NOAA Technical Report NMFS 91

Marine Flora and Fauna

of the Northeastern United States

Echinodermata: Crinoidea

Charles G. Messing

John H. Dearborn

August 1990

u.s. Department of Commerce
NOAA Technical Report NMFS _

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and may be reprinted entirely, reference to source is appreciated.

61. Manual for starch gel electrophoresis: A method for the detection

of genetic variation, by Paul B. Aebersold, Gary A. Winans, David J. Teel,

George B. Milner, and Fred M. Utter. December 1987, 19 p.

62. Fishery publication index, 1980-85; Technical memoradum index,

1972-85, by Cynthia S. Martin, Shelley E. Arenas, Jacki A. Guffey, and

Joni M. Packard. December 1987, 149 p.

63. Stock assessment ofthe Atlantic menhaden, Brevoortia tyrannus, fishery,

by Douglas S. Vaughan and Joseph W. Smith. January 1988, 18 p.

64. Illustrated key to penaeoid shrimps of commerce in the Americas,

by Isabel Perez Farfante. April 1988, 32 p.

65. History ofwhaling in and near North Carolina, by Randall R. Reeves

and Edward Mitchell. March 1988, 28 p.

66. Atlas and zoogeography of common fishes in the Bering Sea and

northeastern Pacific, by M. James Allen and Gary B. Smith. April 1988,

151 p.

67. Index numbers and productivity measurement in multispecies

fisheries: An application to the Pacific coast trawl fleet, by Dale Squires.

July 1988, 34 p.

68. Annotated bibliography II ofthe hard clam Mercenari4 mrrctTUlria, by

J. L. McHugh and Marjorie W. Sumner. September 1988, 59 p.

69. Environmental quality and aquaculture systems: Proceedings of the

thirteenth U.S.-Japan meeting on aquaculture, Mie, Japan, October 24-25,

1984, edited by Carl J. Sindermann. October 1988, 50 p.

70. New and innovative advances in biology/engineering with potential

for use in aquaculture: Proceedings ofthe fourteenth U.S.-Japan meeting

on aquaculture, Woods Hole, Massachusetts, October 16-17, 1985, edited

by Albert K. Sparks. November 1988, 69 p.

71. Greenland turbot ReinJlarthius hippoglossoides ofthe eastern Bering Sea

and Aleutian Islands region, by Miles S. Alton, Richard G. BakkaIa, Gary

E. Walters, and Peter T. Munro. December 1988, 31 p.

72. Age determination methods for northwest Atlantic species, edited

by Judy Penttila and Louise M. Dery. December 1988, 135 p.

73. Marine flora and fauna of the Eastern United States. Mollusca:

Cephalopoda, by Michael Vecchione, Clyde F. E. Roper, and Michael

J. weeney. February 1989, 23 p.

74. Proximate composition and fatty acid and cholesterol content of 22

species of northwest Atlantic finfish, by Judith Krzynowek, Jenny Murphy, Richard S. Maney, and Laurie J. Panunzio. May 1989, 35 p.

75. Codend selection of winter flounder PseudlJpleuronutes omeri&onus, by

David G. Simpson. March 1989, 10 p.

76. Analysis of fish diversion efficiency and su"'ivorship in the fish return

system at San Onofre Nuclear Generating Station, by Milton S. Love,

Meenu Sandhu, Jeffrey Stein, Kevin T. Herbinson, Robert H. Moore,

Michael Mullin, and John . Stephens Jr. April 1989, 16 p.

77. Illustrated key to the genera of free-living marine nematodes of the

order Enoplida, by Edwin J. Keppner and Armen C. Tarjan. July 1989,

26 p.

78. Su",ey of fishes and water properties of south San Francisco Bay,

California, 1973-82, by Donald E. Pearson. August 1989, 21 p.

79. Species composition, distribution, and relative abundance of fishes

in the coastal habitat off the southeastern United States, by Charles A.

Wenner and George R. Sedberry. July 1989, 49 p.

80. Laboratory guide to early life history stages of northeast Pacific fishes,

by Ann C. Matarese, Arthur W. Kendall Jr., Deborah M. Blood, and

Beverly M. Vinter. October 1989, 651 p.

81. Catch-per-unit-effort and biological parameters from the Massachusetts coastal lobster (Homorus americanus) resource: description and Trends,

by Bruce T. Estrella and DanielJ. McKiernan. September 1989, 21 p.

82. Synopsis of biological data on the cobia RiJ&hycentron canadwn (Pisces:

Rachycentridae), by Rosalie Vaught Shaffer and Eugene L. Nakamura.

December 1989, 21 p.

83. Celaphopods from the stomachs ofsperm whales taken ofTCalifornia, by Clifford H. Fiscus, Dale W. Rice, and Allen A. Wolman. December 1989, 12 p.

84. Results of abundance surveys ofjuvenile Atlantic and Gulf menhaden, Brevoortia tyrannus and B. palrunus, by Dean W. Ahrenholz, James F.

Guthrie, and Charles W. Krouse. December 1989, 14 p.
NOAA Technical Report NMFS 91

Marine Flora and Fauna

of the Northeastern United States

Echinodermata: Crinoidea

Charles G. Messing

John H. Dearborn

August 1990

u.s. DEPARTMENT OF COMMERCE

Robert Mosbacher, Secretary

National Oceanic and Atmospheric Administration

John A. Knauss, Under Secretary for Oceans and Atmosphere

National Marine Fisheries Service

William W. Fox Jr., Assistant Administrator for Fisheries
Foreword _

This NOAA Technical Report NMFS is part of the subseries "Marine Flora and Fauna

of the Eastern United States" (formerly "Marine Flora and Fauna of the Northeastern

United States"), which consists of original, illustrated, modern manuals on the identification, classification, and general biology of the estuarine and coastal marine plants

and animals of the eastern United States. The manuals are published at irregular intervals on as many taxa of the region as there are specialists available to collaborate

in their preparation. These manuals are intended for use by students, biologists,

biological oceanographers, informed laymen, and others wishing to identify coastal

organisms for this region. They can often serve as guides to additional information about

speCIes or groups.

The manuals are an outgrowth of the widely used "Keys to Marine Invertebrates

of the Woods Hole Region," edited by R.I. Smith, and produced in 1964 under the

auspices of the Systematics Ecology Program, Marine Biological Laboratory, Woods

Hole, Massachusetts. Geographic coverage of the' 'Marine Flora and Fauna of the

Eastern United States" is planned to include organisms from the headwaters of estuaries

seaward to approximately the 200-m depth on the continental shelf from Maine to

Florida, but can vary somewhat with each major taxon and the interests of collaborators.

Whenever possible, representative specimens dealt with in the manuals are deposited

in the reference collections of major museums.

The "Marine Flora and Fauna of the Eastern United States" is being prepared

in collaboration with systematic specialists in the United States and abroad. Each manual

is based primarily on recent and ongoing revisionary systematic research and a fresh

examination of the plants and animals. Each manual, treating a separate major taxon,

includes an introduction, illustrated glossary, uniform originally illustrated keys, annotated checklist (with information, when available, on distribution, habitat, life history,

and related biology), references to the major literature of the group, and a systematic

index.

Manuals are available from the National Technical Information Service, United

States Department of Commerce, 5285 Port Royal Road, Springfield, VA 22161, either

as a paper copy or microfiche, for a charge. Manuals are not copyrighted, and so can

be photocopied from the NOAA Technical Report NMFS Circulars and Reports

available in most major libraries and listed at the end of this manual.

The National Marine Fisheries Service (NMFS) does not approve,

recommend or endorse any proprietary product or proprietary

material mentioned in this publication. No reference shall be made

to NMFS, or to this publication furnished by NMFS, in any advertising or sales promotion which would indicate or imply that NMFS

approves, recommends or endorses any proprietary product or proprietary material mentioned herein, or which has as its purpose

an intent to cause directly or indirectly the advertised product to

be used or purchased because of this NMFS publication.

Text printed on recycled paper

11
Marine Flora and Fauna of the Northeastern United States

Echinodermata: Crinoidea*

CHARLES G. MESSING

Nova Oceanographic Center

8000 North Ocean Drive

Dania, FL 33004

JOHN H. DEARBORN

Department of Zoology

University of Maine

Orono, ME 041:69

ABSTRACT

The crinoid fauna of the continental margin (0-1500 m) of northeastern North America (Georgia

to Canada) includes 14 species in 13 genera and 5 families. We introduce the external morphology

and natural history of crinoids and include a glossary of terms, an illustrated key to local taxa,

annotated systematic list, and an index. The fauna includes 2 species found no further south

than New England and 8 that occur no further north than the Carolinas and Blake Plateau. Comactinia meridionalis (Agassiz) is the only species commonly found in shallow water «50 m). No

taxa are endemic to the area.

Introduction _

The class Crinoidea includes the most exquisite members

of the phylum Echinodermata and the only extant forms

that bear a stalk. Forms known as sea lilies retain a stalk

throughout their lives. Feather stars, or unstalked crinoids

(order Comatulida), retain a stalk through a postlarval pentacrinoid stage, but discard it and take up a free existence

as juveniles. All living forms have at least a superficial pentamerous radial symmetry and flexible featherlike arms.

The arms bear reproductive organs and extensions of the

water vascular system that serve for food capture and

respiration. Crinoids orient in life with the oral surface uppermost, away from the substrate; both mouth and anus

are on the oral surface.

Extant crinoids belong to the subclass Articulata which

includes almost all post-Paleozoic taxa. Modern forms

display a much narrower range of structural diversity than

their far more numerous fossil forebears. The following

discussion of morphology and natural history refers only

to living species.

Crinoid taxonomy is not stable. Revisions at all taxonomic levels are either ongoing or needed and small

numbers of new taxa are described each year. Between 600

and 700 extant species are generally recognized worldwide;

'Contribution No.3 from the Deep Ocean Society, Miami, FL.

about 70 occur in the North Atlantic. This paper includes

crinoids found along the continental margin (0-1500 m)

of northeastern North America (Georgia and northern

Blake Plateau to Canada and Greenland). Several taxa currently known only as far north as the northern slopes of

the Little Bahama Bank may eventually be found further

north on the Blake Plateau and within the area covered

by this manual. Included taxa belong to the order Bourgueticrinida (family Bathycrinidae) and order Comatulida

(families Atelecrinidae, Comasteridae, Antedonidae, and

Pentametrocrinidae).

External Morphology _. _

Crinoid taxonomy is based chiefly on the endoskeleton of

articulated calcareous plates, or ossicles, that composes a

very large proportion of the whole animal (Figs. 1, 2, 3).

This skeleton is far more obvious in crinoids than in other

echinoderms. Most of the ossicles are covered only with

a thin epidermis; they are clearly visible, although a dissecting microscope is generally required for recognition of

diagnostic features.

The typical crinoid body is more or less lenticular and

rests in a cuplike calyx composed of two circlets usually

of five ossicles each (Figs. 1, 3). Basal ossicles, oriented

interradially, form the aboral circlet. They are reduced in

comatulids and generally not visible without dissection.
2 NOAA Technical Report NMFS 91 _

Figure 1

Lateral view of a stalked crinoid (order Bourgueticrinida) showing upper (proximal) portion of stalk, calyx, and bases of three

rays.

Radla18 ---\--'.--

Baaaia

(fu8ed)-~-

Calyx

Columnals

Radial ossicles, associated with the five water vascular

canals that radiate from the circumoral ring canal, form

the second circlet. Radial and basal ossicles are immovably

joined to each other. Both are at least partly obscured from

external view by an aboral structure that supports the

crinoid above the substrate. In sea lilies, it is a stalk. In

comatulids, the calyx rests on a large modified ossicle, the

centrodorsal, derived from the topmost segment of the

postlarval stalk. (Phylogenetically, the centrodorsal may

represent several fused stalk ossicles.) Centrodorsals range

from disklike through hemispherical to cylindrical and

usually bear numerous unbranched anchoring cirri. A

typical cirrus is more or less prehensile, with the terminal

segment modified as a claw and the penultimate bearing

an opposing spine. Cirri range from short, robust and

highly prehensile to long, slender and relatively straight.

Rootlike segmented branching structures restricted to the

stalk base of Bathycrinidae are also called cirri but are

not homologous with cirri arising from the comatulid

centrodorsal.

Each radial is the first, or most proximal, ossicle of a

ray. Each ray consists of a complex, usually branched,

series of ossicles that extends outward from the body, bearing an open ambulacral groove bordered by small fingerlike tube feet. Ossicles following the radials are called

brachials. In rays that branch, each series of brachials

leading from the radial to a branch point or from one

branch point to another is termed a division series (or

brachitaxis). Each division series is usually composed of

two or four brachials and terminates with a modified axillary (or axil) ossicle that bears two articular faces distally. An axillary may thus bear two unbranched arms, two

additional division series, or one of each. In unbranched

rays, the arms extend from the radials. Young crinoids first

develop five arms. Additional arms develop when one is

shed and two grow back in its place. Many species have

ten arms as a result; other chiefly tropical taxa have as

many as 200.

Each arm bears smaller segmented unbranched appendages, pinnules, arranged in alternating series on either

side. Arms, division series, and pinnules carry extensions

of coelomic, nervous, and water vascular systems. Pinnules

give the arms their characteristic featherlike appearance

and are also the primary site of food collection. One or

more pairs of pinnules near the arm base may be modified

and lack an ambulacral groove. Such oral pinnules are

often longer and more flexible or, in some families, more

robust and spinelike than those succeeding. They are

modified ostensibly for sweeping or protecting the oral surface. Oral pinnules are followed by several pairs of genital

pinnules bearing the gonads. These rarely exhibit skeletal

modifications; they are usually similar to the more distal,

exclusively food-collecting pinnules.

In most modern crinoids, the cuplike structure of the

calyx is lost; the calyx no longer surrounds the visceral
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 3

Figure 2

Lateral diagrammatic view of an unstalked crinoid or feather star (order Comatulida) showing centrodorsal with some cirri, and three

rays. (Left- and right-hand rays show pinnules on only one side of each arm.)

mass. Instead, the viscera rest on top of the radial circlet

and proximal ossicles of the rays. The oral surface of the

visceral mass, the tegmen, or disk, is usually naked integument although it may be invested with small calcareous

pieces. The tegmen bears both the anus, located at the apex

of a small tube or papilla, and the mouth with its radiating

ambulacral trunks. Portions of arms unattached to the

visceral mass and distal to the tegmen are sometimes called

free arms. The mouth is usually located centrally on the

tegmen with the anus displaced to one side. In most Comasteridae the reverse is true.

Most articulations between successive ray ossicles contain both muscles and ligaments and are called muscular

articulations. Moving from the aboral to the ambulacral

surface of a typical brachial articular face, each articulation consists of a large aboral ligament, a transverse fulcral

ridge, a pair of smaller ligaments and a pair of muscles.

Ligaments and muscles insert into depressions, or fossae,

in articular faces.

Exclusively ligamentary articulations have little or no

flexibility. Syzygies are rigid articulations in which flat

ridges radiating from the center of one articular face correspond with ridges on the other face. The ridges alternate with shallow grooves that also correspond on the two

faces. Ridges and grooves do not interlock; the external

appearance of the joint resembles a perforated line.

Syzygies represent fracture points for arm autotomy and

typically occur at regular, sometimes diagnostic intervals.

Synarthries are ligamentary articulations characterized by

a pair ofsemicircular ligaments separated by a fulcral ridge

oriented vertically, that is, through the oral/aboral axis of

the ray. Among comatulids, they are usually restricted to

the middle joint in a division series of two ossicles (or the

first joint in a division series of four ossicles), and the articulation between the first two arm brachials. The aboral

surface around the joint of a pair of brachials united by

synarthry is frequently swollen as a synarthrial tubercle.

Such swellings are typical of many antedonids but are
4 NOAA Technical Report NMFS 91 _

Cirrus Socket

Opposing Spine

Terminal Claw-

'"

Figure 3

Lateral view of an unstalked crinoid (order Comatulida) showing centrodorsal, two cirri, and bases of three rays.
____________ Messing and Dearborn: Echinode~mata, Crinoidea of the Northeastern United States 5

absent in comasterids. In both syzygies and synarthries,

the ossicle proximal to the articulation lacks a pinnule.

Hourglass-shaped columnals ofthe middle and lower stalk

of Bathycrinidae are also united by synarthries.

Symbols _

Because of the highly repetltlve nature of the crinoid

skeleton, taxonomic descriptions are greatly condensed

by using symbols for a variety of skeletal parts. Considerable controversy has surrounded the identification

of homologous ossicles across various crinoid groups, and

the terms and symbols have varied quite a bit among

authors. However, most of the taxa covered in this manual can be described with fair uniformity. We refer the

reader to Ubaghs (1953), Clark (1970), and Breimer (1978)

for more thorough discussions of crinoid structural

homologies.

Basal ossicles are abbreviated B (pI. BB) and radials R

(pI. RR). Each successive division series is indicated by

a roman numeral followed by Br for brachial. An arabic

number following indicates the number of ossicles composing the division series. Thus, IBr2 is the rust division

series composed oftwo ossicles; IIBr4 is a second division

series composed offour ossicles. Individual division series

ossicles are indicated by arabic subscript (e.g., IBr2'

IIIBr~). Brachials of undivided arms are indicated as Brl>

Br2' etc. A plus ( + ) sign indicates a syzygy. For example, IIBr4(3 + 4) indicates a second division series of four

ossicles with the third and fourth joined by syzygy; Brg +10

indicates brachials nine and ten joined by syzygy. In Conocnnus, which has unbranched arms, we follow Gislen (1938)

and Clark (1970) in numbering brachials from the first

ossicle following the radial.

Pinnules are indicated by P. Arabic subscript numbers

indicate successive exterior pinnules, that is, those along

the outside of an arm; subscript lower case letters indicate

interior pinnules (e.g., PI> P2, Pa' Pb)' The interior side

of an arm or division series is that side closest to the extrapolated median axis of the ray. In the ten-armed species

included in this key (except Atelecrinus which has deficient

pinnulation), Br2 bears PI> Pa is on the opposite side of

Br4 [Br3 and Br~ are joined by syzygy (Br3 +~) so there is

no pinnule on Br3], Brs bears P2, and Br6 bears Pb'

Bathycrinids have relatively few pinnules that normally do

not require specific enumeration.

In species descriptions, number of cirri is given in roman

numerals and number of cirrals that compose a cirrus is

given in arabic numerals. LW of cirrals indicates ratio

of length to proximal width. Centrodorsal DH indicates

ratio of greatest diameter to oral/aboral height. Arm LD

indicates ratio of midradial ray length (L) from the distal

margin of a radial to Brg +10, to arm diameter (D) at

Br3 +4'

Distribution _

The region covered by this manual represents a depauperate, transitional area for crinoids. No species are endemic

and only Atelecrinus balanoides and, possibly, Trichometra

cubensis and Conocrinus lofotensis have ranges spanning the

region. Three species, Heliometraglacialis, Poliometra prolixa,

and Hathrometra tmella, represent a boreal and Arctic fauna

that reaches its southern and westermost outposts in the

northern part of this region. Over twenty additional taxa

occur in the northeastern Atlantic. Eight species, Democnnus

conifer, Pentametrocrinus atlanticus, Comatilia iridometriformis,

Comactinia echinoptera, Comactinia meridionalis, Comatonia cristata, Coccometra hagenii, and Zenometra columnaris, represent

elements of the rich tropical western Atlantic fauna that

penetrates no further north than the Blake Plateau and

Carolina Capes. Ofthese, only P. atlanticus occurs on both

sides ofthe Atlantic. About two dozen additional taxa reach

no further north than the Straits of Florida and Bahama

Islands. Stalked crinoids are rare and poorly documented

in the region. By contrast, the northeastern Atlantic (to

about 2000 m) supports a varied fauna comprising ten

species in four (possibly five) families; the tropical western

Atlantic includes at least nine species in three families.

The majority of species occur on the outer continental

shelf and slope. Comactinia meridionalis is the only species

that normally occurs in shallow populations. Shallow

records of other species are either unique (Comatonia cristata, Coccometra hagenit) or occasional (Hathrometra tmella,

Heliometra glacialis) in the region.

Natural History _

All extant crinoids are suspension feeders. They produce

no respiratory/feeding current but rather rely on extrinsic

water movement. For many years, our understanding of

their ecology was both sketchy and fanciful. Crinoids were

once supposed to sit on the sea floor with arms spread out

in an upturned bowl, subsisting on a slow rain of detritus

from above (Hyman 1955; Nichols 1962). It was also

reported that they entangled prey with rhythmic scooping

motions of their arms (Fell 1966). SCUBA and submersible observations have largely disproven both views (e.g.,

Meyer 1973; Macurda and Meyer 1974) and have also indicated that crinoids are not completely passive filter

feeders.

Ecological studies of extant crinoids are largely descriptions of feeding postures and strategies because the foodgathering apparatus comprises such a large proportion of

the animal's structure. It is critical to keep in mind,

however, that the arms and pinnules that support the foodgathering apparatus also function in locomotion in most

extant taxa. This dual obligation requires that the skeleton

be rigid enough to stand erect against a passing current,
6 NOAA Technical Report NMFS 91 _

yet flexible enough to permit movement. The muscular articulations described above fit this requirement as follows:

contractions of muscles on the ambulacral side ofthe fulcral

ridges curl or roll the arms inward toward the mouth and

flex pinnules toward the arm axis. When the muscles relax,

the elasticity of the large antagonistic ligament extends

arms and pinnules outward, allowing them to maintain an

extended posture passively against a current for food

gathering. Individual articulations have limited scope but

an arm of over 200 segments or a pinnule with more than

50 may have great flexibility. Locomotion is achieved in

comatulids and some stalked crinoids (Isocrinida) by arm

crawling. Some comatulids are capable of swimming with

graceful and coordinated arm undulations although they

appear to do so only rarely.

In extant crinoids, the food-gathering apparatus functions as follows: each arm and pinnule bears an open ambulacral groove bordered by fingerlike podia, or tube

feet-terminal extensions of the water vascular system.

Unlike many asteroids, echinoids, and holothuroids,

crinoid tube feet lack a terminal sucker. Tube feet are arranged in groups of three, each group including a short,

medium, and long tube foot. The longest in each group,

0.5-0.85 mm in length (Meyer 1979), is the primary food

gatherer. It is held out at a right angle and flicks passing

food particles into the pinnule's ambulacral groove. Pinnule grooves run into arm grooves which converge like

tributary rivers on the mouth. Captured particles are

wrapped in mucous secretions and are transported mouthward by ciliary tracts that line the groove floor.

Food particle size depends chiefly upon tube-foot spacing and ambulacral groove width, and ranges from about

50 to 400 lAm. Diets include a variety of protists (e.g.,

diatoms and other unicellular algae, foraminiferans, actinopods), invertebrate larvae, small crustaceans, and

detrital particles.

All extant crinoids appear to use the same basic foodgathering system. Nevertheless, they do occupy a variety

of niches based largely on flow regime preferences. Rheophilic forms prefer relatively strong or consistent, uni- or

bi-directional flow. Most raise themselves above the

substrate in order to intercept a satisfactory current. Sea

lilies, of course, have the advantage of a stalk. Rheophilic

comatulids typically cling to advantageous perches such as

sponges, scleractinians, alcyonarians, projecting hard

substrates and even sea lily stalks. Once above the substrate

in a satisfactory "breeze," rheophiles arrange arms and

pinnules in a variety of postures including parabolic and

radial fans, and bi- and multi-layered vertical fans.

Rheophobic species typically conceal their calyx within

crevices or under ledges on reefs and other rugged bottoms.

Though still dependent upon water movement for food and

gas exchange, they prefer a slower, multidirectional flow

regime. Rheophobes typically extend their feeding arms

with no particular orientation, but may direct successive

pinnules at 900 angles to each other, a more efficient arrangement for intercepting particles that may come from

any direction. Some deep-water taxa exposed to low flow

velocities do arrange their rays in a bowl, apparently for

gravitational interception of particles.

Some shallow-water comatulids remain exposed at all

times; others are nocturnal and remain hidden during the

day. Both groups include rheophilic and rheophobic forms.

Species requiring exposed feeding perches are subject

to predation by a variety of fishes and invertebrates

(Mladenov 1983; Meyer et al. 1984; Meyer 1985) but

attacks are rarely fatal. Predators usually crop one or more

arms or pinnules, or remove the central visceral mass. The

latter regenerates in less than a month. Defenses include

diurnal seclusion (most teleosts that feed on calcareous

prey are day-active); protection of the visceral mass by

semicryptic behavior, development oflarge spinelike oral

pinnules, or a dense thicket of arms and pinnules; incorporation of distasteful or toxic compounds; and crawling/

swimming.

Crinoids host many parasites and commensals including

pontoniine shrimp, cyclopoid copepods, isopods, cirripedes, gastropods, ophiuroids, clingfish, and myzostome

polychaetes (the latter found almost exclusively on crinoids). These range from forms that use the crinoid as a

convenient perch, through those that steal food from the

ambulacral grooves, to parasites that feed directly on tissues

and body fluids. Many match both color and pattern of

the host crinoid.

Sexes are separate and almost uniformly identical.

Reproductive modes among comatulids (sea lily development remains unknown) include broadcast spawning (the

majority ofspecies for which spawning is known), and both

external and internal brooding. Free-swimming larvae,

when present, are barrel-shaped, ciliated lecithotrophic

doliolariae or vitellariae. With the exception of Comatilia

iridometriformis, the twenty or so known internally brooding

species are confined to middle and high southern latitudes.

The latter include the only species with easily distinguishable sexes; females brood embryos in a marsupium, a

pouch adjacent to the gonad on each genital pinnule.

Crinoids are difficult to maintain in aquaria because of

their need for a continuous flow of plankton-laden water.

Preservation and Examination _

Crinoids may be fixed in 10% buffered formalin and

transferred to or preserved directly in 70% ethanol. Arm

breakage is often a significant problem. Some species

autotomize arms more easily than others. Similarly, some

taxa respond to preservative by flexing their arms aborally,

others orally. According to D. L. Meyer (Dep. Geology,

Univ. Cincinnati, pers. commun., Jan. 1990), crinoids can

be killed, often without major arm breakage, by immer-
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 7

sion, oral side down and arms outspread, in a shallow dish

filled with 95 % ethanol. Specimens are then transferred

to 70% ethanol for storage. Colors tend to fade upon

preservation, especially if alcohol is used.

Pinnules and cirri are best examined singly; they are

most easily separated from an arm or centrodorsal of dried

specimens by gentle prying with the tip of a fine forceps.

This procedure normally destroys the articular face,

however. In many-armed species with adjacent arms closely apposed, an entire arm or ray may have to be separated

in order to examine the oral and genital pinnules. In taxa

with numerous cirri, many cirri may have to be removed

in order to examine centrodorsal and calyx.

Examination of individual ossicles and articular faces requires maceration in 5% sodium hypochlorite solution

(i.e., commercial liquid bleach). External surfaces of

sutures and articulations are best examined in dried specimens. However, dried (properly preserved) specimens

become brittle; pinnules and arms ofsmaller specimens are

particularly prone to breakage.

Glossary _

Aboral. Away from or opposite to surface bearing mouth

or ambulacrum.

Ambulacral. Toward, on, or associated with ambulacrum-bearing surface of ray, arm, or pinnule.

Ambulacral groove. Furrow in oral surface of tegmen,

arms, and pinnules lined with tracts of cilia and serving

to convey food to mouth.

Ambulacrum. Simple or branched, elongated area on oral

surface of body, extending radially from mouth onto

tegmen, arms, and pinnules, overlying radial water

vascular canal and consisting of groove, marginal lappets, and podia.

Anal tube. Conical or cylindrical projection on oral surface bearing anus at its summit.

Arm. Unbranched, linear series of ossicles arising from

radial ossicle of an unbranched ray, or from distalmost

axillary of a branched ray.

Articular face (or facet). Usually sculptured surface of

an ossicle serving as attachment site for ligaments or

ligaments and muscles that join successive ossicles in a

senes.

Articulation (=Joint). Flexible to nearly immovable

union of successive ossicles in a series.

Axillary ( = Axil). Brachial bearing two distal articulations

and representing the branching point of a ray.

Basal. Any of five interradial plates forming a circlet

aboral to the radial circlet.

Basal ray. Any of five rodlike structures, modified parts

of basal ossicles, lying interradially between radial circlet

and centrodorsal of most comatulids (except Atelecrinidae); often visible as knobs at aboral corners of radials.

Basal tubercle (or knob). The outer tip of a basal ray when

visible externally as a low triangle or rounded swelling

at the aboral, interradial corner of a radial ossicle.

Brachial. Ossicle of an arm or division series.

Calyx. Cuplike central skeleton consisting (in recent species) of basal and radial circlets; reduced in comatulids.

Centrodorsal. Modified uppermost stalk ossicle attached

to aboral surface of comatulids, retained after loss oflarval stalk and commonly bearing cirri.

Cirral. Cirrus ossicle.

Cirrus (pl., cirri). Unbranched, usually hooklike, segmented appendage arising from stalk or centrodorsal;

also refers to branched, segmented "roots" arising from

disal end of stem in Bathycrinidae.

Cirrus socket. Articular face on centrodorsal or columnal for attachment of cirrus.

Column. Unbranched series of ossicles composing stalk

or stem, exclusive of cirri, roots, or holdfast.

Columnal. Column ossicle.

Comb. Modification of distal pinnulars of oral pinnules

producing comblike profile; in Comasteridae and a few

species of Antedonidae.

Crown. In stalked species, the whole crinoid exclusive of

stem.

Disk. Visceral mass that rests on the radial circlet and

arm bases; sometimes its oral surface only (see tegmen).

Distal. Referring to a direction or position away from

the aboral/oral axis, that is, toward tips of rays, arms,

and pinnules; also toward base of stalk.

Distal pinnule. Pinnules distal to genital and oral

pinnules.

Division series ( = Brachitaxis). Series of ossicles following a radial or axillary to and including the next axillary;

series of ossicles between branches on a ray.

Genital pinnule. Gamete-bearing pinnules; usually distal

to one or several pairs of modified oral pinnules.

Interradial. Oriented between rays, that is, between

structures associated with radial water vascular canals.

Ligamentary articulation. Joints bearing only ligaments,

lacking muscle fibers; examples include synarthry and

syzygy.

Muscular articulation. Joint bearing both muscle fibers

and ligaments; opposed articular faces usually characterized by a semicircular aboral ligament fossa, transverse fulcral ridge, axial canal, paired interarticular ligament fossae, and paired muscular fields. In comatulids,

the great majority of arm articulations.

Oral. (adj.) Toward or on the mouth-bearing surface.

(n.) Any offive interradially oriented plates forming circlet surrounding or covering mouth (absent in adult

comatulids).

Oral pinnule. Any proximal pinnule differentiated from

more distal pinnules in structure, function, or both.

OssU:le. Any single calcareous segment of crinoid skeleton

(e.g., columnal, cirral, basal, radial, brachial, pinnular).
8 NOAA Technical Report NMFS 91 _

Pinnular. A pinnule ossicle.

Pinnule. Unbranched, segmented appendage usually

ansmg from alternate sides of successive brachials

(except axillaries and proximal ossicle of pair joined by

syzygy).

Podia (sing.) podion). Tube feet; fine fingerlike projections of water vascular system that line ambulacral

grooves and serve in food capture and respiration.

Proximal. Referring to direction or position toward

aboral/oral axis, that is, toward base of ray, arm, or pinnule; also, toward top of stalk.

Radial. (n.) Most proximal ossicle of a ray, associated

with anyone of five radial water vascular canals that

arise from the circumoral ring canal. (adj.) Associated

or oriented with a ray.

Radial circlet ( = Radial pentagon). Ring of five radials;

sometimes fused into a cone or cup.

Ray. Radial ossicle together with all structures arising

from it.

Rosette. Delicate, perforated structure formed of modified

basal ossicles (excluding basal rods) located within radial

circlet of comatulids (except Atelecrinidae).

Stalk ( = Stem). Series of ossicles attached to aboral surface of crown and serving to anchor crown to and raise

it above substrate; lost in adult comatulids.

Stereom. The fenestrated calcareous material of the

crinoid endoskeleton.

Synarthrial swelling (or tubercle). Rounded or inflated

thickening of aboral surface of ossicles joined by synarthry; lacking in most Comasteridae, present in many

Antedonidae.

Synarthry. Ligamentary articulation in which opposed

articular faces bear an aboral/oral-oriented fulcral

ridge separating two lateral fossae for attachment ofligament bundles; permits limited side-to-side movement of

joint.

Syzygial pair. Two ossicles joined by syzygy; the proximal ossicle lacks a pinnule.

Syzygy. Rigid ligamentary articulation in which ridges

radiating from center of one articular face are apposed

to corresponding ridges on the other face; visible externally as a perforated line; serves as breakage point for

autotomy.

Tegmen. Oral surface of visceral mass bearing mouth,

anus and trunks of ambulacra proximal to free portions

of rays.
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 9

Key to the Crinoidea of the

Continental Margin of Northeastern North America _

la Adult stalked, sessile; basal portion of stalk bears fine, rootlike branches; arms unbranched .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . (Order Bourgueticrinida) 2

lb Adult unstalked, sedentary; aboral centrodorsal bearing segmented, unbranched, usually prehensile cirri;

centrodorsal variously shaped (conical, discoidal, hemispherical or cylindrical) ... (Order Comatulida) 3

2a(la) Basal ossicles completely fused; sutures between basals not visible; sutures between radial ossicles visible

or not; small and delicate; stalk usually no more than 50 mm in length; arms usually five (sometimes

six or seven) in number; known chiefly from the north Atlantic, as far south as Massachusetts (possibly northern Florida) Conocrinus lofotensis (Fig. 4)

,b d

Figure 4

Conocrinus lofotensis. (a through d) Lateral views of upper stalk, calyx, and arm bases

of three specimens. Scales: a, c, d = 2 mm; b = 5 mm. (c and d reprinted with permission from A. M. Clark, 1970).
10 NOAA Technical Report NMFS 91 _

2b(1a) Sutures between basal ossicles usually completely visible; sutures between radial ossicles completely visible; arms five; small and delicate to large and robust; stalk may exceed 200 mm in length; known chiefly

from the Caribbean, as far north as the Blake Plateau Democrinus conifer (Fig. 5)

Figure 5

Democrinus conifer. (a throughf) Lateral

views of upper stalk, calyx, and arm bases

(to the first brachial) of six specimens.

(g) Distal columnal articulation. Scales:

a-f = 2 mm; g = 1 mm.

9

...'y......

",(,,,,:..• ..,

f

b

I

d

3a(1b) Five undivided arms Pentametrocnnus atlanticus (Fig. 6)

3b(1b) Ten arms; each ray with one division series of two ossicles joined by synarthry 4

4a(3b) Basal ossicles form complete thin ring or pentagon usually visible externally as a narrow band completely separating centrodorsal from radial ossicles (but sometimes reduced to low, wide, interradial triangles);

pinnules lacking on proximallO-17 arm segments; centrodorsal almost always taller than wide, conical

or with straight sides near base; cirrus sockets arranged in lO alternating columns; each socket bears

a pair of lateral, often aborally directed tubercles; cirri long slender and rarely retained .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Atelecrinus balanoides (Fig. 7)
_____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 11

c

d

Figure 6

Pentametrocrinus atlanticus. (a) Lateral view

of a specimen showing centrodorsal, bases

of several cirri, bases of three rays, and

interradial portion of tegmen. (b and

c) Lateral views of two centrodorsals with

bases of several cirri. (d) Cirrus. Scales:

left, a-c = 2 mm; right, d = 2 mm.

. . ".' :, .;. \ .. , , ," ~':'" .. . ::.,.. - :: ,.

Figure 7

Atelecrinus balanoides. (a and

b) Lateral views of two specimens showing centrodorsal

and bases of three rays; a:

"balanoides" form; b: "helgae"

form. (c) A pical cirrus.

Scale: 5 mm. (b = basal, r =

radial.)
12 NOAA Technical Report NMFS 91 _

4b(3b) Basal ossicles reduced to an internal rosette with thin radiating rods sometimes visible externally as interradial tubercles; centrodorsal variously shaped; when conical, usually wider than tall; cirrus sockets

without paired lateral tubercles; pinnules present on all or most proximal arm segments 5

5a(4b) Distal segments of proximal one to four pairs ofpinnules each bear a flat, round, triangular, or quadrate

blade; blade-bearing pinnulars, taken together, form a comblike structure 6

5b(4b) Proximal pinnules lacking comblike structure; pinnulars sometimes spinose distally 9

6a(5a) Proximal pinnules all present; PI composed of 25-50 segments 7

6b (5a) Second, third, and fourth pairs of pinnules (pinnules P2- 4, Pb-d) usually absent (P2, Pb, or Pd occasionally present although not all together on the same arm of a specimen); PI of fewer than 25 segments;

whole animal small and delicate with arms no longer than about 30 mm; unknown north of the Blake

Plateau Comatilia iridometriformis (Fig. 8)

Figure 8

Comatilia iridometriformis. (a) Oblique aboral view showing centrodorsal

and bases of cirri, and bases of two rays; (b and c) Centrodorsals (inverted); (d) Cirrus. Scale: 1 mm. (r = radial; 0 = oral surface; p = PI)'
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 13

7a(6a) Centrodorsal conical or hemispheric, almost covered with sockets; cirri slender and weakly curved; division series and first pair of arm segments strongly humped or swollen abo rally (synarthrial tubercles);

combs restricted to first pair ofpinnules (sometimes only PI; rarely also P2), composed of large, round

teeth and occupying all but basal third or fourth of pinnule; fifth to at least eighth brachial with an aboral

spiny knob or keel; mouth central; unknown north of Cape Lookout, North Carolina .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. Comatonia cristata (Fig. 9)

c

Figure 9

Comatonia emtala. (a) Lateral view showing centrodorsal with small apical

and large peripheral cirrus, and bases of three rays showing PI' P2, and

P3; (b) Lateral view showing centrodorsal, bases of cirri and bases of

three rays; (c) Centrodorsal; (d) Side view of arm base (Br4-Br 11 )

showing spinose knobs. Scales: left, a = 5 mm; right, b, c, d = 2 mm..

7b(6a) Centrodorsal a flat disk or pentagon; cirri confined to lateral margins; cirri short, stout and usually curled

terminally; division series and arm bases smoothly rounded, neither humped, swollen or keeled; comb

on first, second, and sometimes third and fourth pinnule pairs, restricted to distal third or fourth of pinnule and composed of 3-11 rounded, triangular or quadrate teeth; comb teeth range from tall and

conspicuous to low and weak; mouth displaced to one side of oral surface (Genus Comactinia) 8
14 NOAA Technical Report NMFS 91 _

8a(7b) Comb usually present on PH; middle comb teeth usually quadrate and often terminally notched;

rounded keel usually present on basal segments of proximal pinnules; aboral distal margin of brachials

lacks spines; color in life reddish flecked with yellow, with yellow pinnules; centrodorsal pale; ossicles

retain pink color after dissociation in 5% sodium hypochlorite (NaGCl) solution; unknown north of

Raccoon Key, South Carolina Comactinia echinoptera (Fig. 10)

,'.-;.'

.,' ,~

Figure 10

Comactinia echinoptera. (a) Aboral view showing centrodorsal and cirrus, and bases

of two rays; (b) Aboral view of small specimen showing centrodorsal and cirrus,

and bases of three rays; (c and d) Cirri; (e) Pi; (f) P2; (g) P3. Scales: upper,

a-d = 5mm; lower, e-g = 2 mm. (r = radial.)
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 15

8b(7b) Comb on Pl-2 (sometimes P3 and P4); teeth rounded or triangular, weak or strong; basal keels absent;

middle segments of proximal pinnules with spiny rims that may develop into strong, spiny flanges; aboral

distal margin of brachials finely spiny; color yellow, red, purple, brown, or a combination; all color lost

upon dissociation in 5% sodium hypochlorite (NaGCI) solution; unknown north of Cape Lookout,

North Carolina Comactinia meridionalis (Fig. 11)

Figure 11

Comactinia meridiona/is. (a) Aboral view showing centrodorsal and cirri, and bases ofthree rays; (b) Aboral

view of small specimen showing centrodorsal and cirrus, and bases of two rays; (c through f) Cirri;

(g) PI; (h) P2; (i) P3. Scales: upper, a-f = 5 mm; lower, g-i = 2 mm.
16 NOAA Technical Report NMFS 91 _

9a(5b) Centrodorsal taller than wide, cylindrical, or truncated conical, with five interradial ridges separating

paired alternating columns of cirri; centrodorsal apex often excavated; division series and arm base ossicles

usually bear fringe of long glassy spines; proximal pinnules compressed, with all segments longer than

wide; unknown north of Blake Plateau Zenometra columnaris (Fig. 12)

Figure 12

Zenometra columnans. (a) Lateral view showing centrodorsal and bases of cirri, and bases of four

rays; (b) Lateral view showing centrodorsal and bases of three rays; (c) Lateral view showing centrodorsal, base of immature cirrus, and bases of three rays (covered with spines); (d)

PI; (e) Cirrus. Scales: upper, a-c, e = 5 mm; lower, d = 5 mm.

9b(5b) Centrodorsal rounded conical or hemispherical, as wide as tall or wider, lacking interradial ridges; cirri

in closely spaced spiral rows (sometimes in columns as well or so crowded that no pattern is visible);

proximal pinnules not markedly compressed and with first two or more segments short or squarish. .. 10

10a(9b) Centrodorsal flattened hemispherical to rounded conical, from 1.5 to more than 2.0 times wider than

tall, with a smooth apical area lacking cirrus sockets; PI (and sometimes one or more subsequent pinnules) very flexible, usually of 35 or more segments, even distal pinnulars short-no more than twice

as long as wide; Pz similar in length to PI (sometimes 3/4 as long as PI> rarely longer) 11

10b(9b) Centrodorsal conical to rounded conical, no more than 1.5 times wider than tall, usually about as wide

as tall, almost completely covered with sockets or with spiny or papillose apex; PI rather stiff, rarely

with more than 35 segments; middle and distal pinnulars thin, elongated and up to several times longer

than wide, usually with expanded spiny distal ends; PI usually 2-4 times longer than Pz 12
___________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 17

lla(10a) Very large; arms often exceeding 200 mm in length and 3 mm in basal width; peripheral cirri up to

70 mm long, usually with 44-55 cirrals none of which are more than twice as long as their narrowest

width; PH similar, of many short segments; PI usually 15-25 mm long, of 42-54 (up to 82) segments;

unknown south of Georges Bank, Massachusetts Heliometra glacialis (Fig. 13)

Figure 13

Heliomelraglai:ialis. Lateral view showing centrodorsal, small apical and large peripheral

cirrus, and bases of three rays showing PI'

P2, and P3. Scale: 5 mm.

llb(10a) Much smaller; maximum arm length 75 mm and basal width 1.4 mm; peripheral cirri rarely more

than 15 mm long, usually of 15-20 segments; longest cirrals 2.5-3.0 times longer than their narrowest

width; PI usually 8-10 mm long, of 30-43 short segments; P2-3 of far fewer, longer segments (P2

may occasionally resemble PI); unknown north of Cape Lookout, North Carolina .

. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Coccometra hagenii (Fig. 14)

12a(10b) Ossicles of division series and arm bases laterally separated and not laterally thickened or flattened,

distal aboral margins not everted and spiny; PI two to four times longer than P2; longest cirrals three

to five times longer than their narrowest width . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .. 13

12b(10b) Ossicles of division series and arm bases laterally thickened, flattened and closely appressed; PI no

more than about twice as long as P2; longest cirrals no more than three and a half times longer than

their narrowest width (Trichometra cubensis) 14
18 NOAA Technical Report NMFS 91 _

Figure 14

Coccometra hagenii. (a) Lateral view showing centrodorsal, small apical and large peripheral cirrus, and bases of three rays showing PI' P2, and P3 (P2 and P3 bear gonads); (b and c) Lateral

views showing centrodorsal and bases of rays; (d) Centrodorsal. Scales: upper, a = 5 mm;

lower, b-d = 2 mm.

13a(12a)

13b(12a)

Large cirrus sockets arranged in distinct vertical columns as well as spirally; cirri usually 30-60 mm

long and composed of 30-50 segments; arms usually with four muscular articulations between successive syzygies; PI less than half the length of peripheral cirrus, with 5-7 short basal segments;

unknown south of Greenland Poliometra prolixa (Fig. 15)

Small cirrus sockets in crowded spiral rows (columns, if present, indistinct); cirri usually 25-30 mm

long and composed of 20-30 (maximum 33) cirrals; arms usually with three muscular articulations

between successive syzygies following Brg + 10; Plat least half as long as peripheral cirrus, with

3-5 short basal segments Hathrometra tenella (Fig. 16)
_____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 19

Figure 15

Poliometra prolixa. Lateral view showing centrodorsal. small apical and

large peripheral cirrus, and bases of three rays showing PI' Pz, and P3 .

Scale: 5 mm.

Figure 16

Hathromitra tmelUl. (a) Lateral view showing centrodorsal, small apical and large

peripheral cirrus, and bases of three rays showing PI. pz• and P3; (b and

c) Lateral views showing centrodorsal and bases of three rays. Scales: upper,

b-c = 2 mm; lower, a = 5 mm.
20 NOAA Technical Report NMFS 91 _

14a(12b) Longest cirrus segments mostly 2.0-2.5 (maximum 3.0) times longer than their narrowest width; distal

cirrus segments as long as or shorter than wide; PI 3-6 mm long and composed of 11-15 segments;

unknown north of the Blake Plateau Trichometra cubensis (sensu stricto) (Fig. 17)

Figure 17

Trichometra cubensis (sensu stricto). Lateral view showing centrodorsal, small apical and

large peripheral cirrus, and bases of three rays showing PI' P2 , and P3 . Scale: 5 mm.
____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 21

14b(12b) Longest cirrals mostly 3.0-3.5 (minimum 2.6) times longer than their narrowest width; distal cirrals

usually slightly longer than wide; PI usually 7-11 mm long and composed of 16-23 segments; as far

south as New England Triclwmetra cubensis (northern form) (Fig. 18)

Figure 18

Trichomara cubmsis (northern fonn). (a) Lateral view showing centrodorsal, cirrus bases,

and bases of three rays; (b) Lateral view showing centrodorsal, small apical and large

peripheral cirrus, and bases of three rays showing PI' P2, and P3. (c) The same of

another specimen (lacking small apical cirrus). Scales: a = 2 mm; b-c = 5 mm.
22 NOAA Technical Report NMFS 91 _

Annotated Systematic Checklist _

This list follows the higher classification proposed by

Rasmussen and Sieverts-Doreck (1978). Genera are arranged alphabetically within families. A new classification

of the subclass Articulata has recently been proposed by

Simms (1988). The only change that directly affects familial

level taxa and below included here is the removal of the

Atelecrinidae from the Paracomatulacea (probably returning it to the Antedonacea although the author does not

specify). Geographic and bathymetric ranges include all

published records plus data from additional specimens examined at the National Museum of Natural History

(N.M.N.H.), Smithsonian Institution; Museum of Comparative Zoology, Harvard University; and Newfoundland

Museum, St. John's, Newfoundland. Temperature and

substrate data are from Clark and Clark (1967) and Clark

(1970) except where indicated.

Class CRINOIDEA

Subclass ARTICULATA

Order BOURGUETICRINIDA _

Family BATHYCRINIDAE *

Conocrinus lofotensis M. Sars, 1868-Norwegian coast

from lat. 58 to 700 N (possibly as far east as the Siberian

Arctic, see Clark 1970); west of Ireland; south and west

of the Faeroe Islands; between southeastern Greenland and

Iceland; Davis Straits southward to Massachusetts; possibly

off northern Florida. 140-3475 m.

Color-Crown yellow or grayish.

Habitat-Unconsolidated substrates: clay, mud, or

combinations of mud and sand.

Remarks-This small sea lily has a stalk length to about

75 mm (usually to 50 mm) and a crown height (base of

calyx to arm tips) to about 22 mm (usually 12-15 mm).

Specimens from American waters were originally recognized as a distinct species (Rhizocrinus verrilli A. H. Clark),

then reduced to varietal status as an extreme form of a continuum with R. lojotensis (Clark 1923). Gislen (1938)

thought any distinction between the two doubtful and the

two have since been treated as synonymous. Roux (1977)

recognized the close similarity among the genera Rhizocrinus, Conocrinus, and Democrinus and placed R. lojotensis

in the genus Conocrinus (Roux 1985). These three genera

have been partly distinguished by calyx characteristics:

-------------------

'Roux (1985) transferred Conocrinus and Democrinus (and Rhizocrinus) to

the family Bourgueticrinidae but cited no reasons for the change.

Rhizocrinus and Conocrinus supposedly lack interbasal (and

often interradial) sutures while Democrinus exhibits sutures

between all calyx ossicles. However, Roux (1985) described

C. cabiochi Roux with partial interbasal sutures and D. parjaiti (Perrier) with both interradial and interbasal sutures

complete, irregular, partial, or not visible. He suggested

that, while the three genera may not be synonymous,

characters other than those offered by the calyx should be

used to distinguish them.

The distribution of C. lojotensis along the east coast of

the United States is uncertain. No records exist between

northern Florida and Massachusetts. Because the Caribbean species Democrinus conifer (A. H. Clark) has been

recorded from the Blake Plateau (see below), occasionally

lacks interbasal sutures, and has calyx proportions overlapping those of C. lojotensis, the identity of the small, poorly

preserved material of C. lojotensis from northern Florida remains unclear. The wide bathymetric range attributed to

this species may also be due to misidentifications.

Democrinus conifer (A. H. Clark, 1909)-B1ake Plateau,

Bahama Islands and Gulf of Mexico south along the coast

of Central and South America, and the West Indies, to

Brazil. 170-1750 m (chiefly 200-800 m). Most records

are from the Straits of Florida south; the species was collected at a single R/V Pillsbury station off Jacksonville,

Florida, on the Blake Plateau in 388-403 m (Meyer et al.

1978).

Color-Yellow or yellow orange with yellow arms.

Habitat-Unconsolidated substrates; mud or muddy

sand.

Remarks-As mentioned under Conocrinus lojotensis, the

distinction between the genera Democrinus, Conocrinus, and

Rhizocrinus is unclear and the material of C. lojotensis from

northern Florida may really represent this species.

Calyx proportions and stalk length are variable in D.

conifer and the taxon may require subdivision. However,

preliminary examination of a large number of specimens

of this species and the sympatric D. brevis (A. H. Clark)

and D. robustus (A. H. Clark) reveals no consistent differences between the three and they are here considered

as synonymous.

Democrinus conifer reaches a much larger size than C.

lojotensis; stalk length may exceed 280 mm and crown height

reaches at least 78 mm. Large specimens sometimes appear to have ceased growing: newly developed thin proximal columnals may be absent so the topmost columnals

are already squarish, and the calyx appears to be overgrown with secondary stereom that obscures interbasal

sutures. The ultrastructure of the stalk was treated by

Grimmer et al. (1984).
___________ Messing and Dearborn: Echinodermata, Crinoidea of the Nor~heastern United States 23

Order COMATULIDA

Superfamily PARACOMATULACEA --

Family ATELECRINIDAE

Atelecrinus balanoides Carpenter, 1881-West ofIreland

and southwest of the Faeroe Islands; Blake Plateau and

Gulf of Mexico to Recife, Brazil. 550 to 1530 m. Specimens

that may represent an undescribed species (see below) may

have been collected as shallow as 178 m.

Color-White in preserved specimens.

Habitat-Unconsolidated substrates; mud or pelagic

ooze. Photographs and observations made from submersibles [tentative identification, Messing (1985)] indicate

bottoms apparently undisturbed by significant horizontal

currents. Long delicate cirri (almost never retained in

dredged material) act as outspread props; arms extend

radially parallel to and just above the sediment with tips

slightly upcurved. Usually dredged singly or in small

numbers.

Remarks-Two bathymetrically allopatric forms exist.

Most specimens collected shallower than 800 m have the

following suite of characteristics: centrodorsal parallel-sided

proximally, with weak interradial ridges and socket

tubercles; basals slightly swollen interradially, if at all; profile of the external part of the radials forming an angle of

40-50° with a plane parallel to the centrodorsal base (normal to the oral/aboral axis); IErl distolaterally cut away;

axils rhombic or hexagonal with lateral margins converging proxi~ally, not laterally thickened; adjacent axils well

separated; synarthrial swellings weak or absent and, most

consistently, Brl and Br2 separated and rounded exteriorly (Fig. 7a: "balanoides" form).

Specimens collected primarily below 1000 m differ in the

following respects: centrodorsal usually tapering from the

base, with more prominent interradial ridges and socket

tubercles; basals swollen interradially; radial/centrodorsal

angle usually 30° or less; IBrl sometimes thickened distolaterally; axils hexagonal, with lateral margins parallel,

thickened and ridge- or knob-like; adjacent axils usually

in contact; synarthrial swellings moderate; Brl and Br2

typically apposed, thickened and expanded or flattened exteriorly (Fig. 7b: "helgae" form).

The shallower-water form, including the type material

of A. balanoides, has been collected between 800 and 900

m at a few stations in the Caribbean Sea; a few specimens

of the deeper-water form have been collected in this depth

range only on the Blake Plateau and off Brazil. The type

of A. helgae, collected in 700 m off Ireland and synonymized with A. balanoides by A. M. Clark (1970), approaches

the deep-water form in having some axils laterally thickened and Brl somewhat flattened exteriorly. Because most

of the distinctions described above are not uniformly clearcut relative to depth, we herein treat the two forms as infrasubspecific variants.

Several specimens collected from the Straits of Florida

differ in several important respects and may represent an

undescribed species. They bear up to 75 cirri in 15 closely

crowded columns rather than ten, with much more prominent socket tubercles. None have been reported from the

area covered by this paper.

Superfamily COMASTERACEA _

Family COMASTERIDAE

Comactinia echinoptera (Muller, 1840)-South Carolina;

southeastern Florida; Bahama Islands and West Indies;

Caribbean coast of Central and South America from the

Yucatan Channel to Cabo Frio, Brazil (and, perhaps, to

Isla de los Alcatraces off Sao Paulo). Found 2 to at least

92 m [possibly to 183 m; older deeper records (Clark 1931)

are probably inaccurate]. Only one specimen has been collected in the area covered by this paper: off Raccoon Key,

South Carolina.

Color-Centrodorsal and cirri white; radials, division

series and arms reddish flecked with yellow; pinnules

yellow. 111 preserved material, radials, division series and

arms range from deep pink to very pale lavender; pinnules,

ambulacra and disk tan or white. Ossicles retain lavender

color after dissociation in sodium hypochlorite solution.

Habitat-Widespread but uncommon on reefs in the

West Indies. It remains cryptic in crevices and under

corals during the day and extends its longer arms to feed

at night; the disk is rarely exposed. Arms form a monoplanar filtration fan normal to water movement with ambulacra facing one direction only (Meyer 1973; Macurda

1973, 1975).

Remarks-Large C. echinoptera develop significantly

asymmetrical rays. Arms arising from the three rays closest

to the mouth (located excentrically to one side of the disk)

tend to be longer than arms arising from the two remaining rays. Ray asymmetry sometimes appears, albeit not

as strongly, in C. meridionalis. See Messing (1978a) for a

complete description.

Comactinia meridionalis (L. Agassiz, 1865)-Cape Lookout, North Carolina to Surinam (possibly to Sao Paulo,

Brazil), including the Bahama Islands, West Indies, and

Gulf of Mexico. Found 3 to at least 373 m (possibly to 549

m). Localized, patchy shallow-water populations exist

along the southeastern United States from the Carolinas

to central Florida and off Central America, but the species

seems to be limited to greater depths (>50 m) off southeastern Florida, the Bahamas, Gulf of Mexico, and West

Indies.

Color-Yellow, sometimes with red (A. H. Clark uses

"carmine," a vivid red) pinnules, often yellow-tipped; red,

sometimes with yellow spots and orange to yellow pinnules
24 NOAA Technical Report NMFS 91 _

(sometimes red- or yellow-spotted or yellow-tipped); purple, sometimes with yellow- and purple-banded pinnules

(after A. H. Clark, 1921). This list may include specimens

referrable to C. echinoptera as A. H. Clark did not distinguish

the two species of Comactinia in his monograph. Shallowwater specimens from the east coast of the United States

have brown arms with narrow yellow bands, brown- and

yellow-beaded pinnules, and the disk brown with yellow

spots (from a photograph courtesy ]. Miller, Harbor

Branch Oceanographic Institute, Fort Pierce, FL).

Habitat-Rough hard bottoms and reefs, in rock

crevices, sponges, or the undersides of corals (Meyer 1973;

Messing 1985 and unpubl. observ.;]. Miller, pers. commun., Dec. 1988).

Remarks-Comactinia meridionalis exists in three reasonably distinguishable morphological forms. Size generally

increases with increasing depth (Clark 1931; Messing

1978a). The shallow-water form (3-25 m; Carolinas to

Central Florida and off Central America) is small; arm

length usually does not exceed 45-50 mm. Specimens from

the southeastern U.S. usually have more than XX cirri

of 10-14 cirrals each. The cirri are more slender, longer

in relation to centrodorsal diameter, and have their longest

cirrals more elongated than in deeper water specimens.

PI through P3 bear relatively strong combs. This is the

true meridionalis form corresponding to the type series. The

larger, deeper water forms may represent a distinct species.

The second form occurs chiefly in 50-200 m in the Straits

of Florida, Gulf of Mexico, and Caribbean Sea. Arm length

reaches about 100 mm; cirri are fewer (usually XVXVIII), stouter, usually with 9-11 cirrals. Combs appear

on PI and Pz. A third, larger robust variant, C. m.

hartlaubi Messing, has a similar distribution (although absent from southeastern Florida) but is characteristic of

greater depths (200-400 m). Arms probably reach 250 mm

in length; robust cirri usually number XX-XXX, with

11-14 short cirrals. PI through P3 (sometimes also Pt )

bear weak combs. See Messing (1978a) for a complete

discussion of intraspecific variation.

Shallow-water specimens extend arms for feeding both

day and night. Arms form a monoplanar filtration fan

normal to wave oscillation with ambulacra facing one

direction only. Meyer (1973) found specimens completely

exposed on shallow reefs off Panama.

Comatilia iridometriformis A. H. Clark, 1909-Blake

Plateau; northern Straits of Florida; Northwest Providence

Channel, Bahamas. 256-686 m.

Color-Yellow.

Habitat-The following information is abstracted from

Messing (1984, and references therein). This species is

almost invariably collected with arborescent, azooxanthellate, bank-forming, scleractinian corals (e.g., Lophelia

prolijera, Enallopsammia profunda) that contribute to dense

thickets and extensive banks in the northeastern Straits of

Florida, Blake Plateau and northeastern Gulf of Mexico.

It occurs in dense (+ 15 individuals/0.25 mZ) patchy

aggregations on dead coral, coral rubble, and low carbonate crusts adjacent to living coral thickets on elongated

carbonate mounds (lithoherms) in the Straits of Florida.

No specimens were observed clinging to living coral. Arms

extend in a planar or shallow bowllike radial flltration fan.

Remarks-This species exhibits paedomorphosis, the

retention of typically juvenile characteristics in an adult.

Paedomorphic features include small size (maximum arm

length "'30 mm), large centrodorsal cavity, large radial

ossicles, apparently delayed metamorphosis of basals from

a complete circlet into a rosette, incomplete pinnulation

(i.e., the last few pinnules that normally develop in comatulids are absent in this species), extreme elongation of

cirrals (relative to other Comasteridae), central mouth,

coarse stereomic (skeletal) microstructure, and elongation

of brachials. Messing (1984) suggested that paedomorphosis in C. iridometrijormis is the result of progenesis [sensu

Gould (1977)], that is, accelerated sexual maturation with

retention of ancestral juvenile form.

Comatilia iridometrijormis is the only internally brooding

crinoid not found in middle or high southern latitudes. One

or two of the most proximal genital pinnules (P5, Pe , or

P6) may bear an enormously swollen gonad containing a

single giant embryo or larva (1.4-1.6 mm long) that appears to lie free in the ovarian lumen. Advanced unreleased

larvae have already developed a stalk, attachment plate,

and calyx ossicles (all still embedded in the larva's globular

body). Pentacrinoid postlarvae attached to cirri of adults

and no larger than unreleased larvae, together with the

species limited geographic range, suggest that a planktonic

stage is absent or very brief.

Pz, P3, Pt , Pb, Pc, and Pd are normally absent in adult

specimens. Pz, Pt , and Pd occasionally occur, although

neither all together on a single arm nor uniformly on all

arms of an individual.

Superfamily ANTEDONACEA _

Family ANTEDONIDAE

Coccometra hagenii (Pourtales, 1867)-Cape Lookout,

North Carolina; Blake Plateau; Straits of Florida; Campeche Bank and Yucatan Channel. 14-1046 m. An extremely abundant species on the Pourtales Terrace south

of the Florida Keys, chiefly in 150-250 m; 567 specimens

were collected in a single trawl at R/V Gerda station 1102

(Meyer et al. 1978). Only three specimens have been collected within the area covered by this paper: off Cape

Lookout, North Carolina in 14 m (the shallowest record)

and on the Blake Plateau in 805 and 1046 m (the deepest

records).

Color-Pale greenish; young specimens with dark

brown spots paired on arm bases but becoming confluent
___________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 25

distally; spots hardly apparent in adults (Pourtales 1868,

1878); brownish (Nutting 1895, cited in Clark and Clark

1967). Some preserved specimens have strongly marked

disks.

Habitat-Hard irregular substrates.

Remarks-Examination of extensive material collected

by the University of Miami's R/V Gerda in the Straits of

Florida indicates a slightly wider range of morphometric

features than given in the most recent description of this

species (Clark and Clark 1967), as follows: maximum centrodorsal diameter 3.1 mm (most specimens 2.2-2.8 mm);

cirri XXVII-LXVIII, 8-25 (usually XLIII-LVIII, 15-

19); PI of up to 43 segments; P2 normally shorter and stiffer than PI with up to 21 segments and bearing a gonad;

P3 occasionally longer than P2. Rarely, P2 may lack a

gonad and resemble PI'

Dredged specimens typically retain their arms but lose

their cirri.

Comatonia cristata (Hartlaub, 1912)-Cape Lookout,

North Carolina; Straits of Florida; Yucatan Channel.

14-580 m. All but three specimens were collected on

the Pourtales Terrace south ofthe Florida Keys in association with Coccometra hagenii, chiefly between 150 and 300

m. The only specimen known from the area covered by

this paper was taken off Cape Lookout, North Carolina

in 14 m.

Color-Yellow.

Habitat-Hard, irregular substrates. A single specimen

was collected with coral rubble derived from arborescent,

azooxanthellate, thicket-forming coral (Lophelia prolifera) in

the northeastern Straits of Florida (Messing, unpubl.).

Remarks-A. H. Clark (1916, 1931) placed C. cristata

in the Comasteridae on the basis of well-developed combs

on the first pair of pinnules (and occasionally also P2). All

other morphological features (e.g., centrodorsal, division

series, arms) indicate antedonid affinities. Messing (1981)

pointed out that the comasteridlike combs are likely

modifications of comb rudiments found in several antedonid genera and followed Gislen (1924) and Meyer (1972)

in including the species in the Antedonidae.

Dredged specimens typically retain some cirri but lose

their arms beyond the first or second syzygies.

Hathrometra tenella (Retzius, 1783)-Scandinavia from

Denmark to northern Norway; southwest oflreland; Shetland Islands westward along the Faeroe Ridge to Iceland;

southern Greenland; Davis Straits southward to offChesapeake Bay. 28-1783 m (46-891 m off North America;

tending to shoal to the north; off New England chiefly in

220-450 m). 2.0-13.9°C.

Color-Dark green dotted with white; cirri white; disk

grayish (Clark 1921).

Habitat-Chiefly uncosolidated substrates: mud or mud

and sand, sometimes in great abundance; Verrill (1882)

reported over 10,000 specimens from a single dredge haul

off Martha's Vineyard, Massachusetts.

Remarks-The distinction between H. tenella and H. sarsi (Diiben & Koren, 1846) seems arbitrary. The latter,

known chiefly from Iceland to Scandinavia, has been distinguished as smaller, with shorter cirri offewer segments.

Cirri of H. sarsi supposedly reach 17 mm long with up to

24 segments (Clark and Clark 1967; Clark 1970) while H.

tenella bears cirri 25-35 mm long with 24-33 segments

(Clark and Clark 1967). However, meristic and morphometric measurements for the two forms overlap widely and maximum reported arm lengths differ by only 10

mm (i.e., 100 vs. 110 mm). Re-examination ofspecimens

collected off eastern Canada and New England (N.M. N. H.

11540, 11541) suggests a continuum; smaller specimens,

identified by A. H. Clark as H. sarsi, bear cirri 11-12 mm

long with 16-20 segments; larger specimens, identified by

him as H. tenella, bear cirri 16-26 mm long with 14-30

segments. In one case, Clark identified both forms in the

same trawl (Albatross stn. 2429).

Bathymetric and temperature ranges given above include

records for H. sarsi from the northeastern Atlantic.

Heliometraglacialis (Owen, 1833)-Wrangel Island (Arctic Ocean) westward to Spitzbergen and northern Norway

(Finmark); off Norway (lat. 62-63°N) westward along the

Faeroe Ridge to Iceland; Greenland, Hudson Bay,

Labrador southward to offCape Cod, Massachusetts; also

in the North Pacific from the Okhotsk Sea to the Korean

Straits (Clark 1970). 14-1358 m. - 1.9 to + 5.8°C (Clark

1970).

Color-Bright yellow, sometimes tinged with purple

(Clark 1970).

Habitat-A variety ofsubstrates including mud, sand,

gravel, and stones; most frequently on gravel/sand and

mud.

Remarks-The largest northern, cold-water comatulid.

Arm length reaches 265 mm (350 mm in individuals from

the northern Pacific); cirri to 90 mm (usually 60-70 mm).

Poliometra prolixa (Sladen, 1881)-East of the Taimyr

Peninsula westward to Novaya Zemlya; Spitzbergen; Norway north of lat. 62-63°N westward along the Faeroe

Ridge to northern Iceland; Greenland; Baffin Bay and

Davis Straits. 20-1960 m. - 1.97 to + 2.0°C.

Color-Arms brown to yellow; pinnules red to yellow;

cirri white (Clark 1970).

Habitat-Chiefly soft mud or clay; sometimes on mud

with sand or gravel.

Remarks-This species reaches North America only off

Baffin and Ellesmere Islands in the Canadian Arctic.

Trichometra cubensis (Pourtales, 1869)-Morocco northward to the Faeroe Islands; south oflceland; Davis Straits

southward to Nova Scotia; southern Blake Plateau(?),
26 NOAA Technical Report NMFS 91 _

northeastern Gulf of Mexico and Bahama Islands southward to St. Vincent and Santa Marta, Colombia. 210-2432

m. 2.4-1O°C. Shallower water material ("'200 m) was collected chiefly off Arrowsmith Bank in the Yucatan Channel where strong northward geostrophic flow tilts isotherms

upward to the west, bringing colder water into shallower

depths along the Mexican side of the channel.

Color-Pale yellowish white.

Habitat-Often found clinging to arborescent cnidarians with narrow branches (e.g., antipatharians, some

alcyonarians).

Remarks-In the northwestern Atlantic, two allopatric

forms may represent distinct species. The southern form

(including the type) occurs from the Bahamas and perhaps

southern Blake Plateau southward and is included here for

comparison. Cirrals are proportionately shorter (LW chiefly 2.0-2.5) and PI is shorter (3-6 mm) with fewer segments (11-15). The northern form is found from Nova

Scotia northward and eastward. Cirrals are proportionately

more elongated (LW chiefly 3.0-3.5) and P j is longer

(7-11 mm) with more segments (16-23). The continuum

in PI length and number of segments between the two

forms is deceiving; specimens bearing minimum northern

PI-values are much smaller than specimens bearing maximum southern PI-values.

We have not separated the two as distinct species because

material identified as Trichometra cubensis from the northeastern Atlantic (Clark 1980) spans much of the range of

variation found in both. Direct comparison of western and

eastern Atlantic material has yet to be made.

Numerous small specimens identified as Trichometra

cubensis collected on the Blake Plateau by the U. S. Fisheries

Commission Steamer Albatross (e.g., N.M.N.H. 14700,

14707, 34608) represent an undescribed antedonid most

easily distinguished by its more flexible pinnules composed

ofshorter, more numerous pinnulars with expanded spiny

distal ends, and by Br2 with a strong acute proximal

margm.

Zenometra columnaris (Carpenter, 1881)-B1ake Plateau;

northern Straits of Florida; northern Caribbean Sea (southwest of Haiti). A single record, originally catalogued as

fragmentary but currently unidentifiable, exists from the

Gulf of Mexico. 504 to 1034 m, chiefly 750-800 m. The

Gulf of Mexico fragment was collected in 308 m.

Color-Unknown.

Remarks-The centrodorsal plate is truncated conical

with weak interradial ridges in smaller specimens; in large

specimens it becomes a five-sided cylinder with strong

interradial ridges and a flat or excavated aboral pole.

Family PENTAMETROCRINIDAE

Pentametrocrinus at/anticus (Perrier, 1883)-Blake

Plateau; Martinique; Morocco; the Canary Islands and

Azores north through the Bay of Biscay to Porcupine Bank

off Ireland. 374-2115 m. In the western Atlantic, the

possible depth range is 374-779 m with four of the five

records shallower than 650 m. In the eastern Atlantic, the

depth range is 650-2115 m with ten of the twelve records

deeper than 1000 m.

Color-Preserved material white or pale tan with brown

tegmen.

Habitat-Probably hard substrates dominated by

sponges, a1cyonarians, and azooxanthellate colonial scleractinians.

Remarks-This is the only five-armed species in the

north Atlantic Ocean. Its morphology, distribution and

relation to congeners were reviewed by Messing (1978b).

Species That May Occur Within the Area

Covered by this Manual But Not Yet Reported _

Order MILLERICRINIDA _

Family HYOCRINIDAE

Hyocrinus (Gephyrocrinus) grimaldii Koehler & Bather,

1902-Reported by Haedrich and Maunder (1984) in

1850-1875 m from Carson Canyon, Newfoundland. Reported elsewhere from the equatorial mid-Atlantic to the

Canary Islands and Madeira.

Order BOURGUETICRINIDA ----_

far west as eastern Greenland, and the Arctic Ocean

as far east as the Laptev Sea, in 1360-2815 m (Clark 1970).

Order COMATULIDA _

Family ANTEDONIDAE

Antedon petasus (Diiben & Koren, 1846)-Chiefly restricted to the coast of Norway in 10 to 326 m; a single

specimen has been taken off southwestern Iceland (Clark

1970).

Family BATHYCRINIDAE

Bathycrinus carpenteri (Danielssen & Koren, 1877)-

Recorded from the deep basin of the Norwegian Sea as

Thaumatometra septentrionalis A.

Known from Baffin Bay to Iceland.

to 3.0°C (Clark and Clark 1967).

H. Clark, 1918-

2075-2623 m. 1.5
_____________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 27

Thaumatocrinusjungerseni A. H. Clark, 1915-Collected

only between Iceland and Greenland. 823-2075 m (Clark

and Clark 1967).

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28 NOAA Technical Report NMFS 91 _

Systematic Index _

Antedon petasus . . . . . . . . . . . 26

Atelecrinus . . . . . . . . . . . . . . . . 5

Atelecrinus balanoides .. 5, 10, II, 23

Atelecrinus balanoides" balanoides" form. . . . . . . . . . . . . . II, 23

Atelecrinus balanoides" helgae" form. . . . . . . . . . II, 23

Atelecrinus helgae . . . . 23

Bathycrinus carpenteri . . . . . . . . . . . . . . . . 26

Coccometra hagenii. . . 5, 17, 18, 24

Comactinia 13

Comactinia echinoptera. . .. 5, 14, 23, 24

Comactinia meridionalis. 5, 15, 23, 24

Comactinia meridionalis hart/aubi. . 24

Comatilia iridometriformis . . . . . . . . . .... . . . . . 5, 6, 12, 24

Comatonia cristata . . . . . . . . . . . . . . . . . . . . . . . 5, 13, 25

Conocrinus . . . . . . . . . . . . . 22

Conocrinus cabiochi . . . . . . . . . . . . . . . . . . . 22

Conocrinus lofotensis. . . .. 5, 9, 22

Democrinus . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 22

Democrinus brevis 22

Democrinus conifer. . . . . .... . . .... . . .. . . .. . . .... .. . ... 5, 10, 22

Democrinus parfaiti . . ... 22

Democrinus robustus. . . . . . . . . . . . . . . . . . . . 22

Hathrometra sarsi. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 25

Hathrometra tenella . . . . . . . . . . 5, 18, 19, 25

Heliometra glacialis .. . . . . . . . . . . . . . . . . . 5, 17, 25

Hyocrinus (Gephyrocrinus) grimaldii 26

Pentametrocrinus at/anticus. . . . . . . . . . . . . . . . . . . . . . . . . . 5, 10, II, 26

Poliometra prolixa . . . . . . . . . . . . . . . . . . . . . . . . . . . . 5, 18, 19, 25

Rhizocrinus . . . .... 22

Rhizocrinus lofotensis . . . . . . . . . . . .. 22

Rhizocrinus verrilli 22

Thaumatocrinus jungerseni . . . . . . . . . . 27

Thaumatometra septentrionalis . . . . . . . . . . . . . . • . . 26

Trichometra cubensis 5, 17, 20, 21, 25, 26

Zenometra columnaris . . . . . . . . . . . . . . . . . . . . . . . . 5, 16, 26

Acknowledgments _

Preparation of this manual was supported in part by a grant

from the National Science Foundation to the Editorial

Board of the "Marine Flora and Fauna of the Eastern

United States."

The authors wish to express their thanks to the following individuals. Frederick M. Bayer (Smithsonian Institution) and the late Gilbert L. Voss (University of Miami)

provided access to specimens collected by R/V Pillsbury and

Gerda. Figures 6-12 and 14 are drawn from this material,

now housed in the National Museum of Natural History.

Access to additional specimens was provided by David L.

Pawson (Smithsonian Institution); Robert Woollacott and

Herbert Levi (Museum of Comparative Zoology, Harvard); Richard Haedrich (Newfoundland Museum); Linda

Pequegnat (Texas A&M University); and A. Conrad Neumann (University of North Carolina, Chapel Hill). Ailsa

M. Clark kindly granted permission to reproduce Figures

4, c and d, previously published in Clark (1970). Figure

6 was previously published in Messing (1978b), Figures

9, b to d, in Messing (1981), and Figures 10 and 11 in

Messing (1978a). Figures 2,8,9, b to d, 10, and 11 were

originally produced for the senior author's master's thesis

(unpubl., University of Miami). The manuscript benefitted

from reviews by David L. Meyer (University of Cincinnati), D. L. Pawson and M. R. Carriker (University of

Delaware).

Coordinating Editor _

Melbourne R. Carriker, College of Marine Studies,

University of Delaware, Lewes, DE 19958.

Editorial Advisers _

A. Ralph Cavaliere, Department of Biology, Gettysburg

College, Gettysburg, PA 17325.

Arthur G. Humes, Boston University Marine Program,

Marine Biological Laboratory, Woods Hole, MA 02543.

David L. Pawson, Department of Invertebrate Zoology,

National Museum of Natural History, Smithsonian Institution, Washington, D.C. 20560.

Kenneth P. Sebens, Marine Science and Maritime Studies

Center, Northeastern University, East Point, Nahant,

MA 01908.

Ruth D. Turner, Museum ofComparative Zoology, Harvard University, Cambridge, MA 02138.

Robert T. Wike, Department of Botany, University of

Massachusetts, Amherst, MA 01002.

In addition to establishing the format for the "Marine

Flora and Fauna of the Eastern United States," the Board

invites systematists to collaborate in the preparation of

manuals, reviews manuscripts, and advises the Scientific

Editor of the National Marine Fisheries Service.

Coordinating Editor's Comments _

Publication of the "Marine Flora and Fauna ofthe Eastern

United States" is most timely in view of the growing universal emphasis on work in the marine environment and

the critical need for precise and complete identification of

organisms related to this work. It is essential, if at all possible, that organisms be identified accurately to species. Accurate scientific names of plants and animals unlock the

great quantities of biological information stored in libraries,

obviate duplication of research already done, and often

make possible prediction of attributes of organisms that

have been inadequately studied.

Charles G. Messing first became fascinated with marine

invertebrates while an undergraduate at Rutgers U niversity from which he graduated in 1970. His professional fate

was decided when he was introduced to the invertebrate
___________ Messing and Dearborn: Echinodermata, Crinoidea of the Northeastern United States 29

collections at the University of Miami's Rosenstiel School

of Marine and Atmospheric Science and the Smithsonian

Institution during his first year of graduate study at the

former. At the suggestion of Frederick M. Bayer in 1972,

he began working on crinoids and has continued to do so

since, despite occasionally straying to dabble in the biology

oftanaidacean crustaceans. He received master's and doctorate degrees from the University of Miami and was subsequently awarded a Smithsonian post-doctoral fellowship

to study crinoids at the National Museum of Natural

History. Several years followed as a lecturer at the U niversity of Miami (including a year as acting coordinator of

the Undergraduate Marine Science Program) and as a freelance consultant and scientific illustrator. He was appointed

assistant professor at Nova University's Oceanographic

Center in 1990.

John H. Dearborn received his undergraduate training

in Zoology at the University of New Hampshire and completed his M.S. degree at Michigan State University. His

professional interests in marine invertebrates developed

during summer employment with the U.S. Fish and Wildlife Service at Boothbay Harbor, Maine and field work for

the Alaska Department of Fish and Game on Afognak

Island. In 1958 he began doctoral studies at Stanford

University and was presented with a unique opportunity

to conduct field work on the ecology ofmarine invertebrates

at McMurdo Sound, Antarctica. These early studies led

to a lifetime passion for polar ecology, especially the biology of invertebrates ofthe Antarctic Shelf and crinoids and

ophiuroids in particular. After completion of the Ph.D.

degree he was awarded a National Science Foundation

Postdoctoral Fellowship for echinoderm studies at the

Museum of Comparative Zoology at Harvard. He joined

the faculty at the University of Maine in 1966 and has been

a Professor of Zoology there since 1976.
30 NOAA Technical Report NMFS 91 _

Published manuals

Marine Flora and Fauna of the Northeastern United States

Annelida: Oligochaeta Cook, David G., and Ralph O. Brinkhurst

Protozoa: Ciliophora Borror, Arthur C.

Higher Plants of the Marine Fringe Moul, Edwin T.

Pycnogonida McCloskey, Lawrence R.

Crustacea: Stomatopoda Manning, Raymond B.

Crustacea: Decapoda Williams, Austin B.

Tardigrada Pollock, Leland W.

Cnidaria: Scyphozoa Larson, Ronald J.

Higher Fungi: Ascomycetes, Deuteromycetes, and Basidiomycetes

Cavaliere, A. R.

Copepoda: Harpacticoida Coull, Bruce C.

Sipuncula Cutler, Edward B.

Echinodermata: Holothuroidea Pawson, David L.

Copepoda: Lernaeopodidae and Sphyriidae Ro, Ju-Shey

Copepoda: Cyclopoids Parasitic on Fishes Ro, Ju-Shey

Crustacea: Branchiura Cressey, Roger F.

Protozoa: Sarcodina: Amoebae Bovee, Eugene C., and Thomas K. Sawyer

Crustacea: Cumacea Watling, Les

Arthropoda: Cirripedia Zullo, Victor A.

Cnidaria: Scleractinia Cairns, Stephen D.

Protozoa: Sarcodina: Benthic Foraminifera Todd, Ruth, and Doris Low

Turbellaria: Acoela and Nemertodermatida Bush, Louise F.

Lichens (Ascomycetes) of the Intertidal Region Taylor, Ronald M.

NOAA Tech Rep.

NMFS Circular no. NTIS no.

374 COM 73 50670

378 73 50888

384 74 50019

386 74 50014

387 7450487

389 74 51194

394 PB 257 987

397 261 839

398 268036

399 268 714

403 273062

405 274999

406 280040

409 281 969

413 222 923

419 285 538

423 296460

425 297 676

438 124 520

439 225 053

440 219 387

446 124 735

NMFS no.

Echinodermata: Echinoidea Serafy, D. Keith, and F. Julian Fell

Echinodermata: Crinoidea Messing, Charles G., and John R. Dearborn

Marine Flora and Fauna of the Eastern United States

Cephalopoda Vecchione, Michael, Clyde F. E. Roper, and Michael J. Sweeney

33

91

73

PC A03/MF A01

PB 89 189 583