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SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY • NUMBER 435

Revision of the Atlantic Brisingida (Echinodermata: Asteroidea), with Description of a New Genus and Family

Maureen E. Downey

SMITHSONIAN INSTITUTION PRESS

City of Washington

1986

ABSTRACT

Downey, Maureen E. Revision of the Atlantic Brisingida (Echinodermata: Asteroidea), with Description of a New Genus and Family. Smithsonian Contributions to Zoology, number 435, 57 pages, 24 figures, 1986.—The Brisingida, a mainly deep-sea order of Asteroidea, is represented in the Atlantic by 23 species belonging to 9 genera and 2 families. The following changes in classification are suggested:

Freyellidae, new family, with Atlantic genera Colpaster, Freyastera, and Freyella, and non-Atlantic genera Belgicella and Freyellaster.

Freyastera, new genus, with the species benthophila, mexicana, sexradiata, and tuberculata, all formerly assigned to the genus Freyella Perrier, 1885c.

Craterobrisinga Fisher, 1916b, synonymized with Brisinga Asbjørnsen, 1856.

Freyella edwardsi (Perrier, 1882) transferred to the genus Colpaster Sladen, 1889.

Astrocles Fisher, 1917 (non-Atlantic), synonymized with Freyella.

Freyella spinosa, bracteata, aspera, abyssicola, and laubieri synonymized with Freyella elegans (Verrill, 1884).

Freyella trispinosa H.L. Clark, 1941, synonymized with Freyella microspina Verrill, 1894.

Brisinga americana Verrill, 1880, transferred to the genus Novodinia Dartnall, Pawson, Pope, and Smith, 1969.

Novodinia elegans (Perrier, 1885a), an invalid homonym of Brisinga elegans (Perrier, 1885c), is renamed Novodinia homonyma, new name.

Novodinia robusta (Perrier, 1885a), synonymized with Novodinia semicoronata (Perrier, 1885a).

OFFICIAL PUBLICATION DATE is handstamped in a limited number of initial copies and is recorded in the Institution's annual report, Smithsonian Year. SERIES COVER DESIGN: The coral Montastrea cavernosa (Linnaeus).

Library of Congress Cataloging in Publication Data

Downey, Maureen E.

Revision of the Atlantic Brisingida (Echinodermata: Asteroidea), with description of a new genus and family

(Smithsonian contributions to zoology; no. 435)

Bibliography: p.

Includes index.

1. Brisingida—Atlantic Ocean—Classification. 2. Echinodermata—Classification. 3. Echinodermata—Atlantic Ocean—Classification. I. Title. II. Series.

QL1.S54 no. 435 [QL384.A8] 591s [593.9′3] 86–6579

Revision of the Atlantic Brisingida (Echinodermata: Asteroidea), with Description of a New Genus and Family

Maureen E. Downey

____________

Maureen E. Downey, Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Introduction

The Brisingida are almost exclusively abyssal, with some Antarctic species occurring more in shallow waters. Hitherto, all brisingidans were considered to belong to one family, the Brisingidae, but careful comparison of all genera clearly shows two well-defined families: the Brisingidae, containing the genera Astrostephane Fisher (1917), Brisinga Asbjørnsen (1856), Brisingaster de Loriol (1883), Brisingella Fisher (1917), Brisingenes Fisher (1917), Midgardia Downey (1972), Novodinia Dartnall et al. (1969), Parabrisinga Hayashi (1943), and Stegnobrisinga Fisher (1916b); and the Freyellidae, with the genera Belgicella Ludwig (1903), Colpaster Sladen (1889), Freyella Perrier (1885c), Freyellaster Fisher (1918), and Freyastera, new genus. These families can be separated both morphologically and ecologically, as discussed below under the family headings.

The Brisingida are related most closely to the Forcipulatida, but are obviously distinctive because of their small, circular disc and long, attenuate arms. Their general appearance is superficially like that of a large, multiarmed ophiuroid. The fused ring of disc plates, spool-like ambulacral column, reduced abactinal plates, single series of marginals, lack of actinal plates, and crossed pedicellariae only are characteristic of the order.

HISTORICAL BACKGROUND.—The first known representative of this order, Brisinga endecacnemos, was described by Asbjørnsen in 1856 from Hardangerfjord, Norway. A second species, Brisinga coronata (= Brisingella coronata) was described by G.O. Sars in 1871 from Lofoten, Norway, and in 1875 Sars recognized the uniqueness of these starfish by erecting a new family, Brisingidae, to accommodate them. Increased deep-sea dredging activity led to further additions to the family by Sladen, Perrier, Verrill, Fisher, and others. Perrier (1885c) based a second genus, Freyella, largely on the presence of bare interradial plates and a consequent gradually curved interradial arc (as opposed to the acute arc in Brisinga). The only major taxonomic and biogeographic survey of the group was that of Fisher (1917, 1919). In 1928 he raised the family to a suborder, Brisingina. In 1958, Tortonese separated the family Brisingidae from the order Forcipulatida and placed it in a new order, Euclasteroidea. I agree fully with this placement, and the content of the order is unchanged. The consensus among current asteroid specialists, however, seems to be that the name Euclasteroidea is of awkward construction and conveys no indication of the content of the order. Accordingly, I have dropped the name Euclasteroidea in favor of the simpler name Brisingida, which, being based on the family name established by G.O. Sars in 1875, is familiar and instantly defines the content of the order.

RANGE EXTENSIONS.—The following range extensions are noted:

Brisinga costata Verrill, 1884, hitherto known only from north of Cape Hatteras in the Western Atlantic, has now been identified from the Gulf of Mexico, Haiti, and Venezuela. The bathymetric distribution is extended to 2377 m.

Brisinga cricophora Sladen, 1889, known only from off the Virgin Islands, has now been identified from Florida, the Bahamas, the Sargasso Sea, and western South Africa. The bathymetric range is extended to 1340 m.

Brisinga hirsuta Perrier, 1894, from the Iberian Basin, has also been collected in the Gulf of Guinea.

Brisingella verticellata (Sladen, 1889), known only from north of Cape Hatteras, has now been collected in the Gulf of Mexico.

Brisingenes multicostata (Verrill, 1894), known only from off New England in depths of over 2000 m, has been identified from the Straits of Florida in 805 m.

Novodinia americana (Verrill, 1880), from Banquereau Bank, has been collected from off Colombia.

Novodinia antillensis (A.H. Clark, 1934), from off Puerto Rico, has been collected in the Gulf of Mexico.

Novodinia pandina (Sladen, 1889), heretofore known only from Faeröe Channel, has been identified from off North Carolina.

Stegnobrisinga splendens H.L. Clark, 1926, from South Africa (∼1000 m), has been identified from the Straits of Florida and from off Venezuela (to 4000 m).

Colpaster scutigerula Sladen, 1889, known only from the type collected by the Challenger southwest of the Canary Islands in 2789 m, has now been collected from the Gulf of Guinea (2525 m) and from off Honduras (933 m).

Freyastera benthophila (Sladen, 1889), previously known from the South Pacific, the Bay of Bengal, and the Bay of Biscay, is now reported from the Eastern Pacific off California.

Freyella elegans (Verrill, 1884), known from Cape Hatteras north to George's Bank, has been collected from the Gulf of Guinea and from off Greenland.

Freyella microspina Verrill, 1894, from off Nantucket and (as Freyella trispinosa) south of Cuba, has also been identified from off Surinam. The bathymetric range is extended to 2734 m.

BIOLOGY.—Little is known about feeding among the Brisingida, but small crustacean exoskeletons are sometimes found within the actinostome. Recent bottom photographs by Pawson and Miller (pers. comm.) and others (e.g., Pawson, 1976) show brisingids with their arms upraised, apparently suspension feeding (Figure 1a,b), whereas the Freyellidae are always photographed lying flat on the bottom (Figure 1d) or with only the distal portion of the arms raised (Figure 1c) and are probably not suspension feeders (see “Discussion” under Brisingidae and Freyellidae).

DIAGNOSTIC METHOD.—In this paper I have used a different approach to the taxon diagnosis, using a master list of characters and character states compiled from the major literature on the Brisingida (“Appendix”). This compilation represents a first step in the development of a comprehensive character list for the entire Asteroidea. The numbers (for characters) and letters (for character states) in square brackets in the diagnoses are described in the appendix. The advantages of having a number and letter system are: (1) it becomes immediately apparent which characters belong at which taxonomic level; (2) characters and character states can be compared in numerous ways to investigate possible relationships; and (3) some evolutionary trends can be detected; less specialized characters are shared at higher taxonomic levels, whereas the greatest degree of specialization appears at the species level or below.

Characters are considered in the following order: general form, disc, arms, terminal plates, inferomarginal plates, adambulacrals, ambulacrals, mouth plates, gonads, pedicellariae. The numbering system for characters is by tens to allow for addition of further characters. Some features are repeated in other forms for ease of evaluation (for example, disc shape and interradial arc frequently offer the same information) and thus tend to weight certain features. Characters were taken en masse from existing literature and in that way reflect a minimal personal bias.

FIGURE 1.—Bottom photographs of various brisingidans showing normal position: a, Novodinia sp.; b, Brisinga sp.; c, Freyastera elegans; d, Freyastera sp. (normally with only six arms).

The Brisingida was selected to test this method primarily because the order was in need of revision and secondarily because it is a relatively small order, with distinctive characters. Revision of another order (or family or genus) would require analyses of a different suite of characters and character states.

Note that the scale on all figures represents one centimeter.

ACKNOWLEDGMENTS.—I am greatly indebted to Stephen Cairns, David L. Pawson, and Frederick M. Bayer, all of the Department of Invertebrate Zoology, National Museum of Natural History, Smithsonian Institution, and to Daniel B. Blake, Department of Geology, University of Illinois, for reading rough drafts of this manuscript and offering valuable comments and suggestions and especially for their encouragement and kindness in urging me to publish results of this somewhat novel approach. Special thanks are due to Molly Griffin, Derwood, Maryland, for the drawings of Brisingella coronata and to Michael Carpenter, National Museum of Natural History, for taking the photographs. I thank David L. Pawson, National Museum of Natural History, and Barbara Hecker of the Lamont Oceanographic Laboratory for the bottom photographs in Figure 1, and John Miller, Harbor Branch Foundation, Fort Pierce, Florida, for coments on possible feeding behavior.

Order BRISINGIDA Fisher, 1928, new status

BRISINGINA Fisher, 1928:4 [suborder].

EUCLASTEROIDEA Tortonese, 1958:1–3.

DIAGNOSIS.—[10.a] Form ophiuroid-like; [20.a] disc size small (∼10–40/1 R/r) [40.a] disc shape circular, frame an inflexible ring of plates fused or united by syzygy; [90.a] position of madreporite more or less lateral, on outer edge of disc (except Freyastera mexicana); [170] arms always more than 5; [180.a] shape of arms long, attenuate, R/r more than 6/1; [200.a] gonadal region of arms differing from rest of arm; [230.a] size of abactinal arm plates relative to marginals small; [300.a,b] marginals not corresponding one to one with adambulacrals (except Brisinga hirsuta); [310.a] marginals armed with spines; [330.a] adambulacral plates more or less cylindrical; [410.a] ambulacral groove broad, open; [440.a] ambulacral plates not crowded (except Novodinia americana), heads abutting end to end; [450] tubefeet in 2 rows; [460.a] tubefeet suckered; [610.a] pedicellariae crossed.

Family BRISINGIDAE G.O. Sars, 1875

BRISINGIDAE G.O. Sars, 1875:1.—Verrill, 1880:139.—Perrier, 1885c:3.—Sladen, 1889:603.—Perrier, 1891c:198.—Verrill, 1895:199.—Ludwig, 1897b:418; 1900:488.—Fisher, 1906:1108.—Koehler, 1907:3; 1906 [1907]:141; 1908:578.—Fisher, 1916b:31; 1917:419; 1918:104; 1919:516,—H.L. Clark, 1920:107.—Fisher, 1928:7.—Hayashi, 1943:136.—Tortonese, 1958:1.—A.M. Clark, 1962:68.—Bernasconi, 1964:248.—Spencer and Wright, 1966:U76.—McKnight, 1975:17.—A.M. Clark and Courtman-Stock, 1976:97.

DIAGNOSIS.—[50.a] Interradial arcs acute; [80.a,b] madreporite large to moderate; [140.b] no bare interradial plates on disc; [190.a] arms deciduous, constricted where they join the disc; [220.a] abactinal arm plates forming costae; [400.a] proximal adambulacrals united interradially.

DISCUSSION.—The Brisingidae, with arms sharply demarked from the disc by a constriction at their base, and with abactinal arm plates in costae across the arms with either bare membrane or thin imbricate fenestrate plates between the costae, raised arms above the disc are admirably adapted to suspension feeding. Only the species of Novodinia are slightly limited in their ability to raise the arms completely over the disc because of the plating between the costae on the proximal part of the arms, but even they are less limited in this regard than are the Freyellidae. The Brisingidae live on hard substrates, and in bottom photographs are easy to mistake for comatulid crinoids because of their feeding position.

Genus Brisinga Asbjørnsen, 1856

Brisinga Asbjørnsen, 1856:95.—G.O. Sars, 1875:1.—Sladen, 1889:604.—Wood-Mason and Alcock, 1891:12.—Alcock, 1893b:170.—Bell, 1893:104.—Perrier, 1894:61.—Verrill, 1894:280; 1895:211.—Fisher, 1916b:31; 1917:427; 1918:103; 1919:516.—Döderlein, 1927:292.—Fisher, 1928:7.—A.M. Clark and Courtman-Stock, 1976:97.

DIAGNOSIS.—[30.a] Disc thick, raised well above plane of arms; [50.a] interradial arcs acute; [60.a] skin thick, dense, opaque; [70.a,b] anal opening barely visible (microscopic) or absent; [140.b] no bare interradial plates; [160.a] no papulae; [170] number of arms 7–13; [190.a] arms deciduous, constricted at base; [220.a] abactinal arm plates imbricating across arms in costae; [400.a] proximal adambulacral plates united interradially; [590.a] gonads numerous; [600.a] gonads serial.

TYPE-SPECIES.—Brisinga endecacnemos Asbjørnsen, 1856, by original designation.

DISCUSSION.—Brisinga formerly contained many species that have since been assigned to other genera within the family, mainly by Fisher (1917, 1919). The type species was monographed and illustrated in detail by G.O. Sars (1875), to whose thorough explication we owe much of our knowledge of this deep-sea genus.

The characters Fisher (1917, 1919) used to separate Craterobrisinga from Brisinga are not reliable. Fisher distinguished Craterobrisinga on the basis of capitate or truncate subambulacral spines in the proximal region of the arms, presence of a second subambulacral spine, and adambulacral plates higher than long. Brisinga costata has 1–3 capitate spines on the proximal adambulacrals, and the adambulacrals are longer than high. Brisinga cricophora (which Fisher assigned to Craterobrisinga) has adambulacrals higher than long proximally, longer than high distally, and only one subambulacral spine. Brisinga endecacnemos has adambulacrals higher than long, and one acicular subambulacral spine. Brisinga hirsuta has adambulacrals longer than high, and one large and one small acicular subambulacral spine. Thus characters Fisher used to distinguish Craterobrisinga are specific, not generic.

The species of Brisinga have large eggs and direct development. Post-larval development is discussed by G.O. Sars (1875).

CHARACTERS.—Those peculiar to the genus Brisinga: None.

Characters shared with other genera of Brisingidae:

| Brisingella: 60.a, 70.a,b, 160.a.
| Brisingenes: 70.a,b, 600.a.
| Midgardia: 30.a, 60.a.
| Novodinia: 30.a, 60.a.
| Stegnobrisinga: 30.a, 70.a,b, 160.a.

Characters in which Brisinga differs from other genera of Brisingidae:

| Brisingella: 30, 590, 600.
| Brisingenes: 30, 60, 160.
| Midgardia: 70, 160, 590, 600.
| Novodinia: 70, 160, 590, 600.
| Stegnobrisinga: 60, 590, 600.

Brisinga costata Verrill, 1884

FIGURE 2

Brisinga costata Verrill, 1884:382; 1885:529; 1894:280; 1895:211.

MERISTICS.—Arms = 12–13, R = 161–381 mm, r = 16–22 mm, R/r = 10–17/1, length of gonadal region = 61–95 mm, length of longest arm spine = 9–15 mm.

DIAGNOSIS.—[80.a] Madreporite large, raised; [100.a] madreporite irregular, coarsely channeled; [110.a,b] abactinal disc plates metapaxillar, tumid, irregular; [120.a] abactinal disc plates widely scattered in membrane; [130.a] abactinal disc plates bearing 1–3 very small sharp spinelets; [170] number of arms 12 or 13; [210.a] abactinal arm plates rod-like; [220.a] abactinal arm plates imbricate across arms in raised ridges (costae), costae well spaced, no plates between or beyond costae; [240.a] abactinal arm plates bearing 0–8 small sharp spinelets in a line on ridge of plate; [280.a] marginal plates very small relative to adambulacrals; [290.a] shape of marginals irregular; [300.b] marginals occurring about every 2–3 adambulacrals; [310.a] marginals bearing one moderately long acicular spine; [320.a] adambulacral plates large relative to marginals, higher than long; [330.b] adambulacrals spool-shaped, higher than long, with sharp inner distal prolongation; [340.a] furrow margin straight; [350] number of furrow spines 1–3; [360.a] furrow spines quite small, slender, acute; [370.a] furrow spines on proximal end of adambulacral; [380] number of subambulacral spines 1(2–3); [390.a,b] subambulacral spines moderately large, acicular, sometimes proximal ones slightly capitate; [420.a] ambulacral plates high, forming distinct ridge along arm; [430.a] head of ambulacrals saddle-shaped; [470.a] structure of tubefeet delicate, thin, translucent; [480.a] mouth plates rather small relative to adambulacrals; [490.a] mouth plates T-shaped; [500] number of preoral spines per mouth plate 1; [510.a] preoral spines very long, fine, acicular; [520.a] position of preoral spines adoral; [530] number of lateral oral spines 2 per mouth plate; [540.a] lateral oral spines small, slender, slightly curved, acicular; [550.a] lateral oral spines on oral edge of mouth plates, beneath preoral spines; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines large, acicular; [580.a] position of suboral spines just behind preorals; [620.a,b,c] pedicellariae tiny, abundant on membrane of disc and in irregular bands on membranous part of arms beyond costae, all spines enclosed in fleshy sacs completely covered with pedicellariae; [630.a] all pedicellariae microscopic.

FIGURE 2.—Brisinga costata: a, abactinal view; b, actinal view.

COLOR.—Unknown.

TYPES.—USNM 7820 (holotype), Albatross Sta 2210, collected 1884 south of Block Island, Rhode Island, 1812 m; USNM 11256 (paratype), Albatross Sta 2533, SE of George's Bank, 1514 m.

MATERIAL EXAMINED.—Holotype and paratype; 1 specimen, USNM 6739, Albatross Sta 2116, off Hatteras, North Carolina, 1624 m; arm fragments, Albatross Sta 2706, off George's Bank, 2173 m; 1 specimen, USNM 18503, Albatross Sta 2116, off Cape Hatteras, North Carolina, 1624 m; arm fragment, R/V Knorr Sta 325, Hudson Canyon, 39°13.3′N, 71°53.4′W, 1974 m; arm fragment, USNM E20942, Oregon Sta 2571, Gulf of Mexico, 26°34′N, 90°31′W, 2377 m; 2 specimens, Pillsbury Sta 1178, Haiti, 19°14′N, 73°14′W, 1903 m; 17 specimens, Pillsbury Sta 748, off Venezuela, 11°24.8′N, 67°10.1′W, 1865 m.

DISTRIBUTION.—Western Atlantic, George's Bank to Venezuela, 1514–2377 m.

DISCUSSION.—Within the gonadal inflation, numerous oval sacs contain one to four very large yolky eggs (up to 3 mm) and a number of smaller eggs. Development is therefore likely direct and probably demersal.

CHARACTERS.—Those peculiar to B. costata: 430.a.

Characters shared with other species of Brisinga:

B. |   cricophora: 80.a, 100.b, 110.b, 130.a, 210.a, 220.a, 240.a, 280.a, 290.a, 300.b, 310.a, 320.a, 330.b, 360.a, 370.a, 390.a,b, 470.a, 480.a, 500.1, 510.a, 540.a, 570.a, 580.a, 620.a,b,c, 630.a.
B. |   endecacnemos: 100.a, 110.a, 130.a, 210.a, 220.a, 240.a, 280.a, 300.b, 310.a, 320.a, 330.b, 360.a, 370.a, 390.a, 420.a, 480.a, 500.1, 510.a, 520.a, 540.a, 560.1, 570.a, 580.a, 620.a,b, 630.a.
B. |   hirsuta: 110.b, 120.a, 210.a, 220.a, 240.a, 280.a, 310.a, 330.b, 360.a, 390.a, 420.a, 470.a, 500.1, 510.a, 570.a, 620.a,b,c.

Characters in which B. costata differs from other species of Brisinga:

B. |   cricophora: 100, 110, 120, 320, 340, 370, 420, 430, 490, 520, 550.
B. |   endecacnemos: 80, 100, 110, 120, 290, 310, 340, 370, 390, 430, 470, 490, 530, 550, 620.
B. |   hirsuta: 80, 100, 110, 130, 290, 300, 320, 340, 370, 430, 480, 490, 520, 530, 550, 580, 630.

Brisinga cricophora Sladen, 1889

FIGURE 3

Brisinga cricophora Sladen, 1889:606–608, pl. 59: figs 6–8.

Craterobrisinga cricophora.—Fisher, 1917:426; 1919:513.

MERISTICS.—Arms = 11, R = 130 mm (estimated), r = 7–13 mm, R/r = 10–19/1, length of gonadal region = 60 mm, length of longest arm spine = 17 mm.

DIAGNOSIS.—[80.a] Madreporite large, raised; [100.b] madreporite irregular; [110.b] abactinal disc plates tumid; [120.b] abactinal disc plates dense in membrane; [130.a] abactinal disc plates sometimes bearing one or more tiny hyaline spinelets or one moderately long hyaline spinelet; [170] number of arms 11; [205.a] gonadal region of arms 60 mm, moderately inflated; [210.a] abactinal arm plates elongate, rod-like; [220.a] abactinal arm plates imbricate, costae delicate, frequently incomplete, bare membrane between and beyond costae; [240.a] abactinal arm plates bearing 1 or 2 tiny thorn-like spinelets; [280.a] marginal plates very small relative to adambulacrals; [290.a] shape of marginals irregular; [300.b] marginals occurring about every other adambulacral; [310.a] first 5–7 marginals without spines, rest with 1 or 2 moderately long acicular spines; [320.a,b] adambulacral plates higher than long proximally, longer than high distally; [330.b] adambulacrals spool-shaped; [340.b] furrow margin slightly indented; [350] number of furrow spines 2; [360.a] furrow spines small, tapering; [370.a,b] one furrow spine at either end of adambulacral; [380] number of subambulacral spines 1; [390.a,b] subambulacral spines extremely long, length increasing distad, first 10–15 stout, capitate, rest slender, aciculate; [420.b] ambulacral plates rather small, delicate; [430.b] head of ambulacral plates quite small, hourglass-shaped; [470.a] tubefeet delicate, slender, translucent; [480.a] mouth plates very small relative to adambulacrals; [490.b] mouth plates trapezoidal, with outer side and distal end concave; [500] number of preoral spines per mouth plate 1; [510.a] preoral spines small, acicular; [520.b] preoral spines directed laterally, across groove; [530] number of lateral oral spines 1–3; [540.a] lateral oral spines small, acicular; [550.b] lateral oral spines on face of mouth plates, near outer side; [560] number of suboral spines per mouth plate 1 or 2; [570.a] suboral spines moderately large, acicular; [580.a] suboral spines behind preoral spines; [620.a,b,c] pedicellariae abundant, all spines enclosed in fleshy sac completely covered with pedicellariae, membrane between and beyond costae with transverse bands of tiny pedicellariae; [630.a] all pedicellariae microscopic.

FIGURE 3.—Brisinga cricophora: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

COLOR.—Orange.

TYPE.—BM(NH) 90.5.7.1043 (holotype), Challenger Sta 24, NW of St. Thomas, Virgin Islands, 18°38.3′N, 65°5.3′W, 713 m.

MATERIAL EXAMINED.—Holotype; USNM E20952, 15 specimens, Gerda Sta 190, Bahamas, 25°57′N, 78°07′W, 733–897 m; USNM E20947, arm fragments, Gerda Sta 403, off Florida, 27°49′N, 78°50′W, 824 m; USNM E20946, arm fragments, Gerda Sta 672, off Florida, 27°53′N, 79°03′W, 796 m.

DISTRIBUTION.—Florida, the Bahamas, the Virgin Islands, Sargasso Sea, western South Africa; 713–1340 m.

CHARACTERS.—Those peculiar to B. cricophora: None.

Characters shared with other species of Brisinga:

B. |   endecacnemos: 120.b, 130.a, 170.11, 205.a, 210.a, 220.a, 240.a, 280.a, 300.b, 310.a, 330.b, 340.b, 360.a, 380.1, 480.a, 500.1, 510.a, 540.a, 550.b, 570.a, 580.a, 630.a.
B. |   hirsuta: 110.b, 210.a, 220.a, 240.a, 280.a, 310.a, 330.b, 340.b, 360.a, 470.a, 500.1, 510.a, 520.b, 560.1–2, 570.a, 620.a,b,c.

Characters in which B. cricophora differs from other species of Brisinga:

B. |   endecacnemos: 80, 100, 110, 290, 320, 390, 420, 430, 470, 490, 520, 620.
B. |   hirsuta: 80, 100, 120, 130, 170, 205, 290, 300, 380, 420, 480, 490, 550, 580, 630.

Brisinga endecacnemos Asbjørnsen, 1856

FIGURE 4

Brisinga endecacnemos Asbjørnsen, 1856:95, pl. 9.—G.O. Sars, 1875:1–112, pl. 4: fig. 41, pl. 7.—Bell, 1892 [1893]:520; 1893:104.—Perrier, 1894:62.—Farran, 1913:27–28.—Gage et al., 1983:284–285.

MERISTICS.—Arms = 11, R = 342 mm, r = 12 mm, R/r = 28.5/1, length of gonadal region = 130 mm, length of longest arm spine = 18 mm.

DIAGNOSIS.—[80.b] Madreporite of moderate size, raised; [100.a] madreporite coarsely channeled; [110.a] abactinal disc plates metapaxillar; [120.b] abactinal disc plates dense, embedded in membrane; [130.a] abactinal disc plates armed with a tiny echinulate spinelet; [170] number of arms 11; [205.a] gonadal region nearly half arm length, moderately inflated; [210.a] abactinal arm plates rod-like; [220.a] abactinal arm plates imbricate, costae meandering, some incomplete, some anastomosing, not widely spaced, membrane between and beyond costae with minute spinelets; [240.a] abactinal arm plates bearing row of fine spinelets; [250.a] terminal plates rather small relative to marginals; [260.a] terminals catsclaw-shaped; [270.a] terminals bearing about 5 slender spines; [280.a] marginal plates very small relative to adambulacrals; [290.b] marginals irregularly oval; [300.b] marginals occurring ring about every other adambulacral; [310.a] marginals bearing each one very long acicular spine; [320.a] adambulacral plates higher than long; [330.b,c] adambulacrals nearly discoid, spool-shaped; [340.b] furrow margin slightly indented; [350] number of furrow spines 1 or 2; [360.a] furrow spines small, acicular; [370.a(a,b)] furrow spines at proximal end of adambulacral if single, one at each end if double; [380] number of subambulacral spines 1; [390.a] subambulacral spines long, acicular; [420.a] ambulacral plates fairly high, heavy; [430.c] head of ambulacrals Y-shaped, larger than base, cylindrical, with inner proximal projection; [470.b] tubefeet rather heavy; [480.a] mouth plates rather small relative to adambulacrals; [490.c] mouth plates longer than broad, subtriangular, truncate; [500] number of preoral spines per mouth plate 1; [510.a] preoral spines acicular, moderately long; [520.a] preoral spines projecting into actinostome; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines small, acicular; [550.b] lateral oral spines at sides of mouth plates; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines moderately long, acicular; [580.a] suboral spines below preoral spines; [620.a,b(f)] pedicellariae on both abactinal and actinal surfaces, on disc confined to mouth spines, on arms mostly confined to adambulacral spines, very few on membranous areas of arms, in bands; [630.a] all pedicellariae microscopic.

FIGURE 4.—Brisinga endecacnemos: a, abactinal view (from Koehler, 1909c); b, Details (from G.O.Sars, 1875).

COLOR.—Bright yellow-orange.

TYPE.—Bergens Museum; Hardangerfjord, Norway, 183–366 m.

MATERIAL EXAMINED.—2 lots, 3 specimens in BM(NH).

DISTRIBUTION.—Norway to Portugal; 183–2000 m.

CHARACTERS.—Those peculiar to B. endecacnemos: None.

Characters shared with other species of Brisinga:

B.
  hirsuta: 80.b, 210.a, 220.a, 240.a, 280.a, 330.b, 340.b, 360.a, 390.a, 420.a, 500.1, 510.a, 530.1, 570.a.

Characters in which B. endecacnemos differs from other species of Brisinga:

B.
  hirsuta: 100, 110, 120, 130, 170, 205, 290, 300, 320, 380, 430, 470, 480, 490, 520, 550, 580, 620, 630.

Brisinga hirsuta Perrier, 1894

FIGURE 5

Brisinga hirsuta Perrier, 1894:66.

MERISTICS.—Arms = 13, R = 148 mm (estimated), r = 8.5 mm, R/r = 17.4/1, length of gonadal region = 45 mm, length of longest arm spine = 5 mm.

DIAGNOSIS.—[80.b] Madreporite of moderate size; [100.c] madreporite subtubercular; [110.b] abactinal disc plates tumid, close but not crowded; [120.a] abactinal disc plates scattered in membrane; [130.b] abactinal disc plates each bearing a moderately long setose spine; [170] number of arms 13; [205.b] gonadal region of arms 45 mm long, not inflated; [210.a] abactinal arm plates rod-like; [220.a] abactinal arm plates imbricate, in costae rather closely and evenly spaced, membrane between and beyond costae usually bare but sometimes with isolated minute platelets; [240.a] costal plates bearing transverse row of small, acute spinelets; [280.a] marginal plates small relative to adambulacrals; [290.c] marginals tubercular; [300.a] marginals corresponding to every adambulacral; [310.a] every other marginal bearing a long, ridged, hyaline spine; [320.b] adambulacrals longer than high; [330.b] adambulacrals spool-shaped, with definite ridge at each end; [340.b] furrow margin slightly indented; [350] number of furrow spines 2 proximally (in gonadal region), 1 distally; [360.a] distal furrow spine small, proximal one very fine, setose; [370.b(b,a)] one furrow spine at distal end of adambulacral and, in gonadal region, one at proximal end also; [380] number of subambulacral spines 2; [390.a] subambulacral spines quite long, acicular, ridged, hyaline, 1 on proximal end of actinal face of adambulacral, one in center of actinal face; [420.a] ambulacral plates large, heavy; [430.b,d] ambulacral plates T-shaped, head hourglass-shaped, with definite ridge at each end; [470.a] tubefeet quite delicate; [480.b] mouth plates of moderate size; [490.d] mouth plates cylindrical, with outer side and distal end indented, broader at base than at apex, somewhat flared at each end; [500] number of preoral spines per mouth plate 1; [510.a] preoral spines small, delicate, acicular; [520.b] preoral spines laterally directed; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines small, delicate, acicular; [550.c] lateral oral spines placed distad on mouth plates; [560] number of suboral spines per mouth plate 1 or 2; [570.a] suboral spines very long, hyaline, acicular; [580.b] suboral spines distad to preorals; [620.a,b,c] pedicellariae on both surfaces of disc and arms, in bands on membranous part of arms; [630.a,b] pedicellariae on costae, mouth plate spines and furrow spine huge, smaller, abundant on other spines.

FIGURE 5.—Brisinga hirsuta: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

COLOR.—Unknown.

TYPE.—Paris Museum 4594 (holotype consisting of one arm fragment); Travilleur Sta 13, between Portugal and the Azores, 2030 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, Pillsbury Sta 266, Gulf of Guinea, 1 ° 13′N, 7°46′E, 2525 m.

DISTRIBUTION.—Iberian Basin, Gulf of Guinea; 2030–2525 m.

DISCUSSION.—A single specimen of Brisinga hirsuta collected by the Pillsbury off Sao Tome in the Gulf of Guinea allows this first description of the disc and the number of arms; Perrier based the species on a single arm fragment, which was, however, distinctive enough to allow identification of the Pillsbury specimen. This species is clearly distinguished from the other species of Brisinga by the relatively large and numerous marginal plates, the unusual size and distribution of the pedicellariae, and the number and distribution of the adambulacral spines, as well as the well-defined, narrow ridges of spines on the costae.

CHARACTERS.—Those peculiar to B. hirsuta: 300.a, 480.b.

Genus Brisingella Fisher, 1917

Brisingella Fisher, 1917:419; 1918:104; 1919:515.—H.L. Clark, 1920:107.—Fisher, 1928:13; 1940:206.—Hayashi, 1943:152.—Djakanov, 1950:103.—John and A.M. Clark, 1954:151.—McKnight, 1975:235.

DIAGNOSIS.—[30.b] Disc raised slightly above plane of arms; [50.a] interradial arcs acute; [60.b] flesh thin, opaque; [70.a,b] anal opening microscopic or absent; [140.b] bare interradial plates absent; [160.a] no papulae; [170] number of arms 8–12; [190.a] arms deciduous, constricted at base; [220.a] abactinal arm plates imbricate, costae widely spaced, spaces between and beyond costae covered with membrane; [400.a] proximal adambulacrals united interradially; [590.b] 2 pairs of gonads per arm; [600.b] gonads in pairs at base of arm.

TYPE-SPECIES.—Brisinga fragilis Fisher (1906: 1115), by original designation.

CHARACTERS.—Those peculiar to the genus Brisingella: None.

Characters Brisingella shares with other genera of Brisingidae:

| Brisingenes: 70.a,b, 400.a.
| Midgardia: 60.a, 400.a, 590.b, 600.b.
| Novodinia: 60.a, 400.a, 600.b.
| Stegnobrisinga: 70.a,b, 160.a, 400.a, 600.b.

Characters in which Brisingella differs from other genera of Brisingidae:

| Brisingenes: 30, 60, 160, 590, 600.
| Midgardia: 30, 70, 160.
| Novodinia: 30, 70, 160, 590.
| Stegnobrisinga: 30, 60, 590.

Brisingella coronata (G.O. Sars, 1871)

FIGURE 6

Brisinga coronata G.O. Sars, 1871:5.—Thomson, 1873:66 (part).—G.O. Sars, 1875:1–112, pls. 1–6.—Ludwig, 1878:216–234, pl. 15.—Perrier, 1882:61.—Danielssen and Koren, 1884:104.—Carus, 1885:91.—Perrier, 1885d:442–444; 1885b:4–5.—Sladen, 1889:598, 601–604, 832.—Bell, 1889:433.—Sladen, 1891:698.—Bell, 1892[1893]:105.—Norman, 1893:347.—von Marenzeller, 1893a:66–70; 1893b:65–67.—Perrier, 1894:50, 51, 54, 68–70, pl. 1: figs 1, 2, 4–6.—von Marenzeller, 1895a:189–191.—Koehler, 1896a:440; 1896b:38–40.—Perrier, 1896:20.—Ludwig, 1897a:308; 1897b:418–438; 1900:488.

Brisinga sp.—Marion, 1883:129–136.

Brisinga mediterranea Perrier, 1885d:442, 444; 1885b:3–4.—Sladen, 1889:602, 603, 834.—Perrier, 1894:51, 70–71, pl. 1: fig. 3, pl. 3: fig. 1.—von Marenzeller, 1895b:137.—Boone, 1933:93.

Brisingella coronata.—Mortensen, 1927:127.—Grieg, 1927b: 127; 1928:1.—John and A.M. Clark, 1954:151.—Gage et al, 1983:285.

MERISTICS.—Arms = 9–12, R = 310 mm (estimated), r = 7–15 mm, R/r = 20.7/1, length of gonadal region = 75 mm, length of longest arm spine = 20 mm.

FIGURE 6.—Brisingella coronata: a, abactinal view of disc and one arm; b, actinal view of disc. (Adapted from G.O. Sars, 1875, by Molly Griffin.)

DIAGNOSIS.—[80.a] Madreporite large; [100.a] madreporite channeled; [110.c] abactinal disc plates small, tubercular; [120.a,d] abactinal disc plates few, in irregular radiating rows, not dense; [130.b] abactinal disc plates bearing 1 moderately long, mobile, denticulate spine; [170] number of arms 9–12; [205.a,c] gonadal region of arms 75 mm long, slightly to highly inflated; [210.a] abactinal arm plates rod-like; [240.b] costal plates each bearing a long, glassy spine; [250.a] terminal plates small relative to marginals; [260.b] terminal plates shield-shaped; [270.a] terminal plates armed with 8 or 9 outwardly directed webbed spines, central 3 longest; [280.a] marginal plates small; [290.a] marginals irregular; [300.b] marginals occurring about every other adambulacral; [310.b] marginals bearing 1 spinelet; [320.a] adambulacrals higher than long; [330.d] adambulacral plates hourglass-shaped; [340.b] furrow margin slightly indented; [350] number of furrow spines 2–4; [360.a] furrow spines tiny, acicular; [370.c] furrow spines medial; [380] number of subambulacral spines 1 or 2; [390.a] subambulacral spines moderately long, acicular; [420.a] ambulacral plates moderately high; [430.c] ambulacrals broadly Y-shaped; [470.b] tubefeet rather heavy; [480.a] mouth plates rather small relative to adambulacrals; [490.e] mouth plates short, broad; [500] number of preoral spines per mouth plate 2; [510.a] preoral spines acute; [520.a] preoral spines adoral; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines acute; [550.b] lateral oral spines lateral; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines acute; [580.c] suboral spines in center of mouth plate; [620.a,b,c] pedicellariae on both surfaces of disc and arms, in bands or rows on arms; [630.a] all pedicellariae microscopic.

COLOR.—Orange to red.

TYPE.—Bergens Museum; Lofoten, Norway, 549 m.

MATERIAL EXAMINED.—1 specimen, BM(NH), from off Ireland; 1 specimen, USNM 18288, HMS Pola, off Samothraki, Turkey, 40°17′N, 25°13′E, 588 m.

DISTRIBUTION.—Norway to Azores and Cape Verde Islands; Mediterranean; 100–2600 m.

DISCUSSION.—Thanks to G.O. Sars (1875), this species is the best known of the brisingidans. Sars had abundant material, including some young specimens (smallest had disc diameter of 2.5 mm), so some idea of growth changes and variation could be given. An even smaller specimen (∼1 mm), with only eight arms, was collected by the Atlantis II off Libya, and my SEM micrographs (unpublished) show the abundant pedicellariae and the early plate development.

CHARACTERS.—Those peculiar to Brisingella coronata: 110.c, 370.c, 490.e.

Characters B. coronata shares with other species of Brisingella:

B.
  verticellata: 100.a, 130.b, 210.a, 280.a, 330.d, 340.b, 360.a, 390.a, 510.a, 520.a, 570.a, 620.a,b,c, 630.a.

Characters in which B. coronata differs from other species of Brisingella:

B.
  verticellata: 80, 110, 120, 170, 240, 290, 320, 370, 420, 430, 470, 480, 490, 530, 540, 550, 580.

Brisingella verticellata (Sladen, 1889)

FIGURE 7

Brisinga verticellata Sladen, 1889:604–606, pl. 59: figs. 9–11.—Verrill, 1894:283; 1895:211.—H.L. Clark, 1941:64–65.

Brisingella verticellata.—Fisher, 1917:427; 1919:524.

MERISTICS.—Arms = 8, R = 150–180 mm (estimated), r = 5–7.5 mm, R/r = 24–30/1, length of gonadal region = 30–40 mm, length of longest arm spine = 8 mm.

DIAGNOSIS.—[80.b] Madreporite of moderate size, not much raised; [100.a] madreporite flat, channeled; [110.d] abactinal disc plates of numerous small papilliform spinelets embedded in membrane; [120.c] abactinal disc plates imbricate, membrane very thin, easily torn; [130.b] abactinal disc plates armed with spines; [170] number of arms 8; [205.a] gonadal region of arm 30–40 mm long, only slightly inflated; [210.a] abactinal arm plates rod-like; [240.a] abactinal arm plates bearing small, robust, conical spinelets; [280.a] marginal plates small; [290.c] marginals tubercular; [320.b] adambulacral plates longer than high; [330.d] adambulacrals hourglass-shaped; [340.b] furrow margin slightly indented; [350] number of furrow spines 2; [360.a] furrow spines acute, glassy; [370.a,b] one furrow spine proximal, one distal; [380] number of subambulacral spines 1; [390.a] subambulacral spines short, robust, conical; [420.b] ambulacral plates small, delicate; [430.d] ambulacral plates T-shaped, head elongate, much larger than base, imbricating distad, but beneath upper distal projection separated by muscle; [470.a] tubefeet delicate; [480.c] mouth plates moderately large; [490.f] mouth plates slightly longer than broad, with prominent median keel; [500] number of preoral spines per mouth plate 1 or 2; [510.a] preoral spines acute; [520.a] preoral spines adoral; [530] number of lateral oral spines 2; [540.b] lateral oral spines blunt; [550.c] lateral oral spines distal; [560] number of suboral spines per mouth plate 0–2; [570.a] suboral spines acute; [580.a,b] one suboral spine immediately behind preoral spine, one near distal end of mouth plate; [620.a,b,c] pedicellariae on both surfaces of disc and arms, on arms in rows or bands; [630.a] all pedicellariae microscopic.

COLOR.—Ashy gray (in alcohol).

TYPE.—BM(NH) 90.5.7.1041 (holotype): Challenger Sta 46, off New Jersey, 40°17′N, 66°48′W, 2469 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, USNM 6619, Albatross Sta 2102, off Delaware Bay, 2211 m; 1 disc, USNM 7818, Albatross Sta 2205, S of Block Island, Rhode Island, 1962 m; 1 specimen, USNM 9016, Albatross Sta 2076, off Martha's Vineyard, 1657 m; 6 discs, 15 arm fragments, USNM 9064, Albatross Sta 2077, off Georges Bank, 2295 m; arm fragments, USNM 15543, Albatross Sta 2734, off Chesapeake Bay, 1538 m; arm fragments, USNM 15548, Albatross Sta 2732, off Chesapeake Bay, 2107 m; arm fragment, USNM 21486, Albatross Sta 2210, S of Block Island, Rhode Island, 1812 m; 3 arms, USNM 21487, Albatross Sta 2229, off Maryland, 2602 m; 5 arm fragments, USNM 21488, off Delaware, 2136 m; 4 arm fragments, USNM 21989, Albatross Sta 2725, off Chesapeake Bay, 2513 m; 3 specimens, USNM 21910, Albatross Sta 2076, off Georges Bank, 1657 m; 1 specimen, USNM E20949, Alaminos Cr. 68A13–8, Gulf of Mexico, 26°18′N, 96°08′W, 732 m; 1 specimen, USNM E20950, no data.

FIGURE 7.—Brisingella verticellata: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

DISTRIBUTION.—Georges Bank to the Gulf of Mexico, 640–2570 m.

DISCUSSION.—This small, delicate brisingid is chiefly distinguished from B. coronata by its widely spaced costae. The disc is remarkably small, flat, and delicate, with very thin membrane, and no plates, and the arms are unusually slender. The subambulacral spines are short, stout, and conical, rather than long and slender as in Brisingella coronata.

CHARACTERS.—Those peculiar to Brisingella verticellata: None.

Genus Brisingenes Fisher, 1917

Brisingenes Fisher, 1917:419,427; 1919:517.

DIAGNOSIS.—[30.c] Disc flat, in same plane as arms; [50.a] interradial arcs acute; [60.b] rather fleshy, skin thin but conspicuous, opaque, many areas plateless; [70.a,b] anal opening microscopic or absent; [140.b] no bare interradial plates; [160.c] one pair of papulae to each radial area of disc; [170] number of arms 13–15; [190.a] arms deciduous, constricted at base; [220.a] abactinal arm plates imbricate; [400.a] proximal adambulacrals united interradially; [590.a] many gonads; [600.a] gonads serial.

TYPE-SPECIES.—Brisinga mimica Fisher, 1917, by original designation.

CHARACTERS.—Those peculiar to Brisingenes: None.

Characters Brisingenes shares with other genera of Brisingidae:

| Midgardia: 160.c, 400.a.
| Novodinia: 400.a.
| Stegnobrisinga: 60.b, 70.a,b, 400.a.

Characters in which Brisingenes differs from other genera of Brisingidae:

| Midgardia: 30, 60, 70, 590, 600.
| Novodinia: 30, 60, 70, 160, 590, 600.
| Stegnobrisinga: 30, 160, 590, 600.

Brisingenes multicostata (Verrill, 1894)

FIGURE 8

Brisinga multicostata Verrill, 1894:280.

Brisingenes multicostata.—Fisher, 1917:426; 1919:513.

MERISTICS.—Arms = 13–15, R = ∼170–400 mm, r = 13–16 mm, R/r = 13–25/1, length of gonadal region = 52–80 mm, length of longest arm spine = 5–13 mm.

DIAGNOSIS.—[80.a] Madreporite large, raised well above plane of arms; [100.b] madreporite warty, irregular; [110.a] abactinal disc plates round, metapaxillar; [120.b] abactinal disc plates dense in membrane; [130.a] abactinal disc plates bearing 1 or 2 small glassy acicular spinelets; [170] number of arms 13–15; [205.a] gonadal region of arms 52–80 mm long, inflated; [210.a] abactinal arm plates rod-like; [240.a] costal plates bearing transverse ridge of small sharp spinelets; [250.a] terminal plates small, about size of last pair of adambulacrals; [260.b] terminals irregularly shield-shaped; [280.a] marginals small; [290.c] marginals tubercular, more or less fused to adambulacral plates; [310.a] marginals unarmed proximally, beyond midgonadal region bearing 1 stout, fluted, bulbous-based acicular spine; [320.a] adambulacral plates higher than long; [330.c] adambulacrals discoidal; [340.b] furrow margin slightly indented, with distal projection; [350] number of furrow spines 2; [360.a] furrow spines short, sharp; [370.a] furrow spines near proximal end of adambulacrals; [380] number of subambulacral spines 2; [390.a] subambulacral spines long, stout, fluted, acicular; [420.a] ambulacral plates large, heavy; [430.e] ambulacrals dumbbell-shaped; [470.b] tubefeet moderately heavy; [480.a] mouth plates small, narrow; [490.g] mouth plates hourglass-shaped; [500] number of preoral spines 1; [510.a] preoral spines small, acicular; [520.b] preoral spines lateral; [530] number of lateral oral spines 0; [560] number of suboral spines 2; [570.a] suboral spines long, acicular; [580.b,c] one suboral spine near center of mouth plate, one distal; [620.a,b,c] pedicellariae on both surfaces of disc and arms, in rows or bands on arms; [630.b] pedicellariae fairly large.

COLOR.—Unknown.

TYPES.—USNM 12074 and 14858 (syntypes), Albatross Sta 2573, SE of Georges Bank, 3186 m, Sta 2685, S of Martha's Vineyard, 2079 m.

MATERIAL EXAMINED.—Syntypes; 4 arm fragments, USNM 12075, Fishing Vessel Grampus, no other data; 1 specimen, USNM E20951, Gerda Sta 226, Straits of Florida, 24°28′N, 80°16′W, 805 m.

DISTRIBUTION.—Georges Bank to the Straits of Florida; 805–3186 m.

DISCUSSION.—Fisher (1917, 1919) placed this species in his new genus, Craterobrisinga; he evidently did not examine specimens of Verrill's species, and Verrill did not mention in his description the presence of papulae on the disc. These papulae, coupled with the serial gonads and the interradially joined first adambulacrals and marginals, clearly place this species in the genus Brisingenes. It is notable for the very large, slender-valved pedicellariae on the spines.

FIGURE 8.—Brisingenes multicostata: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

Genus Midgardia Downey, 1972

Midgardia Downey, 1972:421; 1973:99.

DIAGNOSIS.—[30.a] Disc thick, raised well above plane of arms; [50.a] interradial arcs acute; [60.a] flesh dense, opaque; [70.c] anal opening conspicuous, surrounded by spinelets; [140.b] no bare interradial plates; [160.c] 2 papulae per radial area of disc; [170] number of arms 11–13; [190.a] arms deciduous, constricted at base; [220.a] abactinal arm plates imbricate, in numerous costae; [400.a] proximal adambulacral plates united interradially; [590.b] 2 pairs of gonads to each arm; [600.b] a pair of gonads on either side of arm.

TYPE-SPECIES.—Midgardia xandaros Downey, 1972, by monotypy.

CHARACTERS.—Those peculiar to Midgardia: None.

Characters Midgardia shares with other genera of Brisingidae:

| Novodinia: 30.a, 60.a, 70.c, 400.a, 600.b.
| Stegnobrisinga: 30.a, 400.a, 600.b.

Characters in which Midgardia differs from other genera of Brisingidae:

| Novodinia: 160, 590.
| Stegnobrisinga: 60, 70, 160, 590.

Midgardia xandaros Downey, 1972

FIGURE 9

Midgardia xandaros Downey, 1972:421–426, fig. 1; 1973:99–100, pl. 48A,B,C.

MERISTICS.—Arms = 11–13, R = 600 mm, r = 14.5 mm, R/r = 41.4/1, length of gonadal region = 40–60 mm, length of longest arm spine = 10 mm.

DIAGNOSIS.—[80.a] Madreporite large; [100.b,c] madreporite warty and irregular, tumid, almost conical; [110.d] abactinal disc plates papilliform; [120.a] abactinal disc plates scattered in membrane; [130.a] abactinal disc plates armed with 1 small prickle-like spinelet; [170] number of arms 11–13; [205.c] gonadal region of arm 40–60 mm long, greatly inflated; [210.a,f] thin flat irregular plates embedded in membrane between costae of rod-like plates, bare membrane beyond gonadal region; [220.a] abactinal arm plates imbricate, in close irregular costae; [240.a] costal plates bearing 1 small prickle-like spinelet; [250.a] terminal plates tiny; [270.b] terminals armed with catsclaw-like spinelets; [280.a] marginals small; [290.c,d] marginals broadly triangular proximally, becoming reduced to tubercles distally; [310.a] marginals bearing very long (up to 25 mm) acicular spine; [320.a,b] adambulacral plates about as high as long; [330.c] adambulacrals vertebra-shaped; [340.b] furrow margin slightly indented; [350] number of furrow spines 3 or 4; [360.a] furrow spines setose, small; [370.a,b] furrow spines at either end of adambulacral plate; [380] number of subambulacral spines 1; [390.a] subambulacral spines large, robust (up to 10 mm), acicular; [420.b] ambulacral plates rather small, delicate; [430.d] ambulacrals T-shaped, with very narrow waist, short stout cylindrical head; [470.b] tube feet very long, heavy, stiffened at base by calcareous collar, probably with limited ability to contract; [480.a] mouth plates small; [490.d] mouth plates slightly longer than broad; [500] number of preoral spines per mouth plate 2 or 3; [510.a] preoral spines slender, delicate, small, acicular; [520.a] preoral spines adoral; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines slender, delicate, small, acicular; [550.c] lateral oral spines distal; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines very large, acicular; [580.c] suboral spines in center of mouth plate; [620.a,b] pedicellariae very numerous, none on abactinal surface of disc, a few in ambulacral groove enlarged, subambulacral and marginal spines bearing a flap of tissue at tip covered with pedicellariae, all spines bearing numerous pedicellariae; [630.a] all pedicellariae small.

COLOR.—Bright red.

TYPES.—USNM E11420 (holotype), E11421 (paratype); Alaminos Sta 69–A–11, Gulf of Mexico, 19°02.6′N, 95°27.5′W, 457 m.

MATERIAL EXAMINED.—Holotype and paratype; 1 specimen, USNM E20984, Explorer Sta BC–I, Gulf of Mexico, ∼16°40′N, 82°50′W, 366 m.

DISTRIBUTION.—Gulf of Mexico, 366–457 m.

DISCUSSION.—The two very large specimens of this species taken in a single dredge haul by the Texas A&M vessel Alaminos were mature and shedding gametes when collected. The eggs were large, yolky, and orange; the male gametes were bright red.

CHARACTERS.—Those peculiar to M. xandaros: None.

FIGURE 9.—Midgardia xandaros: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

Genus Novodinia Dartnell, Pawson, Pope, and Smith, 1969

Odinia Perrier, 1885c:9.—Sladen, 1889:598.—Bell, 1982 [1893]: 105.—Perrier, 1894:71.—Verrill, 1894:279; 1895:211.—Koehler, 1895b:420.—Fisher, 1906:1108; 1916b:31; 1917:419.—A.H. Clark, 1916:57.—Fisher, 1918:105; 1919:505.—A.H. Clark, 1934:1.—Hayashi, 1943:136.

Novodinia Dartnell et al., 1969:211 [replacement name].

TYPE-SPECIES.—Odinia semicoronata Perrier, 1885c, by subsequent designation (Fisher, 1917).

DIAGNOSIS.—[30.a] Disc thick, raised well above plane of arms; [50.a] interradial arcs acute; [60.a] flesh dense, opaque, with some plateless areas; [70.c] anal opening usually visible; [140.b] no bare interradial plates; [150.b] papulae numerous, scattered on disc and proximal (gonadal) portion of arms; [160.b] papulae single, long, vermiform; [170] number of arms 13–20; [190.a] arms deciduous, constricted at base; [220.a] abactinal arm plates imbricate; [400.a] first 3–5 pairs of adambulacrals united interradially; [590.c] one pair of gonads to each arm; [600.b] gonads paired.

DISCUSSION.—Unlike most brisingidans, Novodinia is a genus of only moderately deep waters (~250–1500 m). The feeding posture, arms held up into the current over the disc, has been observed and filmed by Pawson et al. (in prep.).

CHARACTERS.—Those peculiar to Novodinia: 160.b.

Characters Novodinia shares with other genera of Brisingidae:

Stegnobrisinga: 30.a, 400.a, 590.c, 600.b.

Characters in which Novodinia differs from other genera of Brisingidae:

Stegnobrisinga: 60, 70, 160.

Novodinia americana (Verrill, 1880)

FIGURE 10

Brisinga americana Verrill, 1880:139; 1894:279; 1895:211.

MERISTICS.—Arms = 15–20, R = 350 mm, r = 12–16 mm, R/r = 22–29/1, length of gonadal region = 45–50 mm, length of longest arm spine = 10 mm.

DIAGNOSIS.—[80.b] Madreporite of moderate size, raised well above plane of arms; [100.a] madreporite button-like, channeled; [110.b] abactinal disc plates tumid, irregularly rounded; [130.a] abactinal disc plates bearing 2–4 small acute spinelets; [170] number of arms 15–20; [205.c] gonadal region of arms 45–50 mm long, high, inflated, laterally compressed, rest of arm depressed and tapering gradually to slender tip; [210.g] costal plates of gonadal region of arm cruciform, between costae membrane embedded with flat, heavy, bizarrely shaped imbricating plates; [220.a] abactinal arm plates imbricate, beyond gonadal region, which ends abruptly, arms covered with plateless membrane; [240.b] most costal plates bearing single short acicular spine; [280.b] marginal plates large relative to adambulacrals; [290.g] marginals cruciform; [300.b] one marginal to each adambulacral on short portion of arm before gonadal inflation, beyond occurring only about every fifth adambulacral; [310.a] marginals unarmed proximally, just beyond highest part of gonadal inflation marginals bear single stout acicular spine, beyond gonadal region marginals bear a transverse series of about 5 long acicular spines; [330.c] adambulacral plates discoidal; [350] number of furrow spines 0; [390.c] form of subambulacral spines long, stout, truncate; [400.a] first 2–4 adambulacrals united interradially; [420.b] ambulacral plates small, delicate; [430.f] head of ambulacrals not much enlarged, discoidal; [440.b] ambulacrals rather crowded; [470.b] tubefeet very heavy; [480.c] mouth plates large; [490.a,c] mouth plates triangular, pair forming broadly rounded T-shape, oral margin broad, in contact across ambulacral groove; [500] number of preoral spines per mouth plate 3; [510.a] preoral spines small, acicular; [520.b] preoral spines lateral; [530] number of lateral oral spines per mouth plate 1–3; [540.a] lateral oral spines small, similar to preoral spines, acicular; [550.b] lateral oral spines lateral; [560] suboral spines absent; [620.a,b] pedicellariae on both surfaces of disc and arms; [630.a] all pedicellariae small.

FIGURE 10.—Novodinia americana: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

COLOR.—“Pale orange-red in alcohol when received, soon fading to whitish…living, probably bright red.” (Verrill, 1880:139).

TYPE.—USNM 33038 (holotype), Gloucester Fisheries Donation, Banquereau Bank, 320 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, Gloucester Fisheries Donation, Banquereau Bank, 732 m; 3 specimens, Pillsbury Sta 776, off Colombia, 12°13.3′N, 72°50′W, 408–576 m.

DISTRIBUTION.—Nova Scotia to Colombia, 320–732 m.

DISCUSSION.—Examination of Verrill's type and additional material in the NMNH shows that this species clearly belongs in the genus Novodinia. The disc was unknown, as Verrill described N. americana solely on the basis of the arms. The three specimens collected by the Pillsbury, although considerably smaller than the type (e.g., height of gonadal inflation 12 mm, vs 32 mm in the type), undoubtedly belong to this species, and allow the first description of the disc. They also represent a considerable extension of geographic range.

CHARACTERS.—Those peculiar to N. americana: 440.b.

Characters N. americana shares with other species of Novodinia:

N. |   antillensis: 110.b, 220.a, 300.b, 330.c, 390.c, 420.b, 430.f, 480.c, 510.a, 520.b, 540.a, 550.b, 560.0, 620.a,b, 630.a.
N. |   homonyma: 100.a, 220.a, 300.b, 330.c, 420.b, 430.f, 470.b, 480.c, 520.b, 540.a, 620.a,b, 630.a.
N. |   pandina: 130.a, 205.c, 220.a, 280b, 300.b, 330.c, 420.b, 430.f, 530.1–3, 630.a.
N. |   semicoronata: 130.a, 210.g, 240.b, 300.b, 330.c, 420.b, 480.c, 520.b, 540.a, 550.b, 620.a,b, 630.a.

Characters in which N. americana differs from other species of Novodinia:

N. |   antillensis: 80, 100, 130, 280, 290, 470.
N. |   homonyma: 80, 110, 205, 210, 240, 290, 390, 490, 500, 510,560.
N. |   pandina: 80, 100, 110, 210, 240, 290, 470, 480, 490, 510, 520, 540, 560, 620.
N. |   semicoronata: 80, 100, 110, 290, 510.

Novodinia antillensis (A.H. Clark, 1934)

FIGURE 11

Odinia antillensis A.H. Clark, 1934:1–3, pl. 1.—Downey, 1973:99, pl. 47c,d.

MERISTICS.—Arms = 13–17, R = 80–250 mm, r = 4–18 mm, R/r = 14–20/1, length of gonadal region = 20–40 mm, length of longest arm spine = 9 mm.

DIAGNOSIS.—[80.c] Madrcporite rather small, raised; [100.e] madreporite with meandering gyri like a brain coral, surrounded by enlarged spine-bearing plates; [110.b] abactinal disc plates close, irregular, tumid, interradially a double row of flat, irregular, more or less bare plates extending from abactinal margin down beveled lateral side of disc and out a short distance on base of arms, which are basally fused; these are the more or less rectangular marginals of the first 4–6 adambulacrals, fused interradially; [130.b] abactinal disc plates bearing 4–10(2–16) small blunt spines; [170] number of arms 13–17; [205.c] gonadal region of arm 20–40 mm long, greatly inflated, arms tapering distally; [210.a,d,g] abactinal arm plates cruciform or rod-like, costae of gonadal region irregular, membrane between costae filled with flat, irregularly shaped plates, bare membrane beyond gonadal region; [220.a] abactinal arm plates imbricating in transverse bands; [240.b,c] many plates of costae bearing a long, slender acicular spine mounted on a tubercle; [280.a] marginal plates small; [290.d,e,h] about first 10 marginal plates flat, rectangular, rest subtriangular; [300.b] about first 10 marginals occurring on every adambulacral plate, on gonadal region proper, on every other adambulacral, and on rest of arm, about every 3–4 adambulacral; [310.a] first 10 marginals unarmed, rest bearing transverse group of 3 long slender acicular spines mounted on large, knob-like tubercles; [320.a] adambulacrals higher than long; [330.c] adambulacral plates discoidal; [350] number of furrow spines 1; [360.a] furrow spines tiny, acute; [370.b] furrow spines on distal prolongation of adambulacral; [390.c] subambulacral spines long, thick, truncate, mounted on tubercle; [400.a] about first 2–6 adambulacral plates united interradially by symphasis to those of adjoining arm, as are the corresponding marginals; [420.b] ambulacral plates small, delicate; [430.f] ambulacrals discoidal; [470.a] tubefeet moderately delicate; [480.c] mouth plates large; [490.a] mouth plate pear-shaped like a broadly rounded T, with lateral prolongation meeting adjoining mouth plates; [500] number of preoral spines per mouth plate 3–4; [510.a] preoral spines small, acicular; [520.b] preoral spines lateral; [530] number of lateral oral spines per mouth plate 1 or 2; [540.a] lateral oral spines small, acicular; [550.b] lateral oral spines lateral; [560] number of suboral spines per mouth plate 0; [620.a,b] pedicellariae on both surfaces of disc and arms; [630.a] all pedicellariae small.

FIGURE 11.—Novadinia antillensis: a, abactinal view; b, actinal view.

COLOR.—Unknown.

TYPE.—USNM E3266 (holotype), Caroline Sta 47, W of Puerto Rico, 18°17.2′N, 67°25′W, 512–622 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, USNM E12621, Oregon Sta 4574, 23°13′N, 87°50′W, 384 m; 1 specimen, USNM E12622, Oregon Sta 4480, 11°10′N, 65°07′W, 402 m; 1 specimen, USNM E12913, Oregon Sta 1408, 28°02′N, 90°15′W, 366 m.

DISTRIBUTION.—West Indies, Gulf of Mexico, 366–622 m.

DISCUSSION.—A quite small specimen (USNM E12621) has only incipient costae and only 1–3 marginal spines. The adambulacral spines are proportionally larger and decidedly capitate.

CHARACTERS.—Those peculiar to N. antillensis: None.

Characters N. antillensis shares with other species of Novodinia:

N. |   homonyma: 80.c, 220.a, 300.b, 330.c, 420.b, 430.f, 480.c, 520.b, 540.a, 620.a,b, 630.a.
N. |   pandina: 100.e, 205.c, 220.a, 300.b, 330.c, 420.b, 430.f, 470.a, 630.a.
N. |   semicoronata: 80.c, 100.e, 300.b, 320.a, 330.c, 420.b, 480.c, 490.a, 520.b, 540.a, 620.a,b, 630.a.

Characters in which N. antillensis differs from other species of Novodinia:

N. |   homonyma: 100, 110, 130, 205, 210, 290, 390, 470, 490, 500, 510, 560.
N. |   pandina: 80, 110, 130, 210, 240, 280, 290, 480, 490, 510, 520, 540, 560, 620.
N. |   semicoronata: 110, 130, 205, 290, 350, 500, 510.

Novodinia homonyma, new name

FIGURE 12

Brisinga elegans Perrier, 1885c:9 [non Brisinga elegans Verrill, 1884:382.]

Odinia elegans.—Perrier, 1894:71–74, pl. 4: fig. 4.

MERISTICS.—Arms = 19, R = 40–79 mm, r = 4–10 mm, R/r = 6.6–10/1, length of gonadal region = 8–16 mm (estimated), length of longest arm spine = 4 mm (estimated).

DIAGNOSIS.—[80.c] Madreporite small, inconspicuous; [100.a] madreporite channeled; [110.f] abactinal disc plates polygonal, thin, flat; [120.c] abactinal disc plates imbricate; [130.a,c] abactinal disc plates bearing 1–3 tubercles with short, capitate spinelets; [170] number of arms 19; [205.a] gonadal region of arms 8–16 mm long, moderately inflated; [210.e,f] abactinal arm plates polygonal, thin, beyond gonadal region arms covered with smooth skin with occasional bands of thin, transparent plates; [220.a] abactinal arm plates imbricate, only a few rudimentary costae on distal portion of gonadal region; [240.a,c] plates of gonadal region of arm with tubercle bearing single moderately long acicular spine; [280.a,b] first 6 marginals rather large, rest quite small; [290.f] marginals squarish; [300.b] first 6 marginals above every adambulacral, closely appressed to those of adjoining arms, their upper edges forming a bare sulcus interradially on disc, other marginals about every third adambulacral; [310.a] first 6 marginals unarmed, others bearing 2 extremely long acicular spines; [320.a,b] adambulacral plates higher than long proximally, longer than high distally; [330.c] adambulacral plates discoidal; [350] furrow spines absent; [390.a] subambulacral spines moderately long, acicular; [420.b] ambulacral plates small, delicate; [430.f] ambulacrals discoidal; [470.b] tubefeet moderately heavy; [480.c] mouth plates large; [490.g] mouth plates long, narrow, hourglass-shaped; [500] number of preoral spines per mouth plate 2; [510.b] preoral spines short, conical; [520.b] preoral spines lateral; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines small, acicular; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines acute; [580.b] suboral spines below center of mouth plate; [620.a,b] pedicellariae on both surfaces of disc and arms; [630.a] all pedicellariae small.

FIGURE 12.—Novodinia homonyma: a, abactinal view; b, actinal view. (From Perrier, 1894.)

COLOR.—Unknown.

TYPE.—Paris Museum 4623 (holotype), Talisman Sta 73, S of Canary Islands, 1056–1435 m.

MATERIAL EXAMINED.—Holotype.

DISTRIBUTION.—Known only from type-locality.

DISCUSSION.—Perrier's Brisinga elegans (1885c) (= his Odinia elegans (1894)) is an invalid homonym of Verrill's Brisinga elegans (1884) (= Freyella elegans) and must therefore be replaced. The name homonyma seems both appropriate and euphonic.

The disc in this species is very flat, and the papulae on the disc occur in regular radiating rows.

CHARACTERS.—Those peculiar to N. homonyma: None.

Characters N. homonyma shares with other species of Novodinia:

N. |   pandina: 120.c, 220.a, 300.b, 330.c, 350.0, 420.b, 430.f, 490.g, 510.b, 560.1, 570.a, 630.a.
N. |   N. semicoronata: 80.c, 120.c, 205.a, 280.a,b, 300.b, 330.c, 350.0, 420.b, 480.c, 510.b, 520.b, 530.1, 540.a, 570.a, 620.a,b, 630.a.

Characters in which N. homonyma differs from other species of Novodinia:

N. |   pandina: 80, 100, 110, 205, 470, 480, 520, 540, 620.
N. |   semicoronata: 100, 110, 210, 240, 290, 390, 490, 500.

Novodinia pandina (Sladen, 1889)

FIGURE 13

Odinia pandina Sladen, 1889:598–601, pl. 59: figs 1–5.—Bell, 1893a:105.

MERISTICS.—Arms = 13–18, R = 150–215 mm, r = 9–15 mm, R/r = 14.3–16.7/1, length of gonadal region = 40–48 mm, length of longest arm spine = 7 mm.

DIAGNOSIS.—[80.a] Madreporite convex, prominent; [100.e] madreporite covered with fine striations; [110.g] abactinal disc plates sturdy, lobed; [120.c] abactinal disc plates imbricate; [130.a] abactinal disc plates bearing 1–14 small, stout, rough-tipped spinelets; [170] number of arms 13–18; [180.a,b] arms robust, rounded in cross-section; [205.c] gonadal region of arms 40–48 mm long, highly inflated; [210.f] abactinal arm plates flat; [220.a] gonadal region of arms covered with thick coriaceous skin, underlaid with small imbricating plates, outer half of gonadal region transversed by 5–7 costae of small prominent imbricating plates, membrane between costae smooth and without pedicellariae or spinelets, beyond gonadal region arms covered with thin membrane bearing small roundish patches of tiny pedicellariae; [240.a] costal plates bearing isolated, robust, conical or clavate spinelets; [280.b] marginal plates large; [290.c,f] beyond first few, marginals transversely elongate, forming a ridge of 2–6 stout, knobby tubercles (the appearance is of a few ankylosed plates, rather than a single plate), first 8–10 marginals rectangular, unarmed, appressed to those of adjoining arm; [300.b] marginals occurring about every fourth adambulacral; [310.a] marginals armed with 2–6 (usually 3) extremely long, acicular spines; [320.b] adambulacrals small, longer than high; [330.c] adambulacrals discoidal; [350] number of furrow spines 0; [390.a,c] subambulacral spines long, stout, chisel-shaped proximally, acicular distally; [400.a] first two adambulacrals joined interradially by a symphasis; [420.b] ambulacral plates small, delicate; [430.f] ambulacral plates discoidal; [470.a] tubefeet rather delicate; [480.a] mouth plates small; [490.g] mouth plates somewhat hourglass-shaped; [500] number of preoral spines per mouth plate 1–3; [510.b] preoral spines short, conical, dogtooth-shaped; [520.a] preoral spines adoral; [530] number of lateral oral spines per mouth plate 1–3; [540.b] lateral oral spines short, conical, touching or overlapping adjacent mouth plate spines, more or less closing ambulacral groove; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines acute; [620.a,g] no pedicellariae on disc, pedicellariae on membranous sheath covering spines and spinelets on arms, tiny pedicellariae in sacculi on bare membrane of distal arms; [630.a] all pedicellariae small.

COLOR.—Yellowish white to pinkish (in alcohol).

TYPE.—BM(NH) 90.5.7.1035 (holotype), Lightning Sta 7, Faeröe Channel, 60°07′N, 5°21′W, 914 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, USNM E15535, off North Carolina, 32°46.7′N, 77°42.3′W, 278 m.

DISTRIBUTION.—Faeröe Channel; North Carolina; 278–914 m.

DISCUSSION.—The recent discovery of a specimen of this species from off North Carolina considerably extends the range of the species.

CHARACTERS.—Those peculiar to N. pandina: 620.g.

Characters N. pandina shares with other species of Novodinia:

FIGURE 13.—Novodinia pandina: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

N.
  semicoronata: 100.e, 110.g, 120.c, 130.a, 300.b, 330.c, 350.0, 420.b, 510.b, 570.a, 630.a.

Characters in which N. pandina differs from other species of Novodinia:

N.
  semicoronata: 80, 180, 205, 210, 240, 290, 320, 480, 490, 520, 540, 620.

Novodinia semicoronata (Perrier, 1885)

FIGURE 14

Brisinga semicoronata Perrier, 1885c:9.

Brisinga robusta Perrier, 1885:11.

Odinia semicoronata.—Perrier, 1894:75–78, pl. 5.

Odinia robusta.—Perrier, 1894:78–81, pl. 6.—Koehler, 1895b:420.

MERISTICS.—Arms = 15–17, R = 275 mm, r = 15 mm, R/r = 18.3/1, length of gonadal region = 40 mm, length of longest arm spine = 5 mm.

DIAGNOSIS.—[80.c] Madreporite small; [100.e] madreporite with deep gyri; [110.g] abactinal disc plates lobed; [120.c] abactinal disc plates imbricate; [130.a] abactinal disc plates bearing 3(1–4) moderately long acute or bifurcate spinelets; [170] number of arms 15–17; [180.b] arms long, robust; [205.a] gonadal region of arms 40 mm long, moderately inflated; [210.g] abactinal arm plates lobed, in usually incomplete costae extending beyond gonadal region for a short distance, between costae thick skin concealing lobed plates, thick skin beyond costae plateless; [240.b] plates of costae bearing acute spines; [250.b] terminal plates large; [260.a] terminals round, convex; [270.a] terminals bearing about 6 long, slender, acute spines; [280.a,b] first few marginals large, others small; [290.b] marginals elongate; [300.b] marginals occurring about every 2–4 adambulacrals; [310.a] marginals bearing 1(2) rather short acute spine(s); [320.a] adambulacral plates higher than long; [330.c] adambulacrals discoidal; [350] furrow spines absent; [390.b,c,d] subambulacral spines long, stout, capitate, truncate or bifurcate; [400.a] first 3 or 4 adambulacrals united interradially; [420.b] ambulacral plates small, delicate; [480.c] mouth plates large; [490.a] mouth plates more or less T-shaped, longer than broad, with lateral extension meeting adjoining mouth plate across ambulacral groove; [500] number of preoral spines per mouth plate 1;[510.b] preoral spines small, conical; [520.b] preoral spines lateral; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines small, acute; [550.b] lateral oral spines lateral; [560] number of suboral spines per mouth plate 1 (not always present); [570.a] suboral spines long, thick, acicular; [620.a,b] pedicellariae on both surfaces of arms and disc; [630.a] all pedicellariae small.

COLOR.—Unknown.

TYPES.—Paris Museum 4630–4631 (2 syntypes), Talisman Sta 73, off Senegal, 25°39′N, 18°26′W, 1056–1435 m.

MATERIAL EXAMINED.—Syntypes; holotype of Brisinga robusta, Paris Museum, Talisman Sta 72, off Senegal, 25°39′N, 18°22′W, 882 m.

DISTRIBUTION.—Known only from off Senegal, 882–1435 m.

DISCUSSION.—Despite the paucity of available material, there can be little doubt that the specimens described by Perrier as Brisinga semicoronata are conspecific with the two specimens described by him as B. robusta from approximately the same locality. Differences (size, length of abactinal spines, one vs one or two adambulacral spines) can be accounted for by ontogeny. The particularly thick skin with embedded lobed plates between the costae is characteristic. The name semicoronata has been chosen over robusta because it has page priority.

The papulae on the gonadal regions of the arms are arranged in irregular transverse rows.

CHARACTERS.—Those peculiar to N. semicoronata: None.

Genus Stegnobrisinga Fisher, 1916

Stegnobrisinga Fisher, 1916b:34; 1917:428.—H.L. Clark, 1926:31.—A.M. Clark and Courtman-Stock, 1976:98.

DIAGNOSIS.—[30.a] Disc thick, raised well above plane of arms; [50.a] interradial arcs acute; [60.b] skin thin, opaque, some areas plateless; [70.a,b] anal opening microscopic or absent; [140.b] no bare interradial plates; [160.a] no papulae; [170] number of arms 11–14; [190.a] arms deciduous, constricted at base; [205.b] gonadal region of arms broad, flat; [220.a] abactinal arm plates imbricate; [400.a] proximal adambulacrals united interradially; [590.c] 2 pairs of gonads per arm; [600.b] gonads paired.

FIGURE 14.—Novodinia semicoronata: a, abactinal view; b, actinal view (from Perrier, 1894).

TYPE-SPECIES.—Stegnobrisinga placoderma Fisher, 1916b, by original designation.

CHARACTERS.—Those peculiar to the genus Stegnobrisinga: None.

Stegnobrisinga splendens H.L. Clark, 1926

FIGURE 15

Stegnobrisinga splendens H.L. Clark, 1926:31–33, pl. 7: figs. 3–4.—A.M. Clark and Courtman-Stock, 1976:98.

MERISTICS.—Arms = 11–14, R = 145–500 mm, r = 9–19 mm, R/r = 16–26/1, length of gonadal region = 22–65 mm, length of longest arm spine = 16 mm.

DIAGNOSIS.—[80.a] Madreporite large, raised, conical; [100.a] madreporite coarsely channeled; [110.a] disc membrane set with small roundish plates; [120.b] abactinal disc plates dense in membrane; [130.a] abactinal disc plates bearing tiny spinelet; [170] number of arms 11–14; [205.a] gonadal region of arms 22–65 mm long, broad, somewhat flat; [210.b,d,f] abactinal arm plates somewhat tumid, irregular, costae low, close, between costae flat irregular plates, beyond gonadal region a few incomplete costae, thin irregular plates, then bare membrane; [240.a] many costal plates bearing single short sharp spinelet, other abactinal arm plates unarmed; [250.a] terminal plates minute; [270.c] terminal plates possibly unarmed; [280.a] marginal plates small; [290.c,g] marginals lobate, with central tubercle; [320.a,b] adambulacral plates about as long as high; [330.f] adambulacrals dumbbell-shaped; [340.c] furrow margin deeply indented; [350] number of furrow spines 4; [360.a] furrow spines fairly long, fine, setose; [370.a,b] 2 furrow spines on distal end of adambulacral, 2 on proximal end; [380] number of subambulacral spines 1; [390.a] subambulacral spines long, acicular; [420.a] ambulacral plates large, sturdy; [430.g] ambulacral plates l-shaped; [470.a] tubefeet rather delicate; [480.a] mouth plates small; [490.b] mouth plates nearly rectangular, with only slight curved indentation on outside lateral edge; [500] 2 preoral spines per mouth plate; [510.a] preoral spines small, acicular, delicate; [520.a] preoral spines adoral; [530] number of lateral oral spines per mouth plate 1; [540.a] lateral oral spines small, acicular; [550.b] lateral oral spines lateral; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines huge, acicular, ridged; [580.b] suboral spines below center of mouth plate; [620.a,b,c] pedicellariae on disc plates, in bands on arms, and on membranous spine covering; [630.b] all pedicellariae large.

COLOR.—Vermilion.

TYPE.—South African Museum (holotype), Pickle Sta 405, Natal, ESE from Durban, 860 m.

MATERIAL EXAMINED.—1 specimen, USNM E20798, Pillsbury Sta 675, off Surinam, 08°26′N, 54°17′W, 1234–1271 m; 1 specimen, USNM E20954, Pillsbury Sta 636, Straits of Florida, 23°54′N, 81°27′W, 3400 m; 1 specimen, USNM E20955, Pillsbury Sta 892, West Indies, 14°17′N, 60°45′W, 4000 m; arm fragment, USNM E20953, Gerda Sta 1112, Straits of Florida, 23°44′N, 81°14′W, 2275–2359 m; 2 specimens, Atlantis II Sta 42–192, Walvis Bay, Angola, 23°02′S, 12°19′E, 2117–2154 m; 1 specimen, uncatalogued, USNM, Pillsbury Sta 747, 11°46′N, 67°05.7′W, 1174 m.

DISTRIBUTION.—South Africa to Angola, Straits of Florida to Venezuela; 860–4000 m.

DISCUSSION.—This species, hitherto known only from South Africa, was collected by the Atlantis II off Angola and in the Straits of Florida, the West Indies, and off Venezuela by the Pillsbury and Gerda. The principal (and only significant) difference between this species and Fisher's (1916b:33) Stegnobrisinga placoderma from the China Sea is in the distribution of the marginal plates; also, Fisher's specimens were smaller.

FIGURE 15.—Stegnobrisinga splendens: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

CHARACTERS.—Those peculiar to S. splendens: 430.g.

Family FREYELLIDAE, new family

DIAGNOSIS.—[50.b] Interradial arcs gradually curved; [80.c] madreporite small; [140.a] bare interradial plates present on disc; [190.b] abactinal surface of proximal arms continuous with that of disc; [220.b] abactinal arms plates not forming costae on proximal part, but a mosaic of abutting plates; [400.b] proximal adambulacrals not united interradially.

DISCUSSION.—The preceding characters separate the Freyellidae from the Brisingidae. In addition, because of the bare interradial plates and the plating on the proximal part of the arms, the Freyellidae have a more limited ability to raise the arms above the disc. They are usually found on soft substrates, and in deeper water (950–5618 m), in contrast to the Brisingidae, which seem to prefer hard substrates and frequently occur in shallower water (100–4500 m). Some of the Freyellidae (e.g., Freyastera) are probably not suspension feeders, as they lack the ability to raise their arms above the disc, while others can raise the arms from a point beyond the gonadal region, but not directly from the juncture with the disc as can members of the Brisingidae.

Genus Colpaster Sladen, 1889

Colpaster Sladen, 1889:647.

DIAGNOSIS.—[30.b] Disc raised slightly above plane of arms; [50.b] interradial arcs gently curved; [60.b] skin thin, opaque; [70.c] anal opening conspicuous; [140.a] conspicuous bare plate interradially, on vertical margin of disc; [160.c] 2 papulae in each radial area of disc; [170] number of arms 7–12; [190.b] abactinal surface of gonadal region of arms continuous with that of disc, arms not constricted at base; [220.a] abactinal arm plates imbricate; [400.b] proximal adambulacrals not united interradially; [590.b] 1 pair of gonads per arm; [600.b] gonads paired.

TYPE-SPECIES.—Colpaster scutigerula Sladen, 1889, by monotypy.

DISCUSSION.—Although I was unable to locate any specimens of Freyella edwardsi Perrier there is little doubt that this species belongs to Colpaster. Based on the descriptions and illustrations of Perrier (1882, 1894) and Koehler (1909a), C. edwardsi possesses the bare, smooth interradial plates on the margin of the disc, two small papulae over each arm (not described, but inferred from the illustrations), the small madreporite, moderately inflated gonadal region of the arms, similar plating on disc and gonadal region of arms, two very small furrow spines, and a similar distribution of pedicellariae.

CHARACTERS.—Those peculiar to the genus Colpaster: None.

Characters Colpaster shares with other genera of Freyellidae:

| Freyastera: 70.c, 400.b.
| Freyella: 30.b, 60.b, 590.b, 600.b.

Characters in which Colpaster differs from other genera of Freyellidae:

| Freyastera: 30, 60, 160, 170, 590, 600.
| Freyella: 70, 160, 220, 400.

Colpaster edwardsi (Perrier, 1882)

FIGURE 16

Brisinga edwardsi Perrier, 1882:61.

Freyella edwardsi.—Perrier, 1885d:441; 1894:82–84, pl. 1: fig. 7.—Sladen, 1889:616,617.—Koehler, 1907:6; 1909c:124–127, pl. 23: figs 1, 2.—Mortensen, 1921:128, fig. 74.

MERISTICS.—Arms = 12, R = 60–80 mm (estimated), r = 11 mm, R/r = 6.4/1, length of gonadal region = 25–40 mm, length of longest arm spine = 4–5 mm.

DIAGNOSIS.—[80.c] Madreporite small but raised and conspicuous; [100.b] madreporite irregular; [110.a] abactinal disc plates small, metapaxillar; [120.b] abactinal disc plates dense in membrane; [130.a] abactinal disc plates bearing tiny, almost granulose, blunt, rough spinelets; [170] number of arms 12; [205.a] gonadal region of arms 25–40 mm long, moderately inflated; [210.c,d] abactinal arm plates metapaxillar, like disc plates, from midgonadal region on, abactinal arm plates tend to form costae or arcs across arm, with irregular plates between costae; [240.a] abactinal arm plates bearing 1–10 spinelets, larger, stouter and more numerous than those of disc plates; [280.a] marginal plates small; [290.a,h] marginal plates irregular, tumid; [310.a] marginals bearing a short, acute spine beyond gonadal region; [320.a] adambulacral plates higher than long; [330.a] adambulacral plates cylindrical; [380] number of subambulacral spines 1; [390.a,c] about first 10 adambulacrals bearing long, stout, truncate subambulacral spine, rest with long, aciculate spine; [530] lateral oral spines absent; [560] number of suboral spines per mouth plate 1; [620.a,b,c] pedicellariae on both surfaces of disc and arms, in rows or bands on arms.

FIGURE 16.—Colpaster edwardsi (from Koehler, 1909c).

COLOR.—Unknown.

TYPE.—Not found; Golfe de Gascogne, 2300 m.

MATERIAL EXAMINED.—None.

DISTRIBUTION.—Golfe de Gascogne, Canary Islands, 1700–2300 m.

DISCUSSION.—I could find no existing material of this species, and neither Perrier nor Koehler discussed several important characters; furthermore they did not figure the mouth plates or pedicellariae. However, despite the lack of specimens, there is little doubt that the species is indeed a Colpaster.

CHARACTERS.—Those peculiar to Colpaster edwardsi: None.

Characters which C. edwardsi shares with other species of Colpaster:

C. scutigerula: 110.a, 130.a, 240.a, 280.a, 320.a, 330.a, 620.a,b,c.

Characters in which C. edwardsi differs from other species of Colpaster:

C. scutigerula: 80, 100, 120, 170, 205, 210, 380, 390.

Colpaster scutigerula Sladen, 1889

FIGURE 17

Colpaster scutigerula Sladen, 1889:648–650, pl. 47: figs. 4–7.

MERISTICS.—Arms = 7–10, R = 135+ mm (estimated), r = 6–10 mm, R/r = 13.5–22.5/1, length of gonadal region = 18–31 mm, length of longest arm spine = 4–7 mm.

DIAGNOSIS.—[80.a] Madreporite large; [100.a] madreporite with a few coarse irregular channels; [110.a] abactinal disc plates small, round, metapaxillar; [120.c] abactinal disc plates imbricate; [130.a] abactinal disc plates bearing 3–6 short, capitate spinelets; [170] number of arms 7–10, robust in proportion to disc; [205.b] gonadal region of arms 18–31 mm long, not inflated; [210.b] abactinal arm plates tumid; [240.a] abactinal arm plates bearing 3–6 short capitate spinelets; [280.a] marginal plates small; [290.d] marginals triangular; [310.a] marginals bearing one long, very robust, frequently capitate spine; [320.a,b] adambulacral plates about as long as high; [330.a] adambulacral plates cylindrical; [340.a] furrow margin straight; [350] number of furrow spines 2, small; [360.a] furrow spines acute; [370.a,b] one furrow spine proximal, one distal; [380] number of subambulacral spines 2; [390.b] subambulacral spines large, robust, frequently capitate; [420.b] ambulacral plates rather small, delicate; [430.d] head of ambulacrals cylindrical, much larger than base; [470.b] tubefeet rather heavy; [480.a] mouth plates small, strong; [490.c,f] mouth plates broader distally than proximally, with pronounced keel distally; [500] number of preoral spines per mouth plate 2; [510.b] preoral spine small, blunt, almost club-shaped; [520.b] preoral spines laterally placed, nearly closing ambulacral groove; [530] number of lateral oral spines per mouth plate 1 (not always present); [540.b] lateral oral spines blunt, similar to preoral spines; [550.a] lateral oral spines proximal on mouth plate; [560] number of suboral spines per mouth plate 1–3; [570.a] suboral spines acute; [580.a] suboral spines above center of mouth plate; [620.a,b,c] pedicellariae on both surfaces of disc and arms, in bands or rows on arms; [630.a] all pedicellariae quite small.

FIGURE 17.—Colpaster scutigerula: a, abactinal view; b, actinal view.

COLOR.—Ashy gray with traces of violet (in alcohol).

TYPE.—BM(NH) 1890.5.7.1081 (holotype); Challenger Sta 3, SW of Canary Islands, 25°45′N, 20° 14′W, 2789 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, USNM E20796, Pillsbury Sta 266, Gulf of Guinea, 1°13′N, 46°E, 2525 m; 1 disc, Oregon II Sta 11217, off Honduras, 26°26′N, 83° 31′W, 933 m.

DISTRIBUTION.—S of Canary Islands; Gulf of Guinea; off Honduras; 933–2789 m.

DISCUSSION.—This species is distinguished mainly by the conspicuous shield-shaped interradial plates and the unusually long and numerous spines of the mouth plates. The specimen from the Gulf of Guinea differs slightly from the type (number of arms, more curved and blunt preoral spines, frequent absence of furrow spines), but I believe that it belongs to this species. Similarly, the specimen from off Honduras (disc only) obviously belongs to this species. It is gratifying to have two more specimens of this peculiar seastar, which has not been seen since the Challenger specimen was collected.

CHARACTERS.—Those peculiar to Colpaster scutigerula: None.

Genus Freyastera, new genus

Freyella.—Korovchinsky and Galkin, 1984:1205–1215 [part].

DIAGNOSIS.—[30.c] Disc flat, in same plane as arms; [50.b] interradial arcs broadly curved; [60.c] flesh thin, delicate, slimey; [70.c] anal opening conspicuous; [140.a] bare interradial plates present; [160.a] no papulae; [170] number of arms normally 6; [190.b] abactinal surface of proximal arms continuous with that of disc, arms not constricted at base; [220.a] abactinal arm plates imbricate; [400.b] proximal adambulacrals not united interradially; [590.c] 1 pair of gonads per arm; [600.a] gonads serial.

TYPE-SPECIES.—Freyella sexradiata Perrier, 1885c.

DISCUSSION.—In addition to the characters listed above, the genus Freyastera is characterized by a very flat disc (on the same plane as the arms), plates of the disc frame are massive, and disc size is small. The adambulacral plates have a peculiar twisted look, and ambulacral plates are quite broad and solid. There is a conspicuous bare smooth plate in each interradius.

CHARACTERS.—Those peculiar to the genus Freyastera: 60.c, 170.6.

Characters Freyastera shares with Freyella: 160.a.

Characters in which Freyastera differs from Freyella: 30, 60, 70, 170, 400, 590, 600.

Freyastera benthophila (Sladen, 1889), new combination

FIGURE 18

Freyella benthophila Sladen, 1889:641–643, pl. 111: figs. 5–8.—Wood-Mason and Alcock, 1891:430.—Alcock, 1893a:121.—Fisher, 1919:538.—Madsen, 1950:184.—Cherbonnier and Sibuet, 1972:1356.—Sibuet, 1975: 292.—Korovchinsky and Galkin, 1984:1215 [key].

Freyellidea benthophila.—Fisher, 1917:429.

MERISTICS.—Arms = 6, R = 82–132 mm, r = 3–5.5 mm, R/r = 14.7–28.3/1, length of gonadal region = 10–29 mm, length of longest arm spine = 2 mm.

DIAGNOSIS.—[80.c] Madreporite small, nearly concealed by scale-like plates; [100.d] madreporite seemingly of two shell-like valves with a suture between; [110.e] abactinal disc plates scale-like, thin, delicate, fenestrate, irregular; [120.c] abactinal disc plates imbricate; [130.c] abactinal disc plates bearing 1–5 minute low tubercles overlaid with thin skin; [180.a] arms long, delicate, very attenuate, almost string-like; [205.b] gonadal inflation slight, length 10–29 mm, less than 1/8 length of arm; [210.e] abactinal arm plates scale-like; [240.c] abactinal arm plates bearing 1–3 minute tubercles, arms beyond gonadal region covered with delicate bare membrane; [250.a] terminal plates small relative to marginals; [260.a] terminal plates bifurcate; [270.a] terminals armed with 1 stout glassy spine on each half; [280.a] marginal plates small; [290.a] marginals irregular; [310.a] marginals bearing 1 long, glassy, acicular spine; [320.b] adambulacral plates very elongate; [330.b] adambulacrals spool-shaped; [340.c] furrow margin deeply indented; [390.a(b)] subambulacral spines moderately long, ridged, glassy, usually acicular but some with slightly expanded rough tip; [420.b] ambulacral plates very elongate; [430.d] head of ambulacral plates large, very elongate; [470.b] tubefeet moderately heavy; [480.c] mouth plates large; [490.b] paired mouth plates subhexagonal; [500] number of preoral spines per mouth plate 1 or 2; [530] number of lateral oral spines per mouth plate 0; [560] number of suboral spines per mouth plate 1; [570.a] suboral spines long, partly sheathed in membrane, acicular, stout, glassy; [580.b] suboral spines below center of mouth plate; [620.a,b,e] all spines except the small preoral spines covered with pedicellariae, which are also numerous on disc but none on gonadal portion of arms except on marginal and adambulacral spines, scattered pedicellariae on arms beyond gonadal region; [630.a] all pedicellariae small.

FIGURE 18.—Freyastera benthophila: a, abactinal view; b, actinal view.

COLOR.—Whitish, with pink tinge (in alcohol).

TYPE.—BM(NH) 90.5.7.1078 (holotype), Challenger Sta 289, mid-South Pacific, 39°41′S, 131°23′W, 4663 m.

MATERIAL EXAMINED.—Holotype; 4 specimens, USNM E29172, Golden Fleece Sta WS-N, off California, 39°31′N, 127°26′W, 4250 m.

DISTRIBUTION.—South Pacific, eastern Pacific off California, Bay of Bengal, mid-Atlantic (between Azores and Spain), Bay of Biscay; 4250–5000 m.

DISCUSSION.—This delicate and attractive species has a cosmopolitan deep-sea distribution. It is notable for the extremely long ambulacral plates and the partial membranous sheath clothing the spines.

CHARACTERS.—Those peculiar to F. benthophila: None.

Characters F. benthophila shares with other species of Freyastera:

F. |   mexicana: 100.d, 110.e, 180.a, 280.a, 470.b.
F. |   sexradiata: 100.d, 110.e,210.e,280.a,430.d, 480.c, 560.1, 630.a.
F. |   tuberculata: 120.c, 280.a, 290.a, 320.b, 420.b, 430.d, 470.b, 480.c, 530.0, 560.1.

Characters in which F. benthophila differs from other species of Freyastera:

F. |   mexicana: 80, 90, 130, 205, 210, 290, 330, 340, 430, 490, 500, 530, 560, 570, 580, 620.
F. |   sexradiata: 80, 130, 205, 240, 290, 330, 420, 470, 490, 500, 530, 620.
F. |   tuberculata: 80, 100, 110, 130, 205, 210, 330, 490, 580, 620.

Freyastera mexicana (A.H. Clark, 1939), new combination

FIGURE 19

Freyella mexicana A.H. Clark, 1939a:442.—Korovchinsky and Galkin, 1984:1213 [key].

MERISTICS.—Arms = 6, R = 110 mm (estimated), r = 5.5 mm, R/r = 20/1, length of gonadal region = 24 mm, length of longest arm spine = 1 mm.

DIAGNOSIS.—[80.a] Madreporite large; [90.b] madreporite lying in smooth, conical pit surrounded by many long spinelets; [100.d] madreporite with single S-shaped pore; [110.e] abactinal disc plates scale-like; [120.b,c] abactinal disc plates dense, slightly imbricate; [130.a] each abactinal disc plate bearing usually 1, sometimes 2 or 3 rough, glassy spinelets; [180.a] arms long, very attenuate; [205.a] gonadal portion of arms slightly inflated, set off abruptly from rest of arm; [210.f] abactinal arm plates flat, squarish, ending abruptly beyond gonadal region, distal portion of arms covered only by thin membrane; [240.a,c] abactinal arm plates bearing about 5 tubercles, each with a rough, glassy spinelet, membrane beyond gonadal region bearing scattered tiny pedicellariae; [280.a] marginal plates small; [290.b,e] marginal plates oval, flat, inconspicuous; [320.b] adambulacral plates longer than high; [330.a] adambulacrals cylindrical; [340.b] furrow margin slightly indented; [350] number of furrow spines 1; [380] number of subambulacral spines 1; [390.a] subambulacral spines large, glassy, mounted on conspicuous tubercle, 2 or 3 very delicate, setose small spines in diagonal line with larger spine, on inner distal edge of adambulacral; [420.a,b] ambulacral plates rather low proximally, quite high distally; [430.a] head of ambulacral plates more or less cylindrical, but not larger than base, waist of plates quite broad, the whole plate looking like a stout anvil: [470.b] tubefeet rather heavy: [490.c] mouth plates triangular, well separated from adjoining mouth plates, about as broad as long, with proximal lateral boss or extension; [500] number of preoral spines per mouth plate 3 or 4, moderately long; [510.b] preoral spines blunt; [520.a] preoral spines behind adoral margin; [530] number of lateral oral spines per mouth plate 2 or 3; [540.a] lateral oral spines small, very delicate, setose; [550.a,c] one lateral oral spine proximal, one or two distal; [560] number of suboral spines per mouth plate 2, large, stout; [570.c] suboral spines blunt; [580.a] suboral spines above center of mouth plate; [620.a,b] pedicellariae abundant on disc, arms, and all spines, those on spines quite large; [630.a,b] both large and small pedicellariae present.

FIGURE 19.—Freyastera mexicana: a, abactinal view; b, actinal view.

COLOR.—Unknown.

TYPE.—USNM E5602 (holotype); Albatross Sta 2379, Gulf of Mexico, 28°N, 87°42′W, 2683 m.

MATERIAL EXAMINED.—Holotype.

DISTRIBUTION.—Known only from type-locality.

DISCUSSION.—The very small, setose spines on the inner distal edge of the adambulacral plates might be thought to be furrow spines, but they sit not on the furrow face of the plate, but on the outer face, in a diagonal line with the large subambulacral spine and so should be considered subambulacral spines.

CHARACTERS.—Those peculiar to F. mexicana: 90.b, 570.c.

Characters F. mexicana shares with other species of Freyastera:

F. |   sexradiata: 100.d, 110.e, 130.a, 205.a, 280.a, 330.a, 500.3–4, 540.a.
F. |   tuberculata: 80.a, 130.a, 205.a, 210.f, 240.a,c, 280.a, 340.b, 380.1, 390.a, 470.b, 580.a, 620.a,b, 630.a,b.

Characters in which F. mexicana differs from other species of Freyastera:

F. |   sexradiata: 80, 90, 210, 290, 390, 430, 470, 490, 560, 580, 620, 630.
F. |   tuberculata: 90, 100, 110, 290, 330, 350, 430, 500, 530, 560.

Freyastera sexradiata (Perrier, 1885), new combination

FIGURE 20

Freyella sexradiata Perrier, 1885c:6; 1894:89–90, pl. 3: fig. 2.—Koehler, 1906[1907]:145; 1909a:129–130, pl. 23: fig. 9; 1921:2.—Grieg, 1927a:20, fig. 10.—Mortensen, 1927:129.—Gage et al., 1983:285.—Korovchinsky and Galkin, 1984:1213 [key].

FIGURE 20.—Freyastera sexradiata: a, abactinal view; b, actinal view (from Perrier, 1894).

MERISTICS.—Arms = 6(5), R = 100+ mm (estimated), r = 5–8 mm, R/r = 12.5–17/1, length of gonadal region = 30 mm, length of longest arm spine = 3 mm.

DIAGNOSIS.—[80.b] Size of madreporite moderate; [100.d] madreporite oval, with single S-shaped pore; [110.e] abactinal disc plates scale-like; [130.a] abactinal disc plates each bearing 1(2) glassy aciculate spinelet; [205.a] gonadal region of arm slightly inflated; [210.e] abactinal arm plates scale-like; [240.a] abactinal arm plates bearing single glassy spinelet; [280.a] marginal plates small; [290.c] form of marginals a small tubercle; [300.b] marginals occurring about every third adambulacral; [310.a] marginals each bearing a moderate-sized spine; [330.a] adambulacral plates cylindrical; [390.b] subambulacral spines moderately long, frequently capitate, spine on actinal face of adambulacral, plus a tiny secondary spine near distal prolongation of adambulacral; [420.a] ambulacral plates large, heavy; [430.d] head of ambulacral plates elongate; [470.a] tubefeet rather delicate; [480.c] mouth plates large; [490.d] mouth plates slightly longer than broad; [500] number of preoral spines per mouth plate 3 or 4; [530] number of lateral oral spines per mouth plate 1 or 2, tiny; [540.a] lateral oral spines acute; [550.c] one lateral oral spine on distal lateral prolongation of mouth plate; [560] number of suboral spines per mouth plate 1, moderately large; [580.b] suboral spines below center of mouth plate; [620.f] pedicellariae occurring only on spines, fewer than normal in most Brisingida; [630.a] all pedicellariae small.

COLOR.—Unknown.

TYPE.—Paris Museum 4598 (holotype); Talisman Sta 134, 42°19′N, 23°36′W, 4060 m.

MATERIAL EXAMINED.—Holotype.

DISTRIBUTION.—Azores, Portugal, Bay of Biscay, 38° to 45°N, 9° to 23°W; 4020–4700 m.

DISCUSSION.—This extremely delicate species has never been collected intact, making it impossible to accurately estimate the length of the arms; obviously, they must have been longer than 100 mm.

CHARACTERS.—Those peculiar to F. sexradiata: 620.f.

Characters F. sexradiata shares with other species of Freyastera:

F.
  tuberculata: 130.a, 205.a, 280.a, 430.d, 480.c, 560.1.

Characters in which F. sexradiata differs from other species of Freyastera:

F.
  tuberculata: 80, 100, 110, 210, 290, 330, 390, 420, 470, 490, 500, 530, 580, 620, 630.

Freyastera tuberculata (Sladen, 1889), new combination

FIGURE 21

Freyella tuberculata Sladen, 1889:638–640, pl. 117: figs. 1–3.—Alcock, 1893a:121.—Korovchinsky and Galkin, 1984:1214[key].

Freyellidea tuberculata.—Fisher, 1917:429.—H.L. Clark, 1920:113.

MERISTICS.—Arms = 6, R = 240 mm, r = 6 mm, R/r = 40/1, length of gonadal region = 30 mm, length of longest arm spine = 7 mm.

DIAGNOSIS.—[80.a] Madreporite huge; [100.c] madreporite subtubercular; [110.f] abactinal disc plates flat, hexagonal; [120.c] abactinal disc plates imbricate; [130.a] abactinal disc plates each armed with a short, sharp spinelet sheathed in membrane bearing many small pedicellariae; numerous larger pedicellariae on disc; [205.a] gonadal region of arms not highly inflated; [210.f] gonadal region of arms covered abactinally with very regular mosaic of squarish, thin plates, larger than those of disc, beyond gonadal region, plates scattered, fewer and thinner, but occurring all the way out the arm; [240.a,c] abactinal arm plates bearing a single tubercle with a glassy spinelet; [280.a] marginal plates small; [290.a] marginals thin, round, reduced; [310.a] marginals bearing single delicate acicular spine; [320.b] adambulacral plates longer than high; [330.f] adambulacral plates dumbbell-shaped, well separated from adjoining plates by muscle tissue in which is sometimes embedded a few small irregular plates or calcite deposits; [340.b] furrow margin slightly indented; [350] furrow spines absent; [380] number of subambulacral spines 1; [390.a] subambulacral spines long, delicate, glassy, borne on tubercle on actinal face of plate; [420.b] ambulacral plates rather small, elongate; [430.d] head of ambulacrals long, cylindrical; [470.b] tubefeet rather heavy; [480.c] mouth plates large; [490.c,f] mouth plates wedge-shaped, with median keel; [500] number of preoral spines per mouth plate 1; [530] lateral oral spines absent [560] number of suboral spines per mouth plate 1; [580.a] suboral spines above center of mouth plate; [620.a,b] all spines covered with delicate membrane bearing many pedicellariae, disc and arm plates bearing larger scattered pedicellariae; [630.a,b] both large and small pedicellariae present.

FIGURE 21.—Freyastera tuberculata: a, abactinal view of disc; b, actinal view of disc; c, abactinal view of arm section; d, actinal view of arm section.

COLOR.—Grayish white, with light pink tinge to gonadal region (in alcohol).

TYPES.—BM(NH) 90.5.7.1–76–1077 (2 syntypes); Challenger Sta 89, between Canary Islands and Cape Verde Islands, 22°18′N, 22°02′W, 4389 m; Sta 346, between W coast of Africa and Ascension Island, 2°42′S, 14°41′W, 4298 m.

MATERIAL EXAMINED.—Syntypes; 1 specimen, USNM E931, Albatross Sta 4742, Eastern Tropical Pacific, 0°03′N, 117°15′W, 4243 m; 1 specimen, USNM E20797, Pillsbury Sta 1372, mid-Atlantic, 23°14′N, 64°49′W, 5618 m.

DISTRIBUTION.—Apparently cosmopolitan; Alcock (1893a) also reported it from the Indian Ocean.

DISCUSSION.—This species is quite close to Freyastera sexradiata, but differs in having fewer mouth spines and more pedicellariae.

CHARACTERS.—Those peculiar to F. tuberculata: None.

Genus Freyella Perrier, 1885

Freyella Perrier, 1885c:5.—Sladen, 1889:618.—Alcock, 1893a:121.—Verrill, 1894:283.—Perrier, 1894:81.—Verrill, 1895:212.—Ludwig, 1905a:267.—Fisher, 1905:319.—Koehler, 1907:3; 1909d:626.—Fisher, 1916b:35.—A.H. Clark, 1916:51.—Fisher, 1918:104.—1919:538; 1928:24.—Grieg, 1932:30.—A.H. Clark, 1939a:442.—H.L. Clark, 1941:65.—Hayashi, 1943:169.—Madsen, 1956:29.—Cherbonnier and Sibuet, 1972:1353.—Sibuet, 1975:281.—Korovchinsky, 1976:1188.—Korovchinsky and Galkin, 1984:1205–1215.

Freyellidea Fisher, 1917:424; 1919:538.—H.L. Clark, 1920:108.—Döderlein, 1927:293.

Astrocles Fisher, 1917:426, 430; 1919:504; 1928:29.

DIAGNOSIS.—[30.b] Disc slightly raised above plane of arms; [50.b] interradial arcs gently curved; [60.b] flesh thin, opaque; [70.a,b] anal opening microscopic or absent; [140.a] a large, bare, smooth interradial plate at each interradial angle; [160.a] papulae absent; [170] number of arms 8–14; [190.b] abactinal surface of proximal arms more or less a continuation with that of disc, arms not constricted at base; [220.b] abactinal arm plates tessellate; [400.a] proximal adambulacrals united interradially; [590.b] 2 pairs of gonads per arm; [600.b] gonads paired.

TYPE-SPECIES.—Freyella spinosa Perrier (1885c:5), by original designation (= Freyella elegans (Verrill, 1884)).

DISCUSSION.—Fisher (1917) attempted to divide the genus Freyella into two groups, Freyella and Freyellidea, but later (1919) decided, quite rightly, that the basis for division was of less than generic importance. The genus Astrocles (Fisher, 1917) differs from Freyella only in having the abactinal arm plates arranged in bands across the arms. As Fisher himself pointed out (1928:29), these plates are more like Freyella plates than the plates that form the costae in Brisinga and related genera. As most of the species of Freyella have the abactinal arm plates beyond the center of the gonadal region also arranged in bands, at least in larger specimens, this can hardly be considered a generic character, and Astrocles is thus a junior synonym of Freyella, and the type and only known species, Astrocles actinodetus, from off British Columbia, is closely related to Freyella microspina.

CHARACTERS.—Those peculiar to Freyella: None.

Freyella elegans (Verrill, 1884)

FIGURE 22

Brisinga elegans Verrill, 1884:382 [non Brisinga elegans Perrier, 1885 (= Novodina homonyma Downey, new name).]

Freyella spinosa Perrier, 1885c:5; 1894:85–89, pl. 7.—Koehler, 1907:6.—Mortensen, 1927:129.—Sibuet, 1975:281.—Gage et al., 1983:285.—Korovchinsky and Galkin, 1984:1213 [key].

Freyella bracteata Sladen, 1889:629–632, pl. 114: figs 1–4.

Freyella elegans.—Verrill, 1894:283; 1895:212.

Freyella spinosa Perrier, 1885c:5; 1894:85–89, pl. 7.—Koehler, 1907:6.—Mortensen, 1927:129.—Sibuet, 1975:281.—Gage et al., 1983:285.—Korovchinsky and Galkin, 1984:1213 [key].

Freyella abyssicola.—A.H. Clark, 1949:375.

Freyella laubieri Cherbonnier and Sibuet, 1972:1353, unnumbered figure.

FIGURE 22.—Freyella elegans: a, abactinal view; b, actinal view.

MERISTICS.—Arms = 9–14, R = 135–191 mm, r = 5–15 mm, R/r = 12–27/1, length of gonadal region = 21–44 mm, length of longest arm spine = 7–9 mm.

DIAGNOSIS.—[80.a,b,c] Madreporite inconspicuous to moderately large; [100.a] madreporite channeled; [110.e,f] abactinal disc plates thin, flat; scale-like; [120.c] abactinal disc plates imbricate; [130.a] abactinal disc plates bearing 2–5 short, sharp, usually divergent spinelets; [170] number of arms 9–14; [205.a,c] gonadal region of arms 21–44 mm long, slightly to highly inflated; flated; [210.f] abactinal arm plates flat, diamond-shaped; [220.b] abactinal arm plates tessellate, closely abutting adjoining plates; [240.a] abactinal arm costae bearing single irregular transverse row of short, sharp spinelets; [250.a] terminal plates tiny; [260.a] terminal plates bifurcate; [270.a] terminal plates armed with 1 or 2 acute spines; [280.a] marginals small; [290.c] marginals tuberculate; [310.a] marginals bearing 1 long stout acicular spine; [320.a] adambulacrals higher than long, with inner distal prolongation; [330.a] adambulacral plates cylindrical; [340.a] furrow margin straight; [350] number of furrow spines 0–2; [360.a] furrow spines short, sharp; [370.b] furrow spines distal; [380] number of subambulacral spines 1; [390.a,b,c] subambulacral spines large, stout, aciculate, capitate, or truncate (usually capitate or truncate in gonadal region, aciculate beyond); [420.a] ambulacral plates rather high; [430.e] ambulacral plates dumbbell-shaped, head large, elongate, spool-shaped; [470.c] tubefeet neither particularly delicate or particularly heavy; [480.a] mouth plates rather small; [490.c] mouth plates slightly longer than broad; [500] number of preoral spines per mouth plate 2 or 3; [510.a,b] preoral spines short, blunt or acicular, divergent; [520.a,b] preoral spines adoral or lateral; [540.a] lateral oral spines short, acicular; [550.c] lateral oral spines at distal angle of mouth plates; [560] number of suboral spines per mouth plate 1–3; [570.a] suboral spines long, stout, acicular; [580.a] suboral spines above center of mouth plate; [620.a,b,e] pedicellariae scattered on both surfaces of disc and arms; [630.a] all pedicellariae small.

COLOR.—Unknown.

TYPES.—Syntypes: USNM 6740, 11 specimens, Albatross Sta 2105, off Virginia, 2551 m; USNM 8076, 5 specimens, Albatross Sta 2229, off Maryland, 2602 m; USNM 9015, 10 specimens, Albatross Sta 2084, off George's Bank, 2359 m; USNM 9016, Albatross Sta 2076, off George's Bank, 1657 m; USNM 9065, 2 specimens, Albatross Sta 2035, off Nantucket Shoals, 2491 m; USNM 18305, 3 arm fragments, Albatross Sta 2077, off Martha's Vineyard, 2295 m; USNM 18455, 14 specimens, Albatross Sta 2105, off Virginia, 2551 m.

MATERIAL EXAMINED.—Syntypes; 1 specimen, USNM 7819, Albatross Sta 2209, S of Block Island, 1829 m; 1 specimen, USNM 7854, Albatross Sta 2211, Cape Hatteras to Nantucket, 1946 m; 1 specimen, USNM 8135, Albatross Sta 2230, off Delaware, 2136 m; 1 specimen, USNM 11257, Albatross Sta 2534, SE of George's Bank, 2257 m; 4 specimens, USNM 11258, Albatross Sta 2564, SW of Martha's Vineyard, 2542 m; 6 specimens, USNM 11259, Albatross Sta 2562, SW of Martha's Vineyard, 2622 m; 8 specimens, USNM 15560, Albatross Sta 2725, off Chesapeake Bay, 2513 m; 1 specimen, USNM 15561, Albatross Sta 2727, off Chesapeake Bay, 2266 m; 1 arm fragment, USNM 18306, Albatross Sta 2192, S of Nantucket, 1939 m; 2 specimens, USNM 38452, Albatross Sta 2229, Cape Hatteras to Nantucket, 2602 m; 32 specimens, USNM 38862, Albatross Sta 2105, between Cape Hatteras and Nantucket, 2551 m; 3 specimens, USNM 38926, Albatross Sta 2229, Cape Hatteras to Nantueket, 2602 m; 4 specimens, USNM E12438, 34°45′N, 75°11′W, 2450 m; 1 arm fragment, USNM E15979, “Knorr” Sta 326, Hudson Canyon; 5 specimens, USNM E15982, “Knorr” Sta 325, Hudson Canyon, 39°13′N, 71°53′W, 1919 m; 2 specimens, USNM E20944, Pillsbury Sta 292, Gulf of Guinea, 0°12′N, 5°11′E, 3587 m; 1 specimen, USNM E20945, Pillsbury Sta 34, Gulf of Guinea, 3°53′N, 2°33′W, 1986 m; 1 specimen, USNM E13828, Chain Sta 249; 2 specimens, USNM 6740, Albatross Sta 2105, off Virginia, 2551 m; 5specimens, USNM 12068, Albatross Sta 2571, SE of George's Bank, 2480 m (the last two lots were originally labelled “Brisinga spinulosa” but this name was never published; Verrill evidently meant it for a replacement name); type of Freyella spinosa Perrier, Paris Museum, about 19°N, 20°W, 2320-4060 m; type of Freyella aspera Verrill, USNM 6301, Albatross Sta 2097, off Martha's Vineyard, 3506 m; type of Freyella bracteata Sladen, Challenger Sta 46, E of New Jersey, 40° 17′N, 66°48′W, 2469 m; type of Freyella laubieri Cherbonnier and Sibuet, Jean Charcot Sta 17, 52°45′N, 41° 12′W, 4340 m.

DISTRIBUTION.—Greenland to Angola (east) and North Carolina (west); 1600-4500 m.

DISCUSSION.—This variable species may have a worldwide distribution in deep water. The size of the gonadal inflation on the arms is dependent on the state of development of the gonads. The most variable features are the preoral mouth spines, the proximal adambulacral spines, and the distribution and abundance of pedicellariae. The oral mouth spines and the proximal adambulacral furrow spines may be straight and acicular, or broad, truncate, and more or less distorted; this seems to be related to distribution, not growth, as is the shape of the adambulacral spines. Possibly Freyella mutabilis Korovchinsky from the Sub-Antarctic (4664-6070 m), F. dimorpha Sladen from Torres Strait (2560 m), F. pennata Sladen from Japan (3429 m), and others also belong to this species. Certainly all the Atlantic species heretofore described that have two or three short preoral spines, one short furrow spine, one long, stout subambulacral spine, and abactinal disc and arm plates as described above belong to the same species, for which Verrill's name elegans has priority.

CHARACTERS.—Those peculiar to F. elegans: 470.C.

Characters F. elegans shares with other species of Freyella:

F. |   microspina: 130.a, 240.a, 280.a, 290.C, 340.a, 390.a,b,c, 430.e, 490.C, 620.a,b,e.
F. |   recta: 100.a, 130.a, 220.b, 240.a, 280.a, 320.a, 330.a, 380.1, 480.a, 540.a, 570.a.

Characters in which F. elegans differs from other species of Freyella:

F. |   microspina: 100, 110, 120, 210, 320, 330, 470, 480, 540, 550, 570, 580, 630.
F. |   recta: 110, 120, 170, 210, 290, 340, 470, 550, 580, 620.

Freyella microspina Verrill, 1894

FIGURE 23

Freyella microspina Verrill, 1894:285.—Korovchinsky and Galkin. 1984:1214 [key].

Freyella trispinosa H.L. Clark, 1941:65–66.—Korovchinsky and Galkin, 1984:1214 [key].

MERISTICS.—Arms = 10–13, R = 95–140 mm (estimated), r = 5–11.5 mm, R/r = 12–19/1, length of gonadal region = 20–33 mm, length of longest arm spine = 7 mm.

DIAGNOSIS.—[80.a] Madreporite large; [100.b] madreporite rough, irregular; [110.b] abactinal disc plates small, tumid, roundish; [120.a] abactinal disc plates scattered in membrane; [130.a] abactinal disc plates bearing central group of 1–5 short, sharp spinelets; [170] number of arms 10–13; [205.a] gonadal region of arms 20–33 mm long, rather sturdy; [210.b] abactinal arm plates tumid; [220.b,c] abactinal arm plates in irregular mosaic; [240.a] abactinal arm plates bearing roughtly 2 rows of short, sharp spinelets; [280.a] marginals small; [290.c] marginals conical; [310.a] marginals bearing one large acicular spine; [320.b] adambulacral plates slightly longer than high, with long truncate prolongation at inner distal edge; [330.e] adambulacral plates knobbed; [340.a] furrow margin straight; [350] furrow spines usually absent, sometimes one short spinelet on distal prolongation; [360.a,b] furrow spines acute or clavate; [370.a,b] one furrow spine proximal, one distal; [380] number of subambulacral spines 1 or 2; [390.a,b,c] subambulacral spines large, stout, truncate or capitate in gonadal region, aciculate beyond; [420.b] ambulacral plates small, delicate; [430.e] ambulacral plates dumbbell-shaped; [470.a] tubefeet rather delicate; [480.c] mouth plates moderately large; [490.c] mouth plates longer than wide, wider at distal than at proximal end; [500] number of preoral spines per mouth plate 3; [510.a] preoral spines moderately long, acicular; [520.b] preoral spines directed laterally, meeting across ambulacral groove, so appearance is of 3 more or less continuous rings of calcite around the actinostome; [530] number of lateral oral spines per mouth plate 1; [540.c,d] lateral oral spines short, flat, truncate, broadly expanded at tip; [550.b] lateral oral spines lying beneath preoral spines and meeting the lateral orals of adjoining mouth plates; [570.b] suboral spines large, usually capitate; [580.b] suboral spines below center of mouth plate; [620.a,b,e] pedicellariae on both surfaces of disc and arms; [630.b] pedicellariae rather large.

FIGURE 23.—Freyella microspina: a, abactinal view; b, actinal view.

COLOR.—Unknown.

TYPE.—USNM 7821 (arm fragments) (holotype), Albatross Sta 2220, off Nantucket, 39°43′N, 69°23′W, 1928 m.

MATERIAL EXAMINED.—Holotype; 1 specimen, USNM 7805, Albatross Sta 2192, S of Nantucket, 1939 m; 1 specimen, USNM E20799, Pillsbury Sta 681, off Surinam, 8°11.5′N, 56°12′W, 2734 m; type of Freyella trispinosa, MCZ 3825, Atlantis Sta 2969A, off Bahia de Guantanamo, Cuba, 1847 m.

DISTRIBUTION.—Off Nantucket; southern coast of Cuba; Surinam; 1847–2734 m.

DISCUSSION.—Although closely related to Freyella elegans, the disc and arm plates of this species are generally thicker and more tumid, the preoral spines are entirely lateral in position, and the small, single furrow spine is sometimes lacking entirely. Freyella trispinosa differs only in the shape of the preoral spines (thinner and more truncate), but this can scarcely be considered a specific difference.

CHARACTERS.—Those peculiar to F. microspina: 330.e, 540.c,d.

Characters F. microspina shares with other species of Freyella:

F.
  recta: 120.a, 130.a, 205.a, 240.a, 280.a, 360.a,b, 510.a, 580.b.

Characters in which F. microspina differs from other species of Freyella:

F.
  recta: 80, 100, 110, 170, 210, 290, 320, 330, 340, 470, 480, 500, 520, 540, 550, 570, 620.

Freyella recta Koehler, 1907

FIGURE 24

Freyella recta Koehler, 1907a:3; 1909:127–129, pl. 21: fig. 7, pl. 22: fig. 4, pl. 23: figs. 3–5.—Korovchinsky and Galkin, 1984:1214 [key].

MERISTICS.—Arms = 8, R = 150+ mm, r = 9 mm, R/r = 17/1, length of gonadal region = 35 mm, length of longest arm spine = 3 mm.

DIAGNOSIS.—[80.c] Madreporite very small, raised; [100.a] madreporite channeled; [110.a] abactinal disc plates metapaxillar, irregularly polygonal; [120.a] abactinal disc plates small, embedded in membrane; [130.a] abactinal disc plates bearing 1–3 short, blunt, rough spinelets; [170] number of arms 8; [205.a] gonadal region of arms 35 mm long, moderately inflated; [210.c,f] abactinal arm plates irregularly polygonal, metapaxillar, larger than those of disc on proximal part of arm, bare membrane beyond gonadal region; [220.b] abactinal arm plates tessellate, forming irregular mosaic; [240.a] abactinal arm plates bearing 4 or 5 short, fine, conical spinelets; [280.a] marginal plates small; [290.b] marginals more or less oval; [310.a] marginals bearing one short acicular spine; [320.a] adambulacral plates higher than long; [330.a] adambulacral plates cylindrical; [340.b] furrow margin slightly indented; [350] number of furrow spines 1; [360.a,b] furrow spines acute or clavate; [380] number of subambulacral spines 1; [390.a,b] subambulacral spines large, first 6 or 7 capitate, rest aciculate; [470.b] tubefeet rather heavy; [480.a] mouth plates rather small; [500] number of preoral spines per mouth plate 2, one tiny, delicate, the other large, stout; [510.a] preoral spines short, sharp, straight; [520.a] preoral spines adoral; [540.a] lateral oral spines acute; [550.a] lateral oral spines proximal; [570.a] suboral spines large, acicular; [580.b] suboral spines below center of mouth plate; [620.a,b,d] pedicellariae in patches on distal membranous area of arms, all spine with membranous sheath covered with pedicellariae (tips of spines exposed).

FIGURE 24.—Freyella recta: a, abactinal view; b, actinal view. (From Koehler, 1909a.)

COLOR.—Unknown.

TYPE.—Musee Oceanographique de Monaco (holotype); Princesse Alice Sta 2111, mid-Atlantic ridge, between Azores and Virgin Islands, 31°45′N, 42°42′W, 3465 m.

MATERIAL EXAMINED.—None.

DISTRIBUTION.—Known only from type-locality.

DISCUSSION.—As I was unable to examine the type, I cannot say with certainty that this species may be another variety of Freyella elegans. Without further material or access to the type, I consider the validity of this species in doubt.

CHARACTERS.—Those unique to Freyella recta: 620.a,b,d.

Appendix

CHARACTERS OF THE BRISINGIDA

  10. | Form:
| a. | Ophiuroid-like
  20. | Disc size:
| a. | Small
  30. | Disc thickness:
| a. | Thick, raised well above plane of arms
| b. | Intermediate, raised slightly above plane of arms
| c. | Flat, in same plane as arms
  40. | Disc shape:
| a. | Circular
  50. | Interradial arcs:
| a. | Acute
| b. | Curved
  60. | Fleshiness:
| a. | Thick, dense, opaque
| b. | Moderate, thin, opaque
| c. | Slight, transparent, usually slimey
  70. | Anal opening:
| a. | Barely visible (microscopic)
| b. | Absent
| c. | Plainly visible
  80. | Size of madreporite:
| a. | Large
| b. | Moderate
| c. | Small
  90. | Position of madreporite:
| a. | Lateral (on edge of disc)
| b. | Near edge of disc
100. | Type of madreporite:
| a. | Channeled
| b. | Irregular
| c. | Subtubercular
| d. | Single pore
| e. | Meandering gyri
110. | Type of abactinal disc plates:
| a. | Metapaxillar
| b. | Tumid
| c. | Tuberculate
| d. | Papilliform
| e. | Scale-like
| f. | Flat
| g. | Lobed or cruciform
120. | Arrangement of abactinal disc plates:
| a. | Scattered in membrane
| b. | Dense in membrane
| c. | Imbricate
| d. | Few or none
130. | Armament of abactinal disc plates:
| a. | Spinelets
| b. | Spines
| c. | Tubercles
140. | Bare interradial plates:
| a. | Present
| b. | Absent
150. | Distribution of papulae:
| a. | On disc only
| b. | On disc and arms
160. | Number of papulae:
| a. | None
| b. | Scattered (single)
| c. | Exact number
170. | Number and/or length of arms
180. | Shape of arms:
| a. | Long, attenuate, R/r more than 6/1
| b. | Long, robust
190. | Attachment of arms to disc:
| a. | Arms deciduous, constricted at base
| b. | Abactinal surface of arms continuous with that of disc, not constricted at base
200. | Gonadal region differing from rest of arm:
| a. | Yes
205. | Size of gonadal region of arms:
| a. | Moderately inflated
| b. | Not inflated
| c. | Greatly inflated
210. | Type of abactinal arm plates:
| a. | Rod-like
| b. | Tumid
| c. | Metapaxillar
| d. | Irregular
| e. | Scale-like
| f. | Flat
| g. | Lobed or cruciform
220. | Arrangement of abactinal arm plates:
| a. | Imbricate
| b. | Abutting (tessellate)
| c. | Scattered
230. | Size of abactinal arm plates relative to marginals:
| a. | Small
240. | Armament of abactinal arm plates:
| a. | Spinelets
| b. | Spines
| c. | Tubercles
250. | Size of terminal plates relative to marginals:
| a. | Small
| b. | Large
260. | Shape of terminal plates:
| a. | Irregular or bifurcate or catsclaw
| b. | Squarish or shield-shaped
270. | Armament of terminal plates:
| a. | Spines
| b. | Spinelets
| c. | Bare
280. | Size of marginal plates relative to adambulacrals:
| a. | Small
| b. | Large
290. | Shape of marginals:
| a. | Irregular
| b. | Elongate
| c. | Tuberculate
| d. | Angular
| e. | Flat
| f. | Squarish or rectangular
| g. | Cruciform or lobate
| h. | Tumid
300. | Position of marginals relative to adambulacrals:
| a. | Corresponding
| b. | Not corresponding
310. | Armament of marginals:
| a. | Spines
| b. | Spinelets
320. | Size of adambulacrals relative to marginals:
| a. | Large (higher than long)
| b. | Small (longer than high)
330. | Shape of adambulacrals:
| a. | Cylindrical
| b. | Spool-shaped
| c. | Discoidal
| d. | Hourglass-shaped
| e. | Knobbed
| f. | Dumbbell-shaped
340. | Furrow margin:
| a. | Straight
| b. | Slightly indented
| c. | Deeply indented
350. | Number of furrow spines
360. | Form of furrow spines:
| a. | Acute
| b. | Clavate
370. | Position of furrow spines:
| a. | Proximal
| b. | Distal
| c. | Medial
380. | Number of subambulacral spines
390. | Form of subambulacral spines:
| a. | Acute
| b. | Capitate
| c. | Truncate
| d. | Furcate
400. | Proximal adambulacrals united interradially:
| a. | Yes
| b. | No
410. | Ambulacral groove:
| a. | Wide (broad, open, often petaloid)
420. | Size of ambulacral plates:
| a. | Large, heavy
| b. | Small, delicate
430. | Shape of ambulacral plates:
| a. | Saddle-shaped
| b. | Hourglass-shaped
| c. | Y-shaped
| d. | T-shaped
| e. | Dumbbell-shaped
| f. | Discoidal
| g. | I-shaped
440. | Spacing of ambulacral plates:
| a. | With heads abutting end to end
| b. | Crowded
450. | Number of rows of tubefeet 2
460. | Type of tubefeet:
| a. | Suckered
470. | Structure of tubefeet:
| a. | Delicate, slender, thin, translucent
| b. | Heavy, thick, opaque
| c. | Neither of the above
480. | Size of mouth plates relative to adambulacrals:
| a. | Small
| b. | Moderate
| c. | Large
490. | Shape of mouth plates:
| a. | T-shaped
| b. | Trapezoidal
| c. | Triangular
| d. | Cylindrical
| e. | Short, broad
| f. | Keeled
| g. | Hourglass-shaped
500. | Number of preoral spines
510. | Shape of preoral spines:
| a. | Acute
| b. | Blunt
520. | Position of preoral spines:
| a. | Adoral
| b. | Lateral
530. | Number of lateral oral spines
540. | Shape of lateral oral spines:
| a. | Acute
| b. | Blunt
| c. | Truncate
| d. | Flattened
550. | Position of lateral oral spines:
| a. | Proximal
| b. | Lateral
| c. | Distal
560. | Number of suboral spines
570. | Shape of suboral spines:
| a. | Acute
| b. | Capitate
| c. | Blunt
580. | Position of suboral spines relative to mouth plate:
| a. | Above center
| b. | Below center
| c. | Center
590. | Number of pairs of gonads per arm:
| a. | Many
| b. | 2
| c. | 1
600. | Position of gonads:
| a. | Serial
| b. | Paired
610. | Type of pedicellariae:
| a. | Crossed
620. | Distribution of pedicellariae:
| a. | Actinal and abactinal
| b. | Disc and arms
| c. | In rows or bands
| d. | In patches
| e. | Scattered
| f. | Only on spines
| g. | Arms only
630. | Size of pedicellariae:
| a. | Microscopic
| b. | Plainly visible

Literature Cited

Alcock, A.

1893a. Natural History Notes from H.M. Indian Marine Survey Steamer Investigator, Commander Oldham, R.N., Commanding, Series 2, No. 7: An Account of the Collection of Deep-sea Asteroidea. Annals and Magazine of Natural History, 11(6):73–121, plates 4–6.

1893b. Natural History Notes from H.M. Indian Marine Survey Steamer Investigator, Commander Oldham, R.N., Commanding, Series 2, No. 9: An Account of the Deep-sea Collection Made during the Season of 1892–1893. Journal of the Asiatic Society of Bengal, 62:169–184, plates 8–9.

Asbjørnsen, P.C.

1856.   Brisinga endecacnemos. In Sars, Koren, and Danielssen. Fauna Littoralis Norvegiae, 2:101, 12 plates. Bergen.

Bell, F.J.

1889.   Report of a Deep-sea Trawling Cruise off the S.W. Coast of Ireland, under the Direction of Reverend W. Spotswood Green: Echinodermata. Annals and Magazine of Natural History, 4:432–445, plates 18, 19.

1892a. On the Echinoderms Collected by the Steamer Fingal in 1890, and by the Steamer Harlequin in 1891, off the West Coast of Ireland. Scientific Proceedings of the Royal Society of Dublin, new series, 7:520–529, plates 23, 24.

1892 [1893].   Catalogue of the British Echinoderms in the British Museum (Natural History). xviii + 202 pages, 16 plates. London.

1893.   On the Names or Existence of Three Exotic Starfishes. Annals and Magazine of Natural History, 6(12):25–29.

Bernasconi, I.

1964.   Asteroideos Argentinos: Claves para los ordenes, familias, subfamilias y generos. Physis, Buenos Aires, 24:241–277, 3 plates.

Boone, L.

1933.   Scientific Results of Cruises of the Yachts Eagle and Ara, 1921–1928, William K. Vanderbilt Commanding. Bulletin of the Vanderbilt Marine Museum (Huntington), 4:68–164, plates 25–102.

Carus, J.V.

1884 [1885].   Prodromus Faunae Mediterraneae sive descriptio Animalium Maris Mediterranei incolarum, & etc. 93 pages. Stuttgart.

Cherbonnier, G., and M. Sibuet

1972.   Resultats scientifiques de la campagne Noratlante: Asteroides et Ophiurides. Bulletin Musee Nationale d'Histoire Naturelle de Paris (Zoologie), 76:1333–1394, 7 figures, 5 tables, 11 maps.

Clark, A.H.

1916.   Six New Starfishes from the Gulf of California and Adjacent Waters. Proceedings of the Biological Society of Washington, 29:51–62.

1934.   A New Starfish from Puerto Rico. Smithsonian Miscellaneous Collections, 91(14):1–3, plate 1.

1939a. Echinoderms of the Smithsonian-Hartford Expedition, 1927, with other West Indian Records. Proceedings of the United States National Museum, 86:441–454, plates 53–54.

1939b. Echinoderms (Other than Holothurians) Collected on the Presidential Cruise of 1938. Smithsonian Miscellaneous Collections, 98(11):1–18, plates 1–5.

1949.   Echinoderms from the Mid-Atlantic Dredged by the Atlantis in the Summer of 1948. Journal of the Washington Academy of Science, 39:371–377.

Clark, A.M.

1962.   Starfishes and Their Relations. 119 pages, 14 plates, 30 figures. London: British Museum (Natural History).

Clark, A.M., and J. Courtman-Stock

1976.   The Echinoderms of South Africa. 277 pages, 276 figures, map. London: British Museum (Natural History).

Clark, H.L.

1920.   Eastern Tropical Pacific Asteroidea. Memoirs of the Museum of Comparative Zoology at Harvard, 39:73–113, plates 1–6.

1926.   Echinoderms from the South African Fisheries and Marine Biological Survey, Part III: Sea-stars (Asteroidea). Report of the Fisheries and Marine Biological Survey, Cape Town, 4(1925):1–33, plates 1–7.

1941.   Reports on the Scientific Results of the Atlantis Expeditions to the West Indies under the Joint Auspices of the University of Havana and Harvard University: The Echinoderms (Other than Holothurians). Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey,” 15:1–154, plates 1–10.

Danielssen, D.C., and J. Koren

1884.   The Norwegian North Atlantic Expedition 1876– 1878, Zoology: Asteroidea. 118 pages, 15 plates, map. Christiana: Grondahl and Son.

Dartnell, A.J., D.L. Pawson, E.C. Pope, and B.J. Smith

1969.   Replacement Name for the Preoccupied Genus Name Odinia Perrier, 1885 (Echinodermata: Asteroidea). Proceedings of the Linnean Society of New South Wales, 93:211.

de Loriol, P.

1883.   Catalogue raisonne des Echinodermes recueillis par M.V. Robillard à l'isle Maurice. Memoires de la Société Physiologique de Geneve, 28(8): 64 pages, 6 plates.

Djakanov, A.M.

1968.   Sea Stars (Asteroids) of the USSR Seas. Izdatel'stvo Akademii Nauk SSSR Moskva-Leningrad, 34:1–203, figures 1–212. [Translation from the Russian: Keys to the Fauna of the USSR, originally published in 1950, same journal.]

Döderlein, L.

1927.   Die Seesterne der Deutschen Sudpolar-Expedition 1901–1903. In Deutsche Sudpolar Expedition 1901–1903, series 19 (Zoology), 11:289–301, plates 11–14. Berlin.

Downey, M.E.

1972.   Midgardia xandaros New Genus, New Species, a Large Brisingid Starfish from the Gulf of Mexico. Proceedings of the Biological Society of Washington, 84(48):421–426, 2 figures.

1973.   Starfishes from the Caribbean and the Gulf of Mexico. Smithsonian Contributions to Zoology, 126:1–158, 48 plates, 2 figures.

Farran, G.P.

1913.   The Deep-water Asteroidea, Ophiuroidea and Echinoidea of the West Coast of Ireland. Fisheries of Ireland Scientific Investigations, 6(1912): 1–66, plates 1, 2.

Fisher, W.K.

1905.   New Starfishes from Deep Water off California and Alaska. Bulletin of the United States Bureau of Fisheries, 24:291–320.

1906.   The Starfishes of the Hawaiian Islands. Bulletin of the United States Fish Commission, 23:987–1130, plates 1–49.

1916a. Notes on the Systematic Position of Certain Genera and Higher Groups of Starfishes. Proceedings of the Biological Society of Washington, 29:1–6.

1916b. New East Indian Starfishes. Proceedings of the Biological Society of Washington, 29:27–36.

1917.   New Genera and Species of Brisingidae. Annals and Magazine of Natural History (London), series 8, 20:418–431.

1918.   Notes on Asteroidea. Annals and Magazine of Natural History (London), series 1, 9:103–111, plate 13.

1919.   Contributions to the Biology of the Philippine Archipelago and Adjacent Regions: Starfishes of the Philippine Seas and Adjacent Waters. Bulletin of the United States National Museum, 100(3):1–712, plates 1–156.

1928.   Asteroidea of the North Pacific and Adjacent Waters, Part 2: Forcipulata (Part). Bulletin of the United States National Museum, 76:1–245, plates 1–81.

1940.   Asteroidea. Discovery Reports (Cambridge), 20:69–306, figures A–M, plates 1–23.

Gage, J.D., M. Pearson, A.M. Clark, G.L. Paterson, and P.A. Tyler

1983.   Echinoderms of Rockall Trough and Adjacent Areas, I: Crinoidea, Asteroidea and Ophiuroidea. Bulletin of the British Museum (Natural History), Zoology, 45(5):263–308, 3 figires, 1 table.

Grieg, J. A.

1927a. Evertebrater fra bankerne ved Spitzbergen indsamlet av Tovik og Armauer Hansen somrene 1925 og 1926 (Fiskenaering og bundfauna). Bergens Museum Aarsbok, 1926(5):l–28.

1927b. Echinoderms from the West Coast of Norway. Nyt Magazin for Naturvidenskaberne, 65:127–136.

1928.   The Folden Fjord: Echinodermata. Tromsø Museum Skrifter, 1(7): 1–12.

1932.   Echinodermata. In Report of the Scientific Research of the “Michael Sars” North Atlantic Expedition, 1910, 3(2):l–47, 5 plates. Bergen.

Hayashi, R.

1943.   Contributions to the Classification of the Sea-stars of Japan, II: Forcipulata, with the Note on the Relationships between the Skeletal Structure and Respiratory Organs of the Sea-stars. Journal of the Faculty of Science of Hokkaido University (Zoology) 8:133–277, figures 1–115, plates 1–11.

John, D.D., and A.M. Clark

1954.   The “Rosaura” Expedition, 3: The Echinodermata. Bulletin of the British Museum (Natural History). Zoology, 2:139–162, figures 1–12, plate 6.

Koehler, R.

1895a. Notes Echinologiques. Revue Biologique Nord France, 7:317–342, plate 9.

1895b. Catalogue raisonne des Echinodermes recueillis par M. Korotnov aux iles de la Sonde. Memoires de la Société Zoologique de France, 8:374–423, plate 9.

1896a. Dragages profonds executes a bord du Caudan dans le Golfe de Gascogne (Aout–Septembre, 1895): Rapport preliminaire sur les Echinodermes. Revue Biologique Nord France, 7:439–496.

1896b. Resultats scientifiques de la campagne du Caudan dans le Golfe de Gascogne (Aout–Septembre, 1895). Echinodermes. Annales de l'Universitaire de Lyon, 26:33–127, plates 1–4.

1906 [1907].   Expedition Antarctique Francaise (1903–1905) commandee par le Dr. Jean Charcot, Sciences naturelles: Documents scientifiques: Echinodermes (Stellerides, Ophiures et Echinides), pages 140–147, 4 plates. Paris: Masson et Cie.

1907.   Note preliminaire sur quelques Asteries et Ophiures provenant des campagnes de la Princesse Alice. Bulletin de l'Institut Oceanographique de Monaco, 99: 47 pages.

1908.   Asteries, Ophiures et Echinides de l'Expedition antarctique nationale ecossaise. Transactions of the Royal Society of Edinburgh, 48:529–649.

1909a. Echinodermes provenant des campagnes du yacht Princesse Alice Asteries, Ophiures, Echinides et Crinoides). Resultats du Campagnes Scientifique de Monaco, 34: 317 pages.

1909b. Echinodermes recueillis dans les mers arctiques par le Mission arctique francaise, commandee par M. Benard. Bulletin de la Musee de Paris, 1909:121–123.

1909c. Echinoderma of the Indian Museum, Asteroidea: An Account of the Deep-sea Asteroidea Collected by the Royal Indian Marine Survey Ship Investigator, 1: Les Asteries de Mer profonde. 144 pages, figures 1–13. Calcutta: Trustees of the Indian Museum.

1921.   Echinodermes (Asteries, Ophiures, Echinides et Crinoides) des dernieres campagnes de la Princesse Alice et de l'Hirondelle II. Bulletin de l'Institut Oceanographique de Monaco, 396:1–8.

Korovchinsky, N.M.

1976.   New Species of Abyssal Starfishes of the Genus Freyella (Brisingidae) from the South Atlantic. Zoologicheskii Zhurnal, 55(8):1187–1194, 5 figures, 3 tables. [In Russian, with English summary.]

Korovchinsky, N.M., and S.V. Galkin

1984.   New Data on the Fauna of Starfishes of the Genus Freyella (Brisingidae). Zoologicheskii Zhurnal, 63(8): 1205–1215, 2 figures. [In Russian, with English summary].

Ludwig, H.

1878.   Zur Kenntniss der Gattung Brisinga. Zoologische Anzieger, 2:216–234, plate 15.

1897a. Diagnosen der Seesterne des Mittelmeeres. Verhandlungen des Naturhistorischen Vereins der Preussischen Rheinlande, Westfalens und des Reg.-Bezirks Osnabrück, 53:281–309.

1897b. Die Seesterne des Mittelmeeres. In Fauna und Flora des Golfes von Neapel und der angrenzenden Meeres-Abschnitte, herausgegeben von der zoologischen Station zu Neapel, 24: 491 pages, 12 plates.

1900.   Arktische und subarktische Holothurien: Arktische Seesterne. In F. Roemer and F.R. Schaudinn, Fauna Arctica, 1(5):447–502.

1903.   Seesterne. In Resultats du voyage du S.Y. Belgica, Zoologie. 72 pages, 7 plates. Anvers: Commission de la Belgica.

1905a. Asteroidea. Memoirs of the Museum of Comparative Zoology at Harvard, 32: 292 pages, 35 plates.

1905b. Asterien und Ophiuren der schwedischen Expedition nach den Magalhaenslaendern 1895–1897. Zeitschrift fur Wissenschaftliche Zoologie, 82:39–79, plates 5–6.

Madsen, F.J.

1950.   The Echinoderms Collected by the Atlantide Expedition 1945–46, I: Asteroidea, with Remarks on Other Sea-stars from Tropical and Northern West Africa. In Atlantide Reports, 1:167–222, figures 1–11, plates 14–16. Copenhagen: Danish Science Press.

1956.   Echinoidea, Asteroidea, and Ophiuroidea from Depths Exceeding 6000 Meters. Galathea Reports, 2:23–32, 1 plate, 5 figures.

Marion, A.F.

1883.   Les progres recents des sciences naturelles. Revue Scientifique, 5:129–136.

McKnight, D.G.

1973.   Additions to the Asteroid Fauna of New Zealand: Families Benthopectinidae, Odontasteridae, Asteriidae and Brisingidae; with Notes on Porcellanaster caeruleus Wyville Thomson (Family Porcellanasteridae). New Zealand Oceanographic Institute Records, 1(16):220–240.

1975.   Classification of Somasteroids and Asteroids (Asterozoa: Echinodermata). Journal of the Royal Society of New Zealand, 5(1):13–19.

Mortensen, T.

1927.   Handbook of the Echinoderms of the British Isles. 471 pages. England: Oxford University Press.

Norman, A.M.

1893.   A Month on the Trondhjem Fiord, [Part 1]. Annals and Magazine of Natural History, 6:xii, 341–367.

Pawson, D.L.

1976.   Some Aspects of the Biology of Deep-sea Echinoderms. Thalassia Jugoslavica, 12(1):287–293.

Perrier, E.

1882.   Note sur les Brisinga. Comptes-rendus Hebdomadaires des Seances de l'Academie des Sciences (Paris), 95:61–63.

1885a. Sur les stellerides recueillis durant la mission du Talisman. Comptes-rendus Hebdomadaires des Seances de l'Academie des Sciences (Paris), 101:884–887.

1885b. Note preliminaire sur les echinodermes, recueilles par Travailleur et Talisman. Nouvelles Archives des Missions Scientifiques et Litteraires, Choix de Rapports et Instructions, series 2, 6:1–154.

1885c. Premiere note preliminaire sur les echinodermes recueilles durant les campagnes de dragages sousmarins du Travailleur et du Talisman. Annales des Sciences Naturelles (Zoologie et Paleontologie), series 6, 19(8):72 pages.

1885d. Sur les Brisingidae de la mission du Talisman. Comptes-rendus Hebdomadaires des Seances de l'Academie des Sciences (Paris), 101:441–444.

1891a. Stellerides nouveaux provenant des campagnes du yacht l'Hirondelle. Memoires de la Société Zoologique de France, 4:258–271.

1891b. Sur les stellerides recueilles dans la Golfe de Gascogne, aux Acores et a Terre-Neuve Pendant les campagnes scientifiques du yacht l'Hirondelle. Comptes-rendus Hebdomadaires des Seances de l'Academie des Sciences (Paris), 112(21):1225–1228.

1891c. Echinodermes de la mission scientifique du Cape Horn, I: Stellerides. Mission Scientifique du Cape Horn, Zoologie, 4: 198 pages, 13 plates.

1894.   Echinodermes, I: Stellerides. In Expeditions Scientifique de Travailleur et du Talisman, 3: 432 pages, 26 plates.

1896.   Contribution à l'étude des stellerides de l'Atlantique Nord (Golfe de Gascogne, Acores, Terre-Neuve). Resultats des Campagnes Scientifiques du Prince de Monaco, fasc. 2, viii + 73 pages, plates 1, 1c.

Sars, G.O.

1871.   Nye Echinodermer fra den Norske Kyst. In Forhandlinger i Videnskabs-Selskabet i Christiania, pages 1–31.

1875.   I: On Some Remarkable Forms of Animal Life from the Great Deeps off the Norwegian Coast, II: Researches on the Structure and Affinities of the Genus Brisinga, Based on the Study of a New Species, B. coronata. 112 pages, 7 plates. Christiania: Christiana University.

Sibuet, M.

1975.   Asterides abyssales de l'Atlantique sud (Resultats de la campagne Walda, juin-juillet-aout 1971). Bulletin de la Musee Nationale d'Histoire Naturel de Paris (Zoology), 199:281–296, 3 figures, plate, 2 tables.

Sladen, W.P.

1889.   Report on the Asteroidea. Report on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1873–76, Zoology, 30(51): xlii + 893 pages, 118 plates.

1891.   Report on a Collection of Echinodermata from the Southwest Coast of Ireland. Proceedings of the Royal Irish Academy, series 3i, 1:687–704, plates 25–29.

Spencer, W.K., and C.W. Wright

1966.   Asterozoans, Part U: Echinodermata. In R.C. Moore, editor, Treatise on Invertebrate Paleontology, 3(1):U4–U107, 89 figures. Lawrence: University of Kansas Press.

Thomson, C.W.

1873.   On the Echinoidea of the Porcupine Deep-sea Dredging Expedition. Philosophical Transactions of the Royal Society of London, clxiv + 719–756, plates 59–71.

Tortonese, E.

1958.   Euclasteroidea: Nuovo ordine di Asteroidi (Echinodermi). Doriana, 2(88):1–3.

Verrill, A.E.

1880.   Notice of Recent Additions to the Marine Invertebrata of the Northeastern Coast of America, with Descriptions of New Genera and Species and Critical Remarks on Others. American Journal of Science, 19:137–140.

1884.   Notice of the Remarkable Marine Fauna Occupying the Outer Banks off the Southern Coast of New England, and of Some Additions to the Fauna of Vineyard Sound. In Annual Report of the U.S. Commissioner of Fish and Fisheries for 1882, pages 378–384.

1885.   Results of the Explorations Made by the Steamer Albatross off the Northern Coast of the United States in 1883. In Annual Report of the U.S. Commissioner of Fish and Fisheries for 1883, pages 503–699, 44 plates.

1894.   Descriptions of New Species of Starfishes and Ophiurans, with a Revision of Certain Species Formerly Described. Proceedings of the United States National Museum, 17:245–297.

1895.   Distribution of the Echinoderms of Northeastern America. American Journal of Science, 49:127–141, 199–212.

von Marenzeller, E.

1893a. Veroffentlichungen der Commission fur Erforschung des Ostlichen Mittelmeeres. (Vorlaufige Mittheilung): Neue Echinodermen aus dem Mittelmeeres. Sitzungsberichte der Mathematisch-Naturwissenschaftlichen Classe der Kaiserlichen Akademie der Wissenschaften, 52(1):66–70.

1893b. Zoologische Ergebnisse der Tiefsee-Expeditionen im ostlichen Mittelmeere auf S.M. Schiff Pola, I: Echinodermen, gesammelt 1890, 1891 und 1892. Anzieger der k. Akademie der Wissenschaften, Mathematisch Naturwissenschaftliche Classe, 8:65–67.

1895a. Echinodermen, gesammelt im ostlichen Mittelmeere, 1893 und 1894. Anzieger der k. Akademie der Wissenschaften, Mathematisch Naturwissenschaftliche Classe, 28:189–191.

1985b. Zoologische Ergebnisse, V: Echinodermen, gesammelt 1893, 1894 Denk Akademie Wien, 52:123–148, 1 plate.

Wood-Mason, J., and A. Alcock

1891.   Natural History Notes from H.M. Marine Survey Steamer Investigator. Annals and Magazine of Natural History, series 6, 7:12–15, 427–433.

Index

(Synonyms and page numbers of principal entries are in italics.)

abyssicola, Freyella, 43

actinodetus, Astrocles, 43

americana, Novodinia, 2, 4, 21, 22

antillensis, Novodinia, 2, 23, 25

aspera, Freyella, 43, 45

Astrocles, 43

Astrostephane, 1

Belgicella, 1

benthophila, Freyastera, 2, 36, 38

bracteata, Freyella, 43, 45

Brisinga, 1, 4, 5, 7, 9, 11, 13, 15, 17, 21, 25, 26, 29, 33, 43, 45

Brisingaster, 1

Brisingella, 1, 2, 4, 5, 13, 15, 16

Brisingenes, 1, 2, 5, 13, 16, 17

Brisingida, 1, 2, 3, 4, 41, 50

Brisingidae, 1, 2, 4, 5, 33

Colpaster, 1, 2, 33, 34, 36

coronata, Brisingella, 1, 4, 13, 15, 16

costata, Brisinga, 2, 5, 7

Craterobrisinga, 5, 7, 17

cricophora, Brisinga, 2, 5, 7, 9

dimorpha, Freyella, 46

edwardsi, Colpaster, 33, 34

elegans, Freyella, 2, 43, 46, 48, 49

elegans, Odinia, 25, 26

endecacnemos, Brisinga, 1, 5, 9, 11

Euclasteroidea, 1, 2, 4

Freyastera, 1, 2, 4, 33, 36, 38, 40, 41, 43

Freyella, 1, 2, 25, 26, 33, 36, 38, 40, 41, 43, 45, 46, 48, 49

Freyellaster, 1

Freyellidae, 33

Freyellidea, 43

benthophila, 36

tuberculata, 41

hirsuta, Brisinga, 2, 4, 5, 7, 9, 11, 13

homonyma, Novodinia, 25, 26, 27, 43

laubieri, Freyella, 43, 45

mediterranea, Brisinga, 13

mexicana, Freyastera, 4, 38, 40

microspina, Freyella, 2, 43, 46, 48

Midgardia, 1, 5, 13, 17, 18, 19

multicostata, Brisingenes, 2, 17

mutabilis, Freyella, 46

Novodinia, 1, 2, 4, 5, 13, 17, 19, 21, 23, 25, 27, 29

Odinia, 21

antillensis, 23

elegans, 25, 26

pandina, 27

robusta, 29

semicoronata, 29

pandina, Novodinia, 2, 23, 25, 26, 27, 29

Parabrisinga, 1

pennata, Freyella, 46

placoderma, Stegnobrisinga, 31

recta, Freyella, 46, 48, 49

robusta, Odinia, 29

scutigerula, Colpaster, 2, 33, 34, 36

semicoronata, Novodinia, 21, 23, 25, 27, 29

sexradiata, Freyastera, 36, 38, 40, 41, 43

spinosa, Freyella, 43, 45

spinulosa, Freyella, 45

splendens, Stegnobrisinga, 2, 31, 33

Stegnobrisinga, 1, 2, 5, 13, 17, 19, 21, 29, 31

trispinosa, Freyella, 2, 46, 48

tuberculata, Freyastera, 38, 40, 41, 43

verticellata, Brisingella, 2, 15, 16

xandaros, Midgardia, 18, 19

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