lobule galeate,
* Bull. Torr. Bot. Club, v, 15. 1874. TRANS. CONN. ACAD., VOL. X. Marcu, 1900
ee 27
398 A.W. Bvans— Hawaiian Hepatice of the Tribe Jubuloidee.
more or less distinctly truncate at base, inflated throughout; stylus minute and slender, three or four cells long: underleaves distant, rhombic-obovate, narrowed and not at all cordate at base, bifid about one half, with acute lobes and sinus, margins angular-unidentate on sides: leaf-cells with somewhat thickened reddish-walls, trigones and intermediate thickenings distinct: @ inflorescence borne on a principal branch; bracts in two or three pairs, increasiag in size toward the perianth, unequally bifid, the lobe broadly ovate, rounded at the apex, entire or slightly sinuate; lobule lanceolate or ovate- lanceolate, acute, or obtuse and apiculate, bearing a lobe-like tooth or stylus at the middle of the inner edge on the innermost bracts, close to the base on the others, otherwise entire ; bracteole slightly connate on one side or free, ovate, bifid about one half with acute lobes and sinus, entire or sparingly laciniate-toothed on the sides; perianth more than half exserted, obovate, truncate above and abruptly narrowed into a short beak, with a broad, usually two- angled keel postically and commonly with one to three low supple- mentary keels on both surfaces, roughened or very sparingly tubercu- late, at least on the keels: ¢ bracts in two to ten pairs, occupying a short branch and forming a globose or oblong spike.
Stems 0.09™" in diameter, lobes of leaves 0.4x0.3™", lobules 0.14 x 0.12™", underleaves 0.14 x 0.15™™, leaf-cells at edge of lobe 12p, in the middle 15, and at the base 18 in diameter, bract I, lobe 0.7x0.5"™, lobule 0.6x 0.17", bracteole I, 0.5 x0.25™™, bract II, lobe 0.5x0.4™", lobule 0.4x0.15™%, bracteole II, 0.35 x0.17"™, perianth 1.1 x 0.75™™.
On trees. Oahu: Nuuanu (Cooke); first collected on the island by Meyen. Kauai: Kipu, Lihue, Half Way Bridge (Cooke).
Frullania Oahuensis is the smallest known Hawaiian Frullania, being even smaller than /) Meyeniana, with which it often grows and which it somewhat resembles. The regularly pinnate habit of this latter species and its clavate instead of galeate lobule will at once serve to distinguish it. The North American F! Virginica Gottsche is a much closer ally of #: Oahuensis, but is a somewhat larger plant in all its parts, and its leaves are more strongly cordate at the base.
The type-specimens of / Oahwensis are apparently not to be found in the Gottsche Herbarium at Berlin. There is, however, a drawing so labeled among the beautiful “Icones Hepaticarum Ineditz,” and this agrees so closely with the specimens collected by Mr. Cooke that I have no hesitancy in pronouncing them the same.
A. W. Evans—Hawaiian Hepatice of the Tribe Jubuloidew. 399
3. Frullania Sandvicensis Angstr. emend.
Frullania Sandvicensis Angsty. Ofversigt af Kong]. Vetensk. Akad. Forhand. xxix, Haft 4, 28. 1872 (in part). Frullania squarrosa Auct. (not F. squarrosa (R. Bl. & N.) Dum.). Frullania arietina Aust. Bull. Torr. Bot. Club, 5:15. 1874 (not Tayl.). Plate XLV., figs. 1-7.
Dioicous: growing in wide depressed tufts, often mixed with mosses or other hepatics, brownish-green, sometimes tinged with reddish: stems irregularly pinnately branched: leaves densely imbri- cated, more or less squarrose when moist, the lobe broadly ovate, arching over the stem and cordate, both at the antical base and at the keel, rounded at the apex, entire, slightly revolute on postical margin; lobule galeate throughout the whole or the greater part of its extent, hood inflated in upper and outer parts, compressed below; stylus small, subulate: underleaves imbricated, broadly orbicular or reniform, bifid about one fifth with broad obtuse or apiculate lobes and lunulate sinus, more or less cordate and channeled at base, margins entire or nearly so: leaf-cells rather thick-walled with very conspicuous trigones and intermediate thickenings: Q inflorescence borne on a short simple branch; bracts in two or three pairs, un- equally bifid, the lobe ovate or obovate, obtuse or rounded at the apex, entire ; lobule ovate, acute or acuminate, sparingly and irreg- ularly toothed on the inner edge, one of the teeth (the stylus) being more distinct and larger than the others ; bracteole connate on one side, ovate, bifid one fourth to one third with acute or acuminate teeth and narrow sinus, sparingly and coarsely toothed on the mar- gins; perianth oblong-obovate, truncate above and abruptly narrowed into a short beak, strongly one-keeled postically and bearing on the surface numerous papilla-like or lobe-like projections, especially on the two lateral keels: ¢ bracts in about six pairs, occupying a short branch and forming a short, oval spike.
Stems 0:2™" in diameter, lobes of leaves 1x0.9™™, lobules 0.32 x 0.28™™", underleaves 0.65 x 0.85™™, leaf-cells at edge of leaf 16m in diameter, in the middle 28x 19», at the base 30m in diameter, bract I, lobe 1.5x0.9"", lobule 1.3.x 0.6™", bracteole I, 0.95 x 0.7™™, bract II, lobe 1.5x1™", lobule 1x0.6™", bracteole H, 0.9 x0.75™™, perianth 2.3 x 1.35™™,
On rocks and trees. Oahu: Lulihi (Wawra); Nuuanu (Heller, Cooke); foot of Konahuanui (Cooke); Luakaha (Cooke); first col- lected on the island by Andersson. Kauai; Hanalei, Kilohana,
400 A.W. Bvans—Hawaiian Hepatice of the Tribe Jubuloider.
Lihue, Half Way Bridge (Cooke). Hawaiian Islands (Hillebrand, Mann and Brigham).
The determination of this species is based on the sterile type- material, preserved in the Royal Academy of Science at Stockholm, ° It is apparently the plant which has been referred by Austin and other authors to the widely distributed and variable J. squarrosa, and it is somewhat questionable as to whether the two are really distinct. When well developed, / Sandvicensis is a little more robust than F. squarrosa, its leaves are less strongly squarrose, and its leaf-cells have somewhat better developed trigones. The underleaves, how- ever, offer the best point of distinction: these are much broader than in & sguarrosa, often completely concealing the lobules, they are less deeply bifid, with broad lobes and sinus, and their margins are usually entire. These differences, although slight, are appar- ently constant. The distinctive characters between