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SMITHSONIAN INSTITUTION
UNITED STATES NATIONAL MUSEUM
Bulletin 76

ASTEROIDEA OF THE NORTH PACIFIC
AND ADJACENT WATERS

BY

WALTER KENRICK FISHER
Professor of Zoology and Director of the Hopkins Marine Station of
Stanford University, California

PART 3. FORCIPULATA (ConcLupeEp)

UNITED STATES
GOVERNMENT PRINTING OFFICE
WASHINGTON : 1930

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TABLE OF CONTENTS

Page

PENCE ee seme chert cee od She Sete eens Se oe ee Se eee ee es 1

PIER RGIS CAC UINION OF GHG TOUR = ooo he ee epee eee bee see ee eenne ek 2

pri teat) UV PUI eee ye ee en i a gk a ara ce ae ae 2
Key to the genera of Asteriinae, Notasteriinae, and Neomorphasterinae of the Northern

FES CSUR OED atee ee eh i ee ah teeta aie marae ain ae eee toate eee 3

STELIRUIS Cl OER ft cI oa oie ei ie eee aan eRe REN aeee 5

RTGS IRCNDARRR Sek ee aces or ee em ee Ok an ome eee eo 23

STI DPR A Rat SICA eae a a a eee a 24

FON ESRD Ra CET el ee a ee Pa te eee ieee 59

PIU OOTIUIN AAV CROMLOT NOR sn note ew is ee ae Ne a en eet 135

CRONIN ICOPURMEER RGN OES cae ern e cee: ie! ape, cern eet ee eR Se eet os 138

MREIAUDN | DOORMAN aaa ob Se ee a a he 139

RTQUIOS CIR DULOLET ROY St eh oe eS eo ee Co eee 156

RAGTIME SR MLIIBEOE WAR ee nr eres eo eon aba rine i norman seis eS 159

RESTA LI SE RCN a te Pe A ee ae Se ee a ieee eee 162

Check list of Asteroidea of north Pacific and adjacent waters______-__.__-.._--____---__ 187
List of species, subspecies, and varieties of Asteriidae in Verrill’s Shallow-water Starfishes,

1914, with their equivalent in the present monograph---....-...--------------------- 203
Summary of the Asteriinae, Notasteriinae, and Neomorphasterinae of the Northern

PORTS NISY CS ats Se a a ee am eR ee ee ine a ee ree as 204
List of species and varietal names applied to Asteriinae, Notasteriinae, and Neomorphas-

PeONaS Of Lhe Norman Pemispheres 2. 22282 Ses tk oe Se Ree 212
Summary of the genera and species of Asteriinae and Notasteriinae of the Southern Hemi-

EN ea ee aera, eg ee ee TG EB ee ee gm 216
Key to the genera of Asteriinae of the Southern Hemisphere--_-__...._.----------------- 217
List of the species names applied to Asteriinae and Notasteriinae of the Southern Hemi-

NS I ae Te a ch cae i pt a at pen oe a a ec tg 244
RSI DRA MEIN ND pe ee rae hers an pee oe te mr ie re Se oe eee 247
PEE UITENGIOTE UN NUOE tenn foe ee Pe eh te a ee en Ieee nee i eee eee 256
Ae SO he Sle as Bs cel gs Oe em a i dg wi we Recent alin 349

2 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

of Bell’s Calvasterias antipodum and Sladen’s Calvasterias stolidota and has supplied
information concerning essential anatomical details, omitted by the describers.

From acknowledgments to many colleagues who have directly or indirectly
aided my work it is to be hoped I have made no serious omissions in the introductory
paragraphs to parts 1 and 2. It is a truism not too often repeated that no serious
piece of scientific work can be completed without the cooperation of others.

In bringing this monograph to a close it is perhaps permitted one to indulge in
the pleasant retrospect of personal contacts and to record the hearty cooperation of
the staff of the United States National Museum which has been more formally
alluded to in the foregoing parts. The late Dr. Richard Rathbun, then assistant
secretary of the Smithsonian Institution, always evinced a lively interest in the
progress of this report—an interest which has happily been continued by his suc-
cessor, Dr. A. Wetmore. Nor should I forget, in matters taxonomic, the wise deci-
sions of such experienced experts as Dr. Theodore Gill and Dr. Leonhard Stejneger;
nor in matters of book making, the seasoned counsel of the editor, Dr. Marcus
Benjamin, cheerfully tendered for over a score of years.

The retrospect, coming nearer home, inevitably focuses on the late Dr. Charles
Henry Gilbert, long professor of Zoology at Stanford University. A scientist of
unusual talents and a controlled and logical mind, he followed to the letter the
Bairdian precept that what is worth doing is worth doing well. To him as professor,
colleague, and friend, I owe much, as well as to our mutual professor, colleague, and
friend, the venerable Dr. David Starr Jordan. But for their confidence the present
monograph—and others—would have been the pleasant task of another.

SYSTEMATIC DISCUSSION OF THE FAUNA
Order FORCIPULATA Perrier '

Suborder ASTERIADINA Fisher’
Family ASTERIIDAE Gray?

Subfamily ASTERIINAE Verrill (emended)

Asteriinae Verriiu, Shallow-water Starfishes, 1914, p. 42.—Fisuer, Ann. and Mag. Nat.
Hist., ser. 9, vol. 12, 1923, p. 250; Bull. 76, pt. 2, 1928, p. 57.

Diagnosis.—Asteriidae with abactinal spines short, slender to stout, conical,
tubercular-subglobose, variously granuliform, sharp to capitate, single or in groups
(but not as a rule prominent, styliform, or acicular, and more or less spaced and iso-
lated); abactinal plates in more or less definite longiseries or irregularly reticulate,
sometimes abortive; actinal area sometimes broad, with upwards of five longiseries
of plates, sometimes without any actinal plates; genital apertures dorsal, lateral, or
ventral; adambuleral spines with or without clusters of pedicellariae.

' Key to the orders of Asteroidea, see pt. 1, p. 16.

? Key to the suborders and families of Forcipulata, see pt. 1, p. 3.
+ Key to the subfamiles of Asteriidae, see pt. 2, p. 56.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 3

KEY TO THE GENERA OF ASTERINAE, NOTASTERIINAE, AND NEOMORPHASTERINAE OF THE NORTHERN HEMISPHERE

a'. Adambulacral spines provided with clusters of straight, or of straight and crossed pedicellariae,
or exceptionally with only single pedicellariae.*
b'. Adambulacrals diplacanthid, or mixed diplacanthid and triplacanthid, or mixed diplacanthid
and monacanthid.
c'. Dorsolateral plates not arranged in fairly regular longiseries; abactinal skeleton an irregular
net, with or without the carinals in an evident longiseries.

d', Gonads opening dorsally; species never carry eggs and young (so far as known).

e'. A single series of actinal plates either spineless and more or less superficially invisible,
or with a small single spine; a single series of actinal papular areas alternating
with the actinal plates; furrow spine of alternate adambulacral plates more or
less advanced into furrow; inferomarginal plates strictly actinal in position, the
superomarginals defining ambitus; cross pedicellariae not unusually large. Type,
PAS IONE ANNO GUN © Siw ie. a be as oh eS Asterias Linnaeus.

e. Actinal plates in three to five prominent series (each plate bearing one or two stout
spines), separated by series of papular areas; furrow spine of alternate adam-
bulacral plates often somewhat advanced into furrow; inferomarginals usually
not actinal but lateral in position; crossed pedicellariae not unusually large.
Tene; Ac iroschelse titimpsoms sono. ate ess 2 ons Seneee ace Evasterias Verrill.

é. No actinal plates; with unusually large, slender-jawed, crossed pedicellariae; abactinal
skeleton weak; marginal plates strong with one prominent superomarginal and one
still longer inferomarginal spine; adambulacral plates mostly diplacanthid, with
sometimes numerous straight pedicellariae along furrow margin; some of these
occasionally attached to base of furrow spines but not usually. Type, Astera-
canthion linckii Miiller and Troschel-_-_--.------------------ Urasterias* Verrill.

@. Gonads opening ventrally; many species known to carry eggs and young; abactinal
skeleton more or less open, the dorsolateral plates usually irregular, but spines
sometimes in poorly defined longiseries; spines usually small, normally with a
small or fairly thick collar of pedicellariae; crossed pedicellariae without enlarged
teeth on the moderately broad terminal lip; one, or in large species, two series of
actinal plates; adambulacrals diplacanthid or mixed diplacanthid and monacanthid;
adambulacral pedicellariae normally in clusters on the spines, but occurring singly
and even very sparingly in individuals which have few pedicellariae elsewhere on
body. Type, Asteracanthion miilleri Sars. ---~-...------------ Leptasterias Verrill.

c@. The dorsolateral plates have the appearance of being arranged in regular or subregular
longiseries; in either case the carinals and marginals are always prominent and regular,
and the dorsolateral small papular areas form longiseries.

d', Ambulacral pores not unusually large; gonads opening ventrally; adambulacrals dipla-
canthid or mixed diplacanthid and monacanthid. (Forms of L. camischatica, L.
aequalis, and L. hexactis vancouveri.) .........---.-------------- Leptasterias, part.

d@. Ambulacral pores unusually large (quadriserial); abactinal and marginal plates broad,
lobed, closely imbricated, with very small papular areas; dorsolateral plates in
three series proximally; plates closely covered with clusters of minute spinelets; two
series of actinals proximally; adambulacrals mostly diplacanthid; gonads unknown.
Type six-rayed. Type, Leptasterias macropora Verrill_.....---- Stenasterias Verrill.

b&. Adambulacral plates monacanthid; with actinal plates in one inconspicuous series or lacking
and gonads opening ventrally (paedophoric); size small; pedicellariae on adambulacral
spines rather few and inconspicuous. (Formae of several species.) ~~ Leplasterias, part.

* Genera not known from north Pacific marked with asterisk. For a key to genera of Southern Hemisphere see p. 217.

4 The clusters of pedicellariae (both crossed and straight, or either in predominance, or either alone) are a characteristic feature
of Asterias, ss., Evasterias, Leptasterias, and Stenasterias, Certain individuals of Leptasterias (as in L. littoralis) which vary (n the
direction of few pedicellariae, may almost or quite lack pedicellariae on the adambulacral spines (or in dried specimens lose the few
they may have had). The character is fundamental and the few exceptions are individual or varietal rather than specific. L.
groenlandica of the arctic Atlantic region usually lacks these pediceliariae; Bering Sea specimens have abundant sdambulscral
spine pedicellariae.

4 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

a?. Adambulacral spines devoid of attached pedicellariae, although large and small straight pedi-
cellariae may occur on the surface of the plates or on fleshy peduncles attached to plate
near base of spines; clusters of the latter in dried specimens oceasionally appear to spring
from the spine sheath but in reality do not.

b!, Abactinal skeleton an irregular net with meshes of various sizes, the plates being sometimes
closely but irregularly imbricated by their lobes; abactinal plates not in rather obvious
longitudinal series (but occasionally in more or less evident transverse series); carinal
series usually but not always distinguishable and frequently very irregular.

cl. Adambulacral plates diplacanthid, at least at base of ray; rarely triplacanthid and tetra-
canthid. :

d'. Crossed pedicellariae unusually large, the jaws narrow and rather definitely hooked
terminally; dorsolateral skeleton a very delicate and irregular meshwork which may
degenerate into more or less disconnected plates; marginal plates conspicuously
larger than dorsolateral; carinal and dorsolateral spines acicular, isolated; actinals
absent.

e!, Crossed pedicellariae very large, and of unusual form, with slender, serrate jaws ter-
minating in an unexpanded unguiculate tip; marginal spines prominent, acicular,
one to a plate, the inferomarginals with a very large cluster or cushion and the
superomarginals with a wreath of crossed pedicellariae; very numerous, large,
compressed-ovoid straight pedicellariae; gonads opening just above superomarginal
plates in interbrachial angle. Type, Asteracanthion linckii Miller and Troschel.

Urasterias* Verrill.

é*. Crossed pedicellariae smaller with relatively stouter jaws, scattered thickly over the
abactinal and lateral surfaces, but not in wreaths or clusters on the abactinal or
marginal spines; superomarginals monacanthid; inferomarginals diplacanthid, or
mixed monaecanthid and diplacanthid distally; gonads opening just above supero-
marginal plates a short distance from base of ray. Type, Asterias panopla Stux-

oe ls Sages <P r ee oo Se ee ee eee Icasterias* Fisher.

@. Crossed pedicellariae not unusually large, but small, with blunt, terminally spatulate,
denticulate jaws—the conventional type.

e!. No actinal plates; dorsolateral plates numerous, irregular, with small spinelets and
showing sometimes a tendency to form transverse series (usually, however, not
well marked); inferomarginal spines two to four in a transverse series, the con-
secutive combs closely placed. Inferomarginal spines bearing crossed pedicella-
riae; abactinal spinelets surrounded by a circlet of crossed pedicellariae.

J’. Papulae compound, each subdivided terminally into upward of eight small papillae;
inferomarginals diplacanthid, superomarginals monacanthid; adambulacrals
diplacanthid; not fissiparous; gonads opening just above superomarginals
abactinally in interbrachial angle; large unguiculate straight pedicellariae.
Type, Aphanasterias pycnopodia Fisher._______________ Aphanasterias Fisher.

f*. Papulae simple, undivided; inferomarginals with three or four spinelets in a trans-
verse comb; superomarginals with several spinelets in a comb; adambulacrals
triplacanthid, at least in part; strongly fissiparous, rays upward of eight;
gonads opening between supero and infero marginal plates, low on side of ray,
in the axillary channel; large bivalved dentate straight pedicellariae. Type,
Asteracanthion albulus Stimpson_._..-_.-._____-_____ Stephanasterias Verrill.

J*. Papulae simple, rays five, not fissiparous; gonads opening just above superomargi-
nals at a distance from interbrachial angle; adambulacrals diplacanthid; infero-
marginals and superomarginals with transverse comb of three or four spines

each; straight pedicellariae small, compressed, lanceolate.

: 7 Type, Asterias
japonica Bell

a hacia pe ce ne eee ee ee Aphelasterias Fisher.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 5

é@. Actinal plates in two or three series (or equivalent). Resembling a Henricia san-
guinolentay plates small with numerous small granuliform, close-set spinelets;
abactinal skeleton close-knit with small papular areas, rather irregularly dis-
tributed dorsolaterally but in definite marginal and actinal longiseries; carinal
plates regular; superomarginals slightly broader than inferomarginals; crossed
pedicellariae surrounding papular areas, not in circlets around spinelets; adam-
bulacral plates with a transverse series of three or four spines proximally, usually
three distally. Type, Asterias rosea Miller_.......-...---- Stichastrella* Verrill.

¢. Adambulacral plates monacanthid.

d', Actinal (ventrolateral) area broad with two to four longiseries of conspicuous spiny
actinal plates; actinostome very sunken; adoral carina narrow and long; gonads
opening dorsally; carinal plates evident; dorsolateral skeleton robust, irregular
with stout spines; in addition to large unguiculate straight pedicellariae, a peculiar
small straight variety having on each jaw a short, sharp, and a long, truncate,
laminate prong; among the adambulacral spines are clusters of straight pedicella-
riae on long peduncles arising from the plates. Type, Asterias ochracea Brandt----

Pisaster Miller and Troschel.

@. No actinal plates, the diplacanthid inferomarginals adjoining the adambulacrals. Re-
sembling Tarsaster; dorsolateral plates numerous, three or four lobed, forming a
close reticulum on either side of the definite carinal series; spines short, cylindrical,
blunt, mostly one on abactinal plates and two on inferomarginals; crossed pedicella-
riae scattered, not in circlets around spines; papular areas small in about three
dorsolateral and one intermarginal series on each side; no actinal papulae; adoral
carina composed of five pairs of contiguous adambulacrals; actinostome small,
sunken; a few fairly large spatulate denticulate, straight pedicellariae; gonads
opening dorsally. Type, Hydrasterias verrilli Fisher ---~-.----- Tarsastrocles Fisher.

b?. Abactinal plates, that is, the carinals and dorsolaterals, close-set, in very regular longiseries.
Abactinal spinelets few, stout, granuliform.

c'. Inferomarginal plates with an oblique series of three or four very flat, bladelike spines
which form a prominent fringe along the edge of ray; above inferomarginals are five
very regular longiseries of plates, the superomarginals broadest, strongly imbricated
in series; whole abactinal surface, including superomarginals, has a granular appear-
ance; five or six very short spinelets per plate; actina] plates small, one corresponding
to each inferomarginal; adambulacrals diplacanthid; tube-feet quadriserial at base
of ray. Type, Gastraster margaritaceus Perrier---------..------- Gastraster* Perrier.

c. Abactinals and marginals sparsely granulated, subhexagonal, imbricated in seven very
regular longiseries; actina' plates with short spinelets, a second series present at base
of ray only; primary apical plates enlarged, occupying all of the slightly sunken cen-
tral portion of disk; adambulacrals diplancathid; resembling abactinally a short-
rayed Zoroaster. Type, Neomorphaster eustichus Sladen-- ---- Neomorphaster* Sladen.

Genus ASTERIAS Linnaeus, emended

Asterias Linnagvus, Syst. Nat., ed. 10, 1758, p. 661; ed. 12, 1766, p. 1098. Type, A. rubens
Lrynaevus.—Gray, Ann. Mag. Nat. Hist., 1840, p. 178; Synopsis, 1866, p. 1—Prrrrer,
Rév. Stell., 1875, p. 38—Bett, Proc. Zool. Soc. London, 1881, p. 492.—StapeEn,
Challenger Asteroidea, 1889, p. 560.—Perrier, Exp. sci. Travailleur et Talisman,
1894, p. 108.—Fisner, Ann. Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 598.

Stellonia Narvo (part), Oken’s Isis, 1834, p. 716. Type, A. rubens, first species, designated,
Fisuer, 1913.—AGassiz, Mém. soc. sci. nat. Neuchatel, 1835, p. 191.

Uraster AGassiz (part), Mém. soc. sci. nat. Neuchatel, 1835, p. 191 (substitute name for
Stellonia).—Fornes, Mem. Wern. Soc., 1839, vol. 8, p. 114.

Asteracanthion Mttter and Troscuet (part), Monatsber. Akad. Wiss. Berlin, 1840, p. 102;
Archiv f. Naturgesch., 6 Jahrg., vol. 1, 1840, p. 320. Type, Asterias rubens Linnaeus.

Asteracanthium Branpt, Middendorff’s Reise in den Aussersten Norden und Osten Sibiriens,
vol. 3, pt. 1, 1851, p.. 28. Emended spelling for Asteracanthion.

63160—30——2

6 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Allasterias Verritu, Amer. Journ. Sci., vol. 29, 1909, p. 65. Type, A. rathbuni Verrill;
Shallow-water Starfishes, 1914, pp. 53, 188. .

Parasterias Verritt, Shallow-water Starfishes, 1914, pp. 53, 187. Type, P. albertensis
Verrill.

Diagnosis.—Asteriinae having the adambulacral spines provided with clusters
of pedicellariae, gonads opening dorsally, superomarginals defining ambitus, infero-
marginals strictly actinal in position, and a single series of actinal plates, either
spineless and more or less superficially invisible, or with a single spine. Abactinal
skeleton an irregular net with meshes of various sizes, the plates being sometimes
closely but irregularly imbricated by their lobes; carinal series usually distinguishable,
often irregular. Adambulacrals diplacanthid or mixed diplacanthid and triplacan-
thid, or mixed diplacanthid and monacanthid; furrow spine of alternate plates more
or less advanced into furrow; gonads large, eggs small; development indirect, through
a free pelagic larva.

Asterias,® as here restricted, includes besides the well known type, A. rubens,
A. vulgaris Verrill and A. forbesi (Desor) of the Atlantic coast of North America;
A. amurensis of the North Pacific and western Arctic; A. amurensis rollestoni (Bell)
and A. versicolor Sladen of Japanese waters.

The genus probably does not occur in the southern hemisphere, being there
replaced by Diplasterias (Podasterias), some species of which closely resemble true
Asterias.

Both Allasterias Verrill and Parasterias Verrill are straight synonyms of Asterias,

sensu stricto.
ASTERIAS AMURENSIS Liitken

Plates 1-5; Plate 6, Figures 2-8, 10, 11; Plate 7

Asterias peclinata Branpt(?), not Linnaeus, Prodromus, 1835, p. 270. Kamchatka.

Asterias amurensis LUrKEN, Videnskab. Meddel. fra d. naturhist. Foren. Kjobenhavn, 1871,
p. 296.—Fisuer, Ann. Mag. Nat. Hist., ser. 9. vol. 12, p. 598.

Asterias rubens Murvocn, Rep. Internat. Polar Exped., Point Barrow, 1885, p. 159.

Allasterias rathbuni Verriti, Amer. Journ. Sci., vol. 28, 1909, p. 65, fig. 5; Shallow-water
Starfishes, 1914, p. 189 (‘‘Maloska””=Unalaska).

Allasterias rathbuni var. nortonensis VERRILL, Amer. Jour. Sci., vol. 28, 1909, p. 66, fig. 7
(Norton Sound).

Allasterias rathbuni var. anomala VeRRILL, Amer. Jour. Sci., vol. 28, 1909, p. 66, fig. 5 (St.
Michael, Norton Sound).

Allasterias rathbuni nortonensis VerriLu, Shallow-water Starfishes, 1914, p. 191, pl. 78,
fig. 2, text figs. 8, 9.

Allasterias anomala VerriLu, Shallow-water Starfishes, 1914, p. 193, pl. 59, fig. 2; pl. 60,
fig. 2; pl. 69, fig. 5; pl. 77, fig. 3; pl. 78, figs. 3, 4.

Parasterias albertensis VERRILL, Shallow-water Starfishes, 1914, p. 187, pl. 57, figs. 1, 2; pl. 70,
fig. 6 (Albert Bay, British Columbia).

Asterias anomala A. H. Ciark, Rep. Canadian Arctic Exped., 1913-1918, vol. 8, 1920, p. 8c.

Asterias nortonensis A. H. Cuarx, Rep. Canadian Arctic Exped., 1920, p. 8c.

The following is a translation of Doctor Liitken’s original description: 7

* For a full summary of species see page 205.
7 Made by Dr. Adam Béving, of the U.S. National Museum, to whom I am greatly indebted.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 7

Asterias amurensis

From the Okhotsk Sea and the coast of Amurland we have a quantity of large specimens of
this starfish. It is very different from the one described by Brandt under the name of Asteracanthion
ochotensis; this I have been able to determine through the kindness of the St. Petersburg Museum.
The specimens from the Amur were collected by Captain Andrea.

All the available specimens (including the one in alcohol) have very broad and flat arms the
dorsal and ventral side of which are separated by a rather sharp border. The madreporic plate is
large and easily seen, and is located nearer the center than the angles between the arms. The dorsal
side is rather densely beset with very short spines the position of which is without regular arrange-
ment. They are placed singly, or two or three, sometimes four, together; along the mid line of the
arms they come together in a wavy line that stands out clearly between the rest of the spinules,
which are scattered and weak. The spinular covering is denser on the disk than on the arms. The
individual spinules are conical or cylindro-conical, covered with asperities, and terminating in one,
two, or three points. Between the reticulations of the skeleton there are numerous papulae; over
the whole dorsal skin there are also a great many irregularly placed 2-bladed pedicellariae, both on
the trabeculae of the skeleton and in the interstices; on the spines themselves I have never observed
them. Along the lateral edges of the arms the spines are densely placed in a multiple series, and
they also have a peculiar flat excavated and apically blunted form. After this dense series of mar-
ginal spines there follow on the ventral side first a zone which is nearly without spines, then a double
(in large specimens triple or quadruple) series of similar but somewhat longer spines; then again a
bare zone; and finally, near the tube feet, a dense double series (in large specimens it is apparently
multiple) of slender, nearly cylindrical, somewhat flattened and excavated ambulacral spines. The
2-bladed pedicellariae which are placed between these three ventral spinous zones are noticeably
larger than those of the dorsal side; but besides these a third form is present that is much smaller
than those of the dorsal side, occurring in great quantities on the ambulacral spines, the marginal
spines, and on the spinous zone between; they are especially present on the more or less excavated
part of those spines which is turned away from the ambulacral groove; at the base of the ambulacral
papillae there are furthermore to be observed quite a few large pedicellariae. The largest specimen
before me has R=120 mm., r=35 mm.; the width of the arms at the base is 38 mm.

The following is Doctor Liikten’s diagnosis:

Diagnosis.—Statura ampla. Branchia quinque lata, depressa, ancipita, acie acuta superficiem
dorsalem a ventrali separante. Spinae dorsales disci densius, brachiorum sparsius despositae,
singulae vel 2-4 juxtapositae, in media brachiorum parte saepe lineam continuam formantes, breves,
conicae vel cylindroconicae, asperae, apice bifidae vel trifidae. Corpus madreporiforme spectabile,
centro propior. Areae poriferae multiporae inter trabeculas sceleti calcarei. Pedicellariae minutae
rectae compressae bidentes numerosae supra totam superficiem dorsalem sparsae, in spinis dorsalibus
nullae. Ad margines laterales brachiorum spinae breves, planae, excavatae, obtusae, pluriseriatae,
dense collocatae. In pagina ventrali zonae longitudinales binae nudae, spinis fere destitutae, cum
serie spinarum similium sed paulo majorum duplici (in parte intima brachiorum saepe triplici vel
quadruplici) alterant. Spinae ambulacrales graciles, cylindricae fere, mediocriter compressae,
canaliculatae seriem duplicem formant vel multiplicem simulant. Pedicellariae dorsalibus con-
formes sed duplo majores zonas illas nudas maximo numero ornant, ad basin spinarum ambulac-
ralium quoque crebre conspiciuntur; minutissimae, confertae, cruciformes latus cavam praecipue,
pedibus averssum, spinarum ventralium, ambulacralium et marginalium frequentant.

Through the kindness of Dr. Th. Mortensen I have received a cotype, one of the
original lot from the Amur country, and have made direct comparisons with numerous
examples from Kamchatka and Alaska, as well as with specimens of Asterias amurensis
rollestoni (Bell) from several localities in Japan.

The detailed description is based upon specimens from station 3242, 11 fathoms,
Bristol Bay, Bering Sea. The cotype, figures of which are given, is closely similar to
these, with the exception of having slightly smaller crossed pedicellariae.

oOo

BULLETIN 76, UNITED STATES NATIONAL MUSEUM

It is obvious that Asterias rubens, Asterias vulgaris, and Asterias amurensis are
very closely related and it is profitless to discuss whether they are distinct species or
subspecies. There is no evidence of geographic intergradation between rubens and
amurensis.

Diagnosis.—Differing from A. rubens Linnaeus in having, when adult, conspic-
uously broader rays, more numerous, usually shorter abactinal spinelets; 5 to 10
channeled or scoop-shaped superomarginal spines instead of two to five more or less
terete, tapered, cylindrical, or clavate ones; in having a more nearly plane actinal
surface (sharply defined by the rather thin edge of the ray, bearing a cheveux-de-
frise of superomarginal spines), and by having a broader, flatter actinal interradial
region; intermarginal spines frequently present. Rays constantly five; abactinal
surface low-arched, resistant or flaccid; actinal surface nearly plane, with an unusually
broad actinal area on proximal third of ray; rays broad at base tapering rapidly to a
blunt point; interbrachia angular; adambulacral spines proximally usually three and
two alternating; inferomarginal spines usually two. Large specimen, R 195 mm., r
53 mm., R=3.68 r; breadth of ray at base 65 to 67 mm., or one-third of r.

Description.—This species is very variable. While large specimens all have the
same general appearance and are easily recognized, a detailed analysis of the struc-
ture will reveal considerable diversity in the size of the skeletal parts, especially of
the dorsum, and in the number, robustness, and form of the spines. The breadth of
the ray increases with age in such a way as to alter very materially the appearance
of the animals. Quite immature specimens are therefore unlike the adults in general
appearance. The following description is based upon a large specimen from station
3242, 11 fathoms, Bristol Bay. (PI. 2, fig. 3; pl. 3, fig. 5.) Some of the variations
exhibited by adults from southern Bering Sea are also noted. Under variations are
listed the principal varieties which are present in a large series of specimens examined.

The Bristol Bay example measures: R 195 mm., r 53 mm., breadth of the ray
at base, 65 to 67 mm., or about one-third R. The actinal surface is low arched,
rather stout and resistant, and is beset with numerous uniform, small but robust
spinelets 1 to 1.50 mm. long, outlining a very irregular and quite open mesh, con-
taining many papulae, some of which are retracted, others inflated. The skin is
pulpy and minutely wrinkled. A carinal series is scarcely discernible, though it is
well marked in other specimens from the same haul by reason of the spines being
much more closely placed. Laterally the spinelets tend to form longitudinal irregu-
lar series. The spinelets are about 0.75 mm. thick, often clavate, irregularly truncate
with a groove or a shallow pit at the tip. The tip may be compressed and incipiently
bifid. The groove may be simply a cleft in the side of the little pit or it may
extend down the side of the spine which becomes a thick-walled half-tube. The
distal spinelets are mostly without pits or channels. There is a good deal of varia-
tion, depending upon age and locality in the forms of the abactinal spinelets. They
tend to become stouter and more gouge-shaped with age. In many small and medium-
sized examples the spinelets are slender clavate or robust clavate, blunt or truncate,
without any groove.

; The superomarginal plates define the ambitus and their spines, which are about
twice as long and two or three times heavier than the dorsolaterals, form a dense
cheveux-de-frise on the very angular margin of the ray. There are six to eight divar-

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——-FISHER 9

icate, spatulate, gouge-shaped, round-tipped, spines to each plate of the proximal
half of the ray [fewer distally and proximally in small specimens] and the clusters are
close together. The channel is usually, but not invariably, on the upper or abactinal
sides of the spine, and in small specimens the spines may not be channeled at all
but simply clavate, and of varying degree of robustness.

Nearly midway between the ambitus and furrow margin is a double row of gouge-
shaped spines separated from the superomarginals, by a conspicuously broad inter-
marginal channel and from the adambulacrals by an actinal channel, both densely
covered with papulae. While in this specimen there are only occasionally three
inferomarginal spines, some examples, as, station 3243, have fairly regularly three
on each plate of the proximal half of the ray, in addition to one or two short
series of intermarginal spines (springing from intermarginal ossicles). The large
specimen from station 3242 has no intermarginal spines but another from the same
haul, with R 156 mm., has a series extending nearly half the length of ray, and
spaced from these, toward the margin, a second row not so long. There are thus on
the proximal third of ray four parallel series of spines, of which two are inferomarginal.
The inferomarginal spines are untapered, or slightly broader at the tip than midway,
or the reverse, and the depth of the channel, which is on the admarginal side, varies
greatly in different examples, being absent in young. In the largest specimen the
spines are 3 to 4 mm. long by 1 to 1.25 mm. broad. The first spine corresponds to
the eleventh adambulacral plate. The marginal and abactinal spines are distally
rough and minutely thorny.

The actinal channel is well marked but not so broad as the intermarginal.
There are no actinal spines, but a single series of small actinal plates, largely obscured
by the inferomarginals, is present.

The adambulacral spines are longer than the inferomarginal (about 5 mm.) and
are so placed as to form three longitudinal rows along the margin of furrow. The
spines of the innermost row are only on alternate plates, are set rather well into the
furrow, on the furrow face of the plate, as in Asterias rubens, and are compressed,
tapered, and bluntly pointed. This is followed by two slightly longer, often tapered,
blunt spines (or by only one on the outer part of the ray) having a variably developed
channel or groove down the outer side. The plates alternating with these have,
proximally, two grooved subambulacral spines; distally only one. There are thus,
proximally, three and two adambulacral spines alternating and distally two and one.
Another large specimen from this station, already referred to, has two spines on both
sorts of plates proximally and distally two and one, alternating. The grooves on the
subambulacral spines becomes less and less well marked in progressively smaller
specimens. It is normlly absent in small specimens even though well marked in
large examples from the same locality. |The position of the furrow spine of alter-
nate plates, near the morphological upper end of the adambulacral plate, led Pro-
fessor Verrill to found the genus Allasterias. But the alternate furrow spines of
Asterias rubens, the type of Asterias, have the same position.]

The mouth plates are sunken and bent down toward the actinostome, as is
characteristic for the genus, but the oral angle although composed of upward of six
pairs of contiguous adoral adambulacral plates, is not a thin carina as in Orthasterias
and Pisaster, but broadens rapidly from the apex and is stout, and crowded with

10 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

prominent spines. The mouth plates are small (but relatively larger in medium-
sized and young examples) and the strongly arched sutural surface is turned toward
the actinostome so that the chord or long axis of the surface is not horizontal but
more nearly vertical, varying to about 45° in younger specimens. There are two
spines on each plate—a slightly curved tapering blunt one at the inner end, directed
over actinostome, and a straight tapering longer one, about the middle, and bent
away from actinostome. ‘The first adambulacral plate is conspicuously larger than
the second, and the second than the third. In the largest specimen the first plate is
actinally about three-fourths as long as the corresponding sutural dimension of the
mouth plates; the second is three-fourths the first; the third, three-fourths the second,
and the fourth, three-fourths the third; the fifth and succeeding plates about the same
as the fourth. Viewed from the furrow face these plates resemble the stones of an
arch, as the series is bent toward the actinostome at the inner ends. In small speci-
mens this arching becomes Jess and less evident with decrease in size, and fewer
plates are joined behind the mouth plates. The first and second adambulacrals,
and sometimes the third also, carry but one spine. The first furrow spine may occur
on the third plate or not until the seventh or eighth.

Abundant abactinal, intermarginal, and actinal papulae, the number increasing
greatly with age.

Pedicellariae very abundant. Small ovate to lanceolate straight pedicellariae,
much smaller than the spinelets, are scattered over the abactinal surface (largest in
interradial depressions), on the superomarginal, intermarginal, and inferomarginal
plates and the bases of the spines, and on the adambulacral and mouth plates and
spines. There is a cluster of about 5 to 10 near the end of the furrow spines, and in
addition to the more numerous crossed pedicellariae, also several near the end of the
other adambulacral spines. Straight pedicellariae of the actinal surface are fully
twice as large as the dorsal; of the former the intermarginal are probably the largest,
though in some specimens the size increases slightly toward the furrow (but the
adambulacral spinal pedicellariae are smaller). The number of pedicellariae varies
greatly with age, and to some extent with locality.

Small crossed pedicellariae present on the abactinal and actinal surfaces, forming
usually a single circle around the dorsal spines, and are distinguishable in dried speci-
mens from the straight pedicellariae by their smaller size and blunt rounded tips.
(The straight pedicellariae are found on the plates and among the papulae, and some
of them are quite as small as the crossed.) The crossed pedicellariae, also, form
clusters on the upper and outer sides of all the marginal and subambulacral spines,
but not on the alternate furrow spines which carry only straight pedicellariae.

The pedicellariae are treated in some detail under Variations, where comparative
measurements are given. The measurements of pedicellariae of a giant specimen
would be somewhat misleading. See also, figures.

Madreporie body large, circular, nearly flat to decidedly convex, situated one-
half r from center. It is 7 mm. in diameter and the fine striae radiate from the center.

Variations —The breadth of the ray and width of intermarginal and actinal
channels increases with age, as do, in general, the number of spines, pedicellariae,
and papulae, the rigidity of the skeleton, and thickness of the spines. These changes

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER ll

operating together, or only in part as sometimes occurs, effect considerable differences
in general appearance.

But in medium-sized and large examples that may reasonably be regarded as
mature there are rather striking variations. The abactinal spines differ somewhat with
practically every locality, and even at the same station exhibit minor differences not
connected with age. They vary from fairly slender, slightly tapering, to stout, cylin-
drical, or clavate. The end may be merely blunt; or bifid or trifid; or swollen; or
compressed a little and therefore sometimes spatulate, usually with a faint to decided
groove. Such gouge-shaped spines predominate on the proximal part of the ray of
some examples which have the spines of the distal half more generalized. Gouge-
shaped spines are rare on immature examples, even though the adult has them well
developed.

The stoutness of the abactinal ossicles and the relative size of the meshes of the
skeleton are variable. The skeleton is sometimes quite flaccid, but this may be due
partly to maceration in weak alcohol. In examples from shallow water both stout
(station 3231) and relatively weak (station 3298) plates are found. The character
of the bottom, and available food may affect this condition, but nothing is known
concerning the cause of the difference. Usually the skeletons composed of weak
plates carry the small terete, unmodified spines.

The marginal spines of shallow-water specimens seem to be at least slightly gouge-
shape or grooved down the outer side in mature, large examples, but the stoutness of
the spine, its form and the size of the groove varies greatly. In extreme cases the
inferomarginal spines, which may be taken as an index, are rather slender, tapered,
and only faintly grooved (Popoff Strait), while from the same locality another large
specimen has spatulate conspicuously furrowed spines relatively two to three times
as broad. From station 3231 (see p. 19) are examples of forma anomala having still
stouter more scoop-shaped spines. There is every gradation between the extremes.
In young, the marginal spines are always slender, and much more constant in form.

Certain specimens without reference to locality have one or two rows of inter-
marginal spines on the proximal part of the ray, the number of spines varying. Some-
times they have the appearance of completely filling the intermarginal channel prox-
imally. This feature occurs in specimens from various parts of the range, for instance,
southern Bering Sea, Norton Sound, Petropavlovsk, and in forms with both slender
and very stout spines. It is present also in examples from deep water (81 fathoms,
station 3257).

While most of the inferomarginal plates carry two spines in an oblique series,
certain plates here and there have three in a triangular group. Frequently at the base
of the ray there appears to be regularly three inferomarginal spines, but the outer is
almost always an intermarginal.

The form of the subambulacral spines varies with that of the inferomarginals.
When the latter are strongly sulcate, the subambulacrals are also more or less deeply
grooved. This is well shown in examples from station 3231 (forma anomala), from
other parts of Bristol Bay, Popoff Strait, Norton Sound, St. Michaels, and Petro-
pavlovsk. The number of adambulacral spines is variable, two and three alternating,
being the maximum. Often there are two on each plate or one and two alternating,
with occasionally one and three. Even in large specimens the maximum number is

12 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

found only at the base of the ray, one and two alternating being the rule on the distal
part.

The number of abactinal crossed pedicellariae is variable, but for accurate
comparison the character is too difficult to handle precisely. The size of the crossed
pedicellariae is, however, useful. There is an increase in size from northern Japan
to Bering Strait, while in eastern Bering Sea there is a further increase with depth.
In shallow water a curious variety with heavier skeleton and spines (forma anomala)
has larger pedicellariae than the typical form. A comparison of the sizes and details
of pedicellariae is taken up in the following section, but the figures will give the best
idea of the variations.

VARIATIONS OF SPECIMENS BY LOCALITY

1. Amur (Gulf of Tartary).—This is the cotype, collected by Captain Andrea.
As I have but the one specimen, no notes can be given on the range of variation at
the type locality; but there is no reason to expect that it is any less than at Petro-
pavlovsk. Liitken, however, has described the types fairly completely, and I shall
cive a few figures of the spines and pedicellariae. R 115 mm.; rays flattened and
distorted.

The abactinal spines give the impression, at first sight, of being mostly conical
and pointed, but many of them are two or three pointed as Liitken writes. Such
spines appear to have a notch or a pit at the tip, especially on the radial region; still
others are frankly gouge-shaped at the tip. The abactinal crossed pedicellariae
are few compared to the larger (but still small) straight pedicellariae, which are
broadly lanceolate, acute, and about 0.375 mm. long. The abactinal crossed pedi-
cellariae are about 0.23 or 0.24 mm. long. (PI. 6, fig. 2.)

There are eight or nine gouge-shaped superomarginal spines, proximally, and two
rows of intermarginal spines; inferomarginal spines two, broadly grooved or gouge-
shaped; adambulacral spines usually two to a plate, the alternate plates with the
characteristic furrow spine.

The actinal straight pedicellariae are broadly lanceolate, sharp and much larger
than the dorsal (about 0.75 mm. long proximally on the intermarginal furrow).

Comparing the size of the actinal crossed pedicellariae of the cotype with com-
parable examples from Kamchatka, and of rollestoni from Japan, we have the
following:

rollestoni (R 125 mm.), Hokkaido,’ 0.17 to 0.28 mm. (PI. 6, fig. 1.)

amurensis (R 115 mm.), Amur, 0.25 to 0.28 mm. (PI. 6, fig. 2a.)

amurensis (R 130 mm.), Petropavlovsk, 0.3 to 0.36 mm. (Pl. 6, figs. 3, 3a.)

The actinal crossed pedicellariae of A. amurensis rollestoni are therefore about
three-fifths the length of those of A. amurensis amurensis from Kamchatka. The
cotype (Amur) is intermediate, but rather nearer to the northern specimens.

2. Petropavlovsk, shore (Pl. 1, fig. 3, pl. 3, figs. 2, 3).—There is a good series
ranging from R 12 mm. to R 130 mm. (the last with r 35 mm., R=3.7 r.). The

* Specimens from Mororan on the south coast. Most of the pedicellariae are small, 0.18 to 0.21 mm., but rarely larger ones
occur which measure 0.26 mm. These are very much in the minority, however. In Kamchatkan and Bering Sea examples occa-
sional pedicellariae are found which are considerably larger than the average. Specimens of rollestoni from farther south on the east
coast of Japan have actinal pedicellatiae as large as 0.27 mm. and assmal!as0.18mm. In typical rollestoni the abactinal spines are
fewer, stouter, and more widely spaced.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 13

abactinal surface of large specimens is rather uniform in appearance. The dorsal
spinelets are relatively small, a trifle shorter and slightly less numerous than in the
cotype. They are rather more stubby and subclavate, but are, however, scoop-
shaped in many cases or incipiently cleft at the tip; epidermis thick; straight
pedicellariae rather few. There are six or seven superomarginal spines to a plate,
usually one short intermarginal series, regularly two inferomarginal spines, and proxi-
mally two and three adambulacral spines, then two and two, and distally two and one.
The ventral spines have the usual armature of pedicellariae.

The superomarginal and inferomarginal spines are robust and scoop-shaped,
varying to a slenderer form, with or without a groove along the outer side. The
marginal spines vary in stature from that characteristic of Verrill’s ‘‘nortonensis”’
(yet Norton Sound examples may have thick spines) to the broadly spatulate scoop-
shape form of forma anomala—medium slender to spatulate.

The subambulacral spines vary in size or robustness corresponding to the devel-
opment of the marginal spines, and they usually have a well-marked groove down
the outer side. A mature example, with R 89 mm. has, however, slender subambu-
lacral and marginal spines without a groove or with only an indication near the tip.
So far as form of spine is concerned, this series includes nearly the extremes of
variation.

The pedicellariae are larger than in the cotype, yet not so large as in some exam-
ples from the eastern part of Bering Sea.

The details of the crossed pedicellariae, always somewhat variable, are pretty
close to those of the cotype’s pedicellariae. The straight pedicellariae, however,
begin to show a tendency, much better marked in eastern Bering Sea specimens, to
develop irregular spines at the tip of the jaws. (PI. 7, fig. 1.) No two pedicellariae
are exactly alike and a comparison of figures will give a better idea of the structure.
In the cotype there is only the faintest suggestion of this. (PI. 6, figs. 10, a, }, ¢.)

The larger straight pedicellariae of the actinal surface are about 0.75 mm. long,
or sometimes somewhat larger. The crossed pedicellariae of the inferomarginal
spines, in a specimen with R 130 mm., are 0.315 to 0.36 mm., while in one with R 89
mm., they measure 0.29 to 0.325 mm. The latter is a specimen with slenderer spines
than the first and the pedicellariae are a shade less robust. (Compare pl. 6, figs. 3
and 3a.) These crossed pedicellariae, as noted in the preceding section, are larger
than in the cotype.

The young, of course, have narrower rays and in very small examples (R 20
mm.) the intermarginal channel is lateral instead of ventral, while the ray is more
nearly terete. This gives the creature an entirely different aspect. There are no
actinal papulae, only one intermarginal papula to an area, and one or two abactinal
to an area. Superomarginal spines two or one; inferomarginal two; adambulacral
two, two or one, two.

3. Port Clarence, Alaska, near Bering Strait—Two specimens. One has R 105
mm., r 25 mm., R=4.2 r.; breadth of ray at base 24 to 27 mm, The other has R
82 mm. In both specimens the abactinal spinelets are small as in the Petropavloysk
specimens, slightly more numerous, slightly tapered in some cases, blunt or incipiently
_ grooved in the larger, and more often bluntly pointed in the smaller, The larger has

14 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

10 or 12 superomarginal spines to the plate proximally, while the smaller has about six.
The larger has frequently three or four inferomarginal spines, and the intermarginal
space of the proximal half of the ray is filled irregularly with similar grooved spines;
the smaller has regularly two inferomarginal spines and a single regular very short
series of intermarginals. The spines are stouter in the larger specimens. Both have
the adambulacral spines 2-3 proximally, but in the larger the sequence is irregular;
a very few plates have four spines. The abactinal crossed pedicellariae are not
numerous and do not form complete circles around the spinelets. The actinal crossed
pedicellariae are very closely similar in size and details to those from Kamchatkan
specimens, as are also the straight pedicellariae.

In the larger specimen the carinal series of spines is not discernible at all; in the
smaller it is well marked.

4, Norton Sound, Alaska (pl. 2, fig. 2; pl. 3, fig. 4)——There are five large speci-
mens, no. 7621, from which the type of Verrill’s nortonensis was selected. The largest
measures R 120 mm., r 36 mm., R=3.3 r; breadth of ray at base variable, about
equal to minor radius.

These specimens are quite comparable to those from Petropavlovsk, being
characterized by small but variable abactinal spinelets, few abactinal crossed pedicel-
lariae, but numerous small straight ones, six or seven stout, grooved, superomarginal
spines, moderately slender to distinctly robust gouge-shaped inferomarginal spines,
and slender ungrooved to stout, grooved subambulacral spines. There are usually
two and three, or two and two adambulacral spines proximally, though the regularity
is often broken.

Two large specimens of the same size differ in that one has nearly twice as many
abactinal spines as the other. The example with the more numerous spines has them
stouter, often broadened and gouge-shaped at the tip, and with fairly complete
circumspinal wreaths of pedicellariae—about as numerous as in southern Bering
Sea examples. The other one has more nearly cylindrical and bluntly pointed spine-
lets and perhaps half as many crossed pedicellariae.

In his description of Allasterias rathbuni nortonensis Verrill says that the type
and only specimen seen “‘presents several rather strongly marked characters’? but
does not enumerate them beyond saying that ‘“‘the characters that seem most impor-
tant are the smaller size and the stouter form of the major straight pedicellariae. The
dorsal and marginal spines are longer than in the typical form and the papulae are
more numerous.”

The straight pedicellariae (pl. 7, figs. 1, 1a) are not more robust than in specimens
from other localities, there being variation in this regard. In an example with R
114 mm., the actinal straight pedicellariae are 0.75 to 0.85 mm., and hence pretty
close to those of the cotype and of Kamchatkan specimens. The tips of the jaws are
more elaborately spiny than in either of the above.

The crossed pedicellariae (pl. 4, fig. 4) are closely similar to those of the cotype,
though larger. The abactinal are about 0.26 mm. long, and the actinal from 0.27 to
0.315.; hence they are nearly as in Petropavlovsk examples. The form is also closely
similar, the jaws being of the stouter type that is found in some of the heavier spined
Petropavlovsk specimens.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——-FISHER 15

The subambulacral spines are slenderer than is usual, but are grooved at least
on the distal part of the spine. (See Verrill, 1914, pl. 78, fig. 2.) Professor Verrill’s
type of nortonensis is an unusually slender-spined specimen; two of mine have some-
what stouter subambulacrals. These, as well as the inferomarginals, are stoutest in
the large specimen with the fewest abactinal spinelets, noted above.

5. Specimens from 50 fathoms or deeper.—Stations 3225, 3257, 3310, 3311, 3334,
3335 (pl. 4, figs. 4, 5), 3538, 3539, 3547.

There are, altogether, 40 specimens from depths of 50 fathoms to 93 fathoms,
taken north of Unimak and Unalaska, and from the vicinity of the Pribilof Islands.
They are therefore from southern Bering Sea.

The largest specimen (station 3335, with R 145 mm., r 36 mm., R=4 r; pl. 4,
figs. 4, 5) is fortunately from the greatest depth. It is comparable with large fully
grown examples from shallow water in the same region, and there is surprisingly
little difference, if the comparison is made with the slenderer spined specimens.
The abactinal spinelets are smaller and a little more numerous. They are about the
same as in the specimens from Petropavlovsk and northern Bering Sea. In form
they are tapering and blunt or else the tip may be compressed. The superomarginal
and inferomarginal spines are distinctly grooved, but the spines are often slightly
tapered and of medium width. Occasionally the superomarginals are slightly
spatulate. There is a single series of intermarginal spines extending about a third
the length of the ray. The subambulacral spines are rather slender, tapered slightly,
with a groove on the outer side. They are slenderer and relatively longer than in
the Petropavlovsk examples, but are not different from specimens from station 3242,
and other shallow water localities. There seems to be no constant difference between
this large fully grown example from 93 fathoms and shallow water specimens.

In medium-sized and smallish specimens from deep water (as station 3257,
81 fathoms) the abactinal spinelets are so short that they are often no longer than
the numerous pedicellariae, the abactinal surface appearing as if covered with a
close rather uniform nap. In other specimens the pattern of the abactinal skeleton
can be readily traced by reason of the stouter spinelets.

Specimens from intermediate depths show all sorts of variations, with the
exception of the very heavy spatulate ventral spines. An example from station
3281, 36 fathoms, has coarse, well-spaced abactinal spines which are notched, pitted,
and grooved as in shallow-water examples.

Specimens with the weakest, most flaccid abactinal skeleton are from station
3242, only 4.5 fathoms.

The crossed pedicellariae are surprisingly large. In a specimen from station
3225 (R about 115 mm.) the abactinal crossed pedicellarie of the abactinal surface
range from 0.27 to 0.31 mm.; of superomarginal spines, 0.27 to 0.36 mm.; of the
inferomarginal spines, 0.31 to 0.36 mm. (see pl. 6, fig. 7); of the subambulacral
spines 0.36 to 0.38 mm. The actinal straight pedicellariae range from about 0.6 to
0.9 mm., the form being shown in Plate 7, Figures 10, lla. The crossed pedicellariae
are about as large as those of the heavily spined forma anomala and are larger than
those of comparable shallow-water examples having typical spines. For instance,
the inferomarginal crossed pedicellariae of the deep-water specimens are usually
between 0.34 and 0.36 mm. (station 3225), while in a comparable example from

16 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

station 3244, Bristol Bay, 4.5 fathoms, these pedicellariae are 0.25 to 0.315 mm.;
in an example from station 3242, 11 fathoms, 0.27 to 0.315 mm.; in a specimen from
3248, 21 fathoms, a maximum of 0.315 mm.; Popoff Strait, Shumagin Islands, low
water, 0.27 to 0.315 mm. These measurements are in agreement with those of
crossed pedicellariae of comparable specimens from Petropavlovsk, Port Clarence,
and Norton Sound. The larger crossed pedicellariae are therefore a characteristic of
deep water specimens as they are of the thick-spined forma anomala.

Growth stages and young (pl. 5).—The change in form due to increase in size is
shown by the following measurements of specimens from stations 3241 and 3242.
The younger stages have both narrower rayed and broader rayed forms:

So 1 esl 2 a Algae ol 2 6 7 8 9 10 u
Station soos soe 32i) | 3241 | 3241 | 3241 3241 3241 3241 3242 | 3242 3242 3242
Re cesta cetedle o | 12 19 25 29 | 32 35 45 | 70 79 150
PAS 2 on ee ea ena ane 3 4 6 65 /} 9 7.5 12 | 20 22 45
Reef oe. Abs aan eee. Cues oe 4 4.75 4.16 4.46 3. 55 4.6 4.2 3.75 | 3.5 3. 59 3.3
Breath of ray in mm-_-_.____......--- 3.5 5 | 7.5 | 6.5 | 10.5 10 12 14 | 21 27 52

The ratio between R and the breadth of the ray at the widest part, close to the
base, varies from 3.4 : 1 in No. 1 to 2.8 :1in No. 11. No 5 is a broader rayed form,
while 4 and 6 have relatively slenderer rays. In specimens of the size of No. 8, and
larger, the proximal third of the ray increases more rapidly in width relatively than
does the distal portion, so that the difference in appearance with increasing age is
more marked than the figures seem to indicate. With the broadening of the ray at
the base the intermarginal channel makes its appearance and quickly widens, leaving
space also for an actinal channel. The presence of these regular channels is one of
the characteristic features of this species, as of rubens.

In specimens as small as Nos. 1 and 2 the inferomarginal spines are lateral, with
the superomarginal directly above them, the rays being subterete. They lack
entirely the characteristic flattened appearance of the adult. In Nos. 5 and 6 the
superomarginals are situated a little more laterally than the inferomarginals, while
in No. 7 the inferomarginals are proximally frankly actinal in position, and the ray is
acquiring the characteristic flattened appearance.

In No. 1 and other small examples the abactinal skeleton has relatively large
meshes (naturally much fewer than in large examples) with one or two papulae to a
mesh. The abactinal spinelets, 0.3 to 0.4 mm. long, are slender, slightly tapered,
not close together and are tipped by several thornlets. Each mostly 3-lobed primary
plate bears a spinelet, but the oblong or elliptical connecting plates are often spineless.
In No. 1 there are no abactinal pedicellariae. These appear in No. 2.

In Nos. 1 and 2, the marginal plates are close together, one above another on
the side of the ray, four lobed, and the lower or descending love of the superomarginal
covers the dorsal lobe of the inferomarginal as in the various genera allied to Coscin-
asterias. There are one superomarginal and two inferomarginal spinelets and one
to three forcipulate pedicellariae on them. The intermarginal papular areas are
very small with a single papula to each. No actinal papulae in No. 1; a few make

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——FISHER 17

their appearance in No. 2, and in No. 3 the series extends half the length of the ray.
In No. 3 the superomarginal spines have increased to two or three; in Nos. 5 and 6,
three is the commonest number, occasionally four, while in No. 8, four or five are
found.

No. 1 and No. 2 have proximally two, then alternately one and two, adam-
bulacral spines, and while the plates upon which the later furrow spine will stand
can be recognized, the spine itself is not yet differentiated from the others. This is
very definitely beginning in No. 3, and the spines are well differentiated in No. 5.
Adambulacral spinal pedicellariae appear first in No. 3.

The small specimens have no actinal plates. In Nos. 1 and 2 there is only
one pair of adambulacral plates in contact behind the mouth plates; in Nos. 5 to 8
there are three, and in No. 10 there are five.

In No. 8 the superomarginal, inferomarginal, and subambulacral spines show
the beginning of a definite shallow groove on the outer side, and the marginal spines
are slightly broader toward their rounded tips. Considerably larger specimens, as
for instance 9 and 10, may have narrower marginal spines; and the subambulacral
spines are less evidently grooved—in fact the furrow shows faintly at the top only.
The amount of grooving, after the animal reaches the age when it should appear, is
not dependent upon size alone, but is subject to a very wide range of variation often
in specimens from the same station, as in the case of the Petropavlovsk examples.

Anatomical notes —Skeleton: As noted under description and variations there is
considerable diversity in the stoutness of the skeleton—enough certainly to justify
the assumption of more than one species if only a few isolated extremes were avail-
able. The skeleton is especially heavy in forma anomala.

The abactinal skeleton is the open irregular mesh type of rvbens, with usually
an irregular carinal series of plates forming a wavy line along the radius. The
plates of the dorsolateral regions have generally three irregular lobes, or three sides,
while the carinals start out in small specimens fairly regularly four lobed but become
irregular with age. The 3-sided dorsolateral plates stand at the nodes of the net
while the connecting bars are composed of one or two narrowly to broadly elliptical
overlapping ossicles, some of which bear a single small spine. The primary plates
usually bear a single spine, but some of them especially in the carinal series bear
two or three. The breadth of the abactinal plates and the degree of overlapping
determines the openness of the skeleton. The number of the meshes from one side
of the ray to the other varies with age, and is difficult to count accurately—perhaps
12 to 14 is a fairly correct estimate for medium-sized specimens.

The superomarginal plates start out in young specimens in the normal 4-lobed
form for the family but the dorsal lobe is later suppressed, while the ventral becomes
less conspicuous as the plate lengthens, until a sort of lozenge form results. The
plates are strongly imbricated with the adoral end outside. The inferomarginals
are larger, more definitely four lobed, and are connected with the superomarginals
by from one to three overlapping ossicles (except distally and in young specimens).
One or even two series of these intermarginal ossicles may bear solitary spines
proximally. (Normally the plates adjacent to inferomarginals bear a short series
at base of ray.) The actinal plates are inconspicuous except near the mouth plates;
there is a single incomplete series, which appears rather late, and they are spineless.

18 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

They have each, however, a long internal process which helps to support or strengthen
the dorsal surface (Vide infra). ie

The most characteristic features of the internal skeleton are the extensive inter-
brachial septum; the series of accessory septa extending from the outer end of every
second or third ambulacral ossicle to the border of the abactinal surface; and the
comparative distinctness of the odontophore. The interbrachial septum is mem-
branous with parallel dorsoventral columns of elongate, imbricated plates, the inner
border being concave and strongly reinforced by plates, which are irregular and form
a band much wider than the other columns. Between thé dorsoventral columns
there is, in small and medium-sized examples, an area of unfortified membrane,
upon which the plates encroach more and more with advancing age.

The transverse, parallel septa which are found along the border of the ray are
composed of long blunt processes which grow outward and upward obliquely from
the inner surface of the actinal, inferomarginal, and intermarginal plates, the septum
being continued dorsally and mesially as a membrance with a free concave mesial
border. That part of the septum which joins the abactinal surface is strengthened
by keellike extensions of the dorsolateral plates or their lower and outer ends. The
condition of these septa, which are pretty evenly spaced, will depend upon age and
the general “calcareousness” of the specimen. They extend to the end of the ray,
becoming smaller as the ray attenuates, and are pretty regularly opposite alternate
ambulacral plates. The ventral processes from the dorsolateral plates are a little
more conspicuous in these distal septa.

The odontophore (pl. 4, fig. 6, O) is clearly outlined, in most specimens, from the
pair of enlarged ambulacral ossicles which surround the first ampulla pore, whereas
in Orthasterias, Stylasterias, and allies, it is hidden by tissue so that its contour is not
evident. Its interradial dimension is about 0.45 of the extreme width.

The enlarged summit of the first ambulacral ossicle is separated from the
companion by a broad muscular suture, oval in form, whose length equals that of
the following seven or eight ambulacral plates.

The ambulacral ossicles are slender and long. The portion which forms the
ambulacral ridge is especially compressed and thin, being only half the width of the
intervening muscular sutures and therefore narrower than the corresponding part in
Orthasterias and allied genera. The ambulacral pores are narrow (on the outside)
with flaring sides. There are two rows on either side of the radial line and at the
base of the ray the inner ends of the outer row interdigitate with the outer ends of the
inner about one-third their length; far along ray about one-half. The first ampulla
pore, on the inside, is of conspicuous size.

Internal anatomy.—The viscera do not present any differences from the typical
Asterias arrangement. The gonads, of the racemose type, open dorsally on the inter-
radial line at about the middle of r. The intestinal coecum is divided into two parts
and each of these into about three lobulated subequal divisions, the longest not
reaching the middle ofr. The spacious stomach has strong retractors and the hepatic
coeca are voluminous. The ampullae are obviously single, rather large, and there
are conspicuous Tiedemann’s glands, but no Polian Vesicles.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 19

Forma ANOMALA (Verrill)
Plate 3, Figure 1; Plate 4, Figures 1-3; Plate 6, Figures 5, 6, 11; Plate 7, Figures 2, 2a—2d, 4
Allasterias anomala VeRRILL, 1914.

Cape Lisburne, Arctic Ocean; St. Michael, Norton Sound; Bristol Bay (stations
3228, 3231, 3233, 3234). These specimens, 51 in number, are referable to Verrill’s
Allasterias anomala, the type of which, No. 3821, is one of a series of four dried
specimens from St. Michael, of which I have three. These were collected November
1874, summer 1875, and August 5, 1876, by L. M. Turner. On page 67 (1914)
Verrill states that the species is from Siberia, probably a lapsus.

Two of the St. Michael specimens, in poor condition, belong to forma anomala,
while the third is forma amurensis. The dredged specimens, taken by the Albatross,
are in fine condition.

Anomala differs from the type form in having much thicker and more spatulate
scoop-shaped superomarginal and inferomarginal spines. The subambulacral spines
are rather heavy, in extreme examples, frequently broadened at the tip, and very
definitely grooved. The abactinal spines are short but thick, rather tubercular,
and stouter than in the more generalized form. They are about 0.9 to 1.3 mm. long
and about 0.7 to 0.8 mm. in broadest diameter, therefore often fully two-thirds as
thick as high. But among the thicker spines there are other secondary slenderer
ones. In form they run pretty much the whole gamut of shapes characteristic of
the slenderer spined varieties—cylindrical, subclavate, round tipped, or even sub-
capitate; variously notched or sunken at the end, or without any specual modifica-
tion; often scoop shaped. The skeleton is generally stouter than in the forma
amurensis, and conspicuously heavier than in deep-water examples.

There is, however, a good deal of variation in the stoutness of the ventral
spines. This is evident enough from a series of specimens from stations 3233 and
3234. Examples from stations 3304 (1), 3290 (1), 3244 (1), and 3247 (1) seem to
be about intermediate between this variety and the stouter specimens of the usual
Bristol Bay form, as described in detail. It is worth noting that in a considerable
series, such as that from station 3233, some examples—extremes of the variety—
exhibit a heavy rather rigid dorsal skeleton with fewer stubby spines, while a
minority have a rather flaccid abactinal wall with much more numerous and
delicate spines. The latter also have less spatulate ventral spines and are in fact
quite intermediate with normal Bristol Bay specimens of that size (R about
90 mm.).

The crossed pedicellariae are larger than in the slender-spined forms from the
same region (except those from deep water). Actinal pedicellariae (from specimens
about 90 mm. major radius) range from 0.27 to 0.36 mm., the majority being over
0.32 while the abactinal crossed pedicellariae are around 0.26 and 0.27 mm. The
minor details of the pedicellariae are of course variable, and the breadth is also
slightly variable.

Professor Verrill (1914, pls. 59, 60, 69) has given good photographie figures of
his type of anomala, and on Plate 78, Figure 3, a drawing of the ventral spinulation.
His figures of the pedicellariae are so poor as to be misleading. On Plate 77,
Figures 3, 3a (labeled Allasterias forficulosa by mistake in the explanation, p. 156),

20 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

are shown crossed and straight pedicellariae, and on Plate 78, Figures 4, a to e,
straight pedicellariae. The crossed pedicellaria, Plate 77, Figure 3a, is quite inac-
curately drawn, as it is unlike any found in this form. The straight pedicellariae are
also incorrectly drawn.

The two badly beach-worn examples from Cape Lisburne, Alaska, were picked
up at the “coal station” after a northwest gale of four days, September 11 to 15,
1885, by Henry D. Woolfe. They constitute the only undoubted Arctic Ocean
record and are notable for their thick, mostly cylindrical, round-tipped spines about
0.9 to 1 mm. long and 0.45 to 0.6 mm. thick. The marginal and ventral spines are
also thick, and not particularly different from St. Michael or Bristol Bay specimens.
The straight pedicellariae of the ventral surface (pl. 7, fig. 4) are, if anything, a little
larger than in Bristol Bay examples, many reaching a length of 0.9 mm. (0.65 to 0.9
mm.). They are relatively narrower. The abactinal straight pedicellariae average
0.45 mm., sometimes smaller, and are about the shape of the ventral pedicellariae
of the Bristol Bay specimen, station 3231. (PI. 7, fig. 2.) The ventral crossed pedi-
cellariae are about 0.315 to 0.35 mm. (some as small as 0.23 mm.) and are similar in
detail to the abactinal, which are a trifle smaller (0.28 to 0.3 mm.). Figures of the
Bristol Bay (pl. 6, figs. 5, 5a) and Cape Lisburne (pl. 6, fig. 6) specimens will bring
out the slight differences.

Type.—In the University Zoological Museum, Copenhagen.

Type locality—Amur country (northern part of Gulf of Tartary) eastern Asia.

Distribution —Arctic Ocean, north of Bering Strait; Bering Sea; Okhotsk Sea,
and the Asiatic coast south to northern Japan where the species intergrades with
rollestoni. On the American coast the Shumagin Islands constitute the only certain
record® south of the Alaskan Peninsula. Vertical distribution, low tide to 93 fathoms.
Temperature records 32.2° to 45.5° F., the latter probably several degrees too high.

Specimens eramined.—Eight hundred and sixteen.

Specimens of Asterias amurensis examined

Num-
Sta- ; r Bottom
tion Locality Depth Nature of bottom temper- pera Remarks
ature | Thens
" Fathoms NR
3225 | Northof Unimak Island, Aleutian Islands_| 85 Blackistones :2=--2.=--.-==<- 38. 6 2
3228 | Bristol Bay, Alaska, 58° 39’ 20” N., 157° 17” 8 Gray stones and pebbles--—-_|_______- 1 | anomala.
30” W.
TON ES Eig SS TV poll? “el | Se 12 Stones...... 20 Do
3233 | 58° 23’ 45’’ N., 157° 42’ 45/7 W__ Pi 7.25 | Sand, pebbles-_- 13 Do
8234 | 58°27’ N., 157952’ W__ 22... ssfotie! 5 Gray sand- - 5 3 Dol
3235 | 58° 16’ 30’ N., 158° 13’ W.____. As ll Black'sand-=....----._. 142 ;
8236 a it N37 SNS OD OOo Wc emo sco nsantenes 14.75 | Gravel, sand, shells___- = 89
on eS ns ae — ae ees opthddeeasted 19 Gray sand, gravel, shells____|_.______ 32
238 | 58° 03' 40” N., 158° 37’ 30” W_. 18 Fine gray sand.......___- 15 j (
3239 | 58° 22’ 20” N., 159° 23’ 15” W__ eb eee dovee ne ee a 4 ae?
3240 | 58° 30’ N., 159° 35’ 50’ W_____- 14.5 Fine black sand 8
3241 | 58° 38’ 30’ N., 159° 33’ 30’ W___- 14 Black mud j
sak | 58° 4 90” NC 10° 08 45” W faa ieee Pi Wega) Raia as

* The case of Parasterias albertensis, labeled as from British Columbia, is very perplexing. The species appears to be Asterias

amurensis. There is a possibility that the specimen may have been tr: i i
“ s ansported by ship; or that it may hav: i
a “curio” and labeled in good faith by the donor. , Rae e Sel

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 21

Specimens of Asterias amurensis examined—Continued

Bottom
Depth tem per- Remarks
ature
Fathoms iF.
58° 45’ N., 160° 28’ W 4.5.) Fine gray.eand....... cccestsn]e nes. One-half bushel re-
jected.
Mouth of Bristol Bay, 58° 37’ 20” N., 161° SE ei 00 aio nn cian nenmenaweulpepecees
05’ W.
§8° 26’ 30 N., 161° 36’ W 17.5\| Gravel. 5.-...-.-..<a-caense 38 8 | 1 bushel rejected.
§8° 40’ 45’’ N., 162° 08’ 307 W_- . 7 Pebbles, stones.....---- -| 40.6 1
§8° 34’ 15’ N.., 162° 22’ W 21 Fine gray sand, gravel. --..- 43 16
58° 27’ 30’ N., 162° 36’ W.--- 13.5 | Fine gray sand, broken shells} 37 15
68° 11/ 30” N., 163° 02’ 45” W 17.6} Gray sand............-..--- 46.2 6
Bering Sea, 57° 35’ 50’ N., 164° 05’ W- 25.5 37.5 2
North coast of Unimak Island 81 39 M“
Be Tere is eat 40.6 27 | About 100 rejected.
13. | Fine black sand-....-. 42 3
27 41.2 2
40 40.5 5
88 | Black sand............. 39.8 8
IN ari OO saectoe piesa a 42 5
aT | Neeteprel 41 7
Bb eee O00 co a eeen en 41.9 13
31 42 8
39 Gray sand, mud---.-- -| 38.5) 5
North of end of Alaskan Peninsula; 55° 34’ 19 Black sand, shells........---|.-.----- 1
30’ N., 162° 31 45” W.
47 +‘| Fine gray sand.........----- 1
36 ©6| Gray sand, black specks -- -~|..-.---- 1
85. ‘}i.-.-00- 55-2 - eee 3
North of Alaska Peninsula, mouth Bristol 37 4| One-half bushel re-
Bay. jected.
North of Alaska Peninsula, 56° 33’ N., 30 | Coarse black sand.......-.-- 43 | 1 peck rejected
160° 14" W.
North of Alaska Peninsula, 56° 26’ 30 N., 15 | Black gravel_.......-------- 3
160° W.
North of Alaska Peninsula, 56° 50’ 30” N., 16 14 | 2 bushels rejected.
26 =| Black sand, gravel _._..-.--- 21
Bae be eae cena laate enn 8 | One-half bushel re-
jected.
30 3 Do.
30 3
1.5 3 Do.
24 7 | 1 bushel rejected.
26 5
20 14 | 1% bushels rejected.
20 Fine gray sand, 2 Do.
specks.
15 | Reb DeG soe n cae es aa 1| One-half bushel re-
jected.
WwW 6 | 1 bushel rejected.
30 2| One-half bushel re-
jected.
1 Do,
6
BB) lava ane en conestacers= 1 Do.
58 2
85 | Green mud......... 4
3333 i a | 2
BUM cou) 5
3335 Bh ial cts .2 6. ste 4
3508 | Bering Sea, 58° 33’ N., 169° 49° W....-..-- 3 Fine gray sand, shelis.- -| 4 16
3520 | Bering Sea, 50° 28’ N., 170° 577 W......... 3s Green mud, fine sand. ...... 32.2 1

22 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Specimens of Asterias amurensis examined—Continued

| |

| Num-
Sta- i h Nature of bottom rs ber of Remarks
tion Locality Dept | Seas oi ature | Spee S
= = = ise
Fathoms F.

3538 | Bering Sea, vicinity of Pribilof Islands--_-. 59 | Green mud, sand --- ae 38 3
3539 | Bering Sea, east of Pribilof Islands--..----- | 57 38.9 1
3547 | Northeast of Unalaska.--.......----- | 51 45 1
8777 | Off Kamohatka.........-..-.---.... 13 | Sticky soft green mud-------|.-.----- 10

Norton Sound, 64° 12’ N., 162° 25’ W..-.-- Ai aay SRR aL ey 1| No. 15809,

Cape Lisburne, Alaska, Arctic Ocean.....|_---.....]----....---..----.----------<- Lee 2| Henry D. Woolfe,

| anomala.

5 | No. 7621, Murdoch.

3 | No. 10999.

3] No. 2821, L. M
| Turner.
Hagemeister Island, Bristol Bay. -....-.-.|--------- 1| W. H. Dall.

Tliuliuk Harbor, Unalaska -- --....- 2 Do.
Popoff Strait, Shumagin Islands -- 5 | No. 3452.
Unga Island, Shumagins_--......- 1] W. H. Dall.
Petropavlovsk, Kamchatka--....- 13 | Albatross, 1906.
Port Clarence, Alaska-...........-- sy ee 2 | Stanford University.
ATHUTAETOVINOG; AMIR soe oe ae cotati hen ee | 1 | Cotype, Copenhagen
| | Museum.

1 Low water.

Remarks.—The most conspicuous points of difference between A. rubens and A.
amurensis have already been listed. I have examined specimens of rubens from
Iceland, Faroe Islands, Dublin Bay, Ireland, Port Erin, Isle of Man, Farsund, Norway,
and Kristineburg, west coast of Sweden. The species A. rubens, like amurensis, is
extremely variable, and certain variations approach fairly close to amurensis. The
general run of specimens, however, is rather easily distinguished. A. rubens
resembles more nearly A. amurensis rollestoni of Japan, but I have never observed
specimens of rollestoni as large as rubens grows to be.

It is to be noted that the furrow spine of alternate adambulacral plates is, in
rubens, advanced into the furrow, as in amurensis. The genus Allasterias, for A.
rathbuni (nearly typical amurensis) was based upon this character and consequently
becomes a synonym of Asterias, ss.

Parasterias albertensis Verrill, 1914, is said to have been taken at Albert Bay,
British Columbia (no. 5537, Yale Museum). I have examined the type which is a
small Asterias having the characteristic arrangement of adambulacral spines and
marginal plates. The specimen is small (R 54 mm.) and as is well shown by Verrill’s
figure (1914, pl. 57, fig. 1), the abactinal skeleton is rather irregularly reticulated, an
irregular carinal series being clearly discernible. The abactinal spinelets are mostly
one to a plate, short, slightly tapered, blunt with rough tips. Supermarginal spine-
lets, faintly capitate, proximally three, distally two, somewhat longer and stouter
than abactinals; inferomarginals two, longer and slenderer, tapered; sometimes one
or two isolated intermarginals. Adambulacrals on proximal part of ray alternately
two and three spined; distally, diplacanthid. The triplacanthid plates have the
inner spine set deeper in the furrow. Abactinal straight pedicellariae small, numer-

ous, the larger about 1.5 the length of the crossed; actinal, large, numerous, similar to
the slenderer sorts of typical amurensis.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 23

If this specimen really came from British Columbia, which is somewhat doubtful,
it is a form of amurensis. The southern-most record of amurensis is Popoff Straits,
Shumagin Islands. It is rather odd that, in spite of all the collecting that has
been done in Alaskan waters by the steamer Albatross, no specimens have been taken
anywhere near British Columbia. Yet we have the curious case of Leptasterias
polaris katherinae which is founded upon “‘chance’’ specimens.

Professor Verrill remarks that the ‘species looks more like the typical species of
Asterias of the North Atlantic than any other west coast species known to me.”’
This is partly due to the small size. The young of amurensis all have the narrower
rays typical of A. forbesi, for instance.

Allasterias forficulosa Verrill® is a form of Asterias amurensis rollestoni Bell.
Professor Verrill compares it with Alaskan specimens, but does not say wherein
it differs from the common Japanese form. I have examined the type.

T have also examined the types of Sladen’s Astercanthion rubens v. migratum" from
the Straits of Korea. The largest has R only 17 mm., and is consequently too small
to compare with adult specimens. It is likely a form of amurensis. A. amurensis
might be expected to occur in the cold area of the Straits of Korea.

I have examined examples of Bell’s Asterias rollestoni from Hokkaido (Hakodate
and Mororan, on the southern coast); Aikawa, Rikuzen; Suruga Bay. While these
specimens are variable, they agree in having somewhat narrower rays, more widely
spaced abactinal spines, and especially in having smaller crossed pedicellariae. The
inferomarginal pedicellariae of the Mororan example range in length from 0.17 to
0.23 mm., with a relatively few giants attaining 0.26 mm. In a comparable Kam-
chatkan specimen, with R 130 mm., the same pedicellariae measure 0.315 to 0.34
mm., with a minority attaining 0.36 mm. (PI. 6 fig. 3a.) The type of amurensis
is intermediate but nearer to the northern extreme. There is little doubt that
perfect intergradation exists between A. amurensis and A. amurensis rollestoni.

Genus LEPTASTERIAS Verrill

Leptasterias Verritt, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 350. Type, Aster-
acanthion miilleri Sars; Shallow-water Starfishes, 1914, p. 116.—Stapen, Challenger
Asteroidea, 1889, p. 563 (subgen.).—Perrier, Expéd. Travailleur et Talisman, 1894,
p. 108.—Fisner, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 599.

Ctenasterias VerRILL, Shallow-water Starfishes, 1914, p. 148. Type, Asterias spitzbergensis
Danielssen and Koren (=Leptasterias groenlandica f. cribraria).

Diagnosis.—Asteriinae having adambulacral spine pedicellariae, as in Asterias,
but differing in having gonads which open in the actinal interradial angle, usually
on a low papilliform prominence, two to each interradius; eggs relatively few and large
(much yolk); the small species known to brood eggs and young (in one case in the
stomach); abactinal skeleton more or less open, the dorsolateral skeleton usually
irregular, but the spines sometimes in poorly defined longiseries; spines usually
small, normally with a small or fairly thick collar of pedicellariae; actinal plates in
one or two series, sometimes feebly developed; adambulacral plates diplacanthid,
or mixed diplacanthid and monacanthid, or exceptionally mostly monacanthid;
adambulacral pedicellariae normally in clusters on the spines, occurring singly, or

19 1914, p. 104, type No. 1183, Mus. Comp. Zodl,, Japan. « Journ. Linn. Soc., vol. 14, 1879, p. 432

24 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

very sparingly in individuals which have very few: pedicellariae without enlarged
teeth on the moderately broad terminal lip; straight pedicellariae variable—small
to large—lanceolate to broadly ovoid, or even clam-shaped, edentate to unguiculate.

Subgenus LEpTASTERIAS Verrill

Leptasterias miilleri group; size small, symmetry pentamerous. North Atlantic,
north Pacific, Arctic.

Includes the following species *: arctica (Murdoch), austera (Verrill), jfishert
Djakonov, groenlandica (Liitken), hylodes Fisher, hyperborea (Danielssen and Koren),
leptalea Verrill, littoralis (Stimpson), miilleri (Sars), ochotensis (Brandt), orientalis
Djakonovy, tenera Stimpson.

LEPTASTERIAS ARCTICA (Murdoch)
Plate 8, Figures 5, 5a-5h; Plate 9; Plate 10; Figures 1-4; Plate 14-16

Asterias arctica Murvocu, Report of the International Polar Expedition to Point Barrow,
Alaska, under Lieutenant Ray, 1885, p. 159——Von Horsten, Die Echinodermen des
Risfjords, Kiing. Svenska Vetenstapsakad. Handl., vol. 54, no. 2, 1915, p. 63.

Asterias nautarum BrLu (nomen nudum), Sladen, Challenger Asteroidea 1889, p. 824
(nomen nudum). See Fisaer, Ann. and Mag. Nat. Hist., ser. 9, vol. 18, 1926, p. 198.

Aslerias miilleri Lupwia, Fauna Arctica, vol. 1, 1900, p. 482.

Asterias hyperborea Verritt, Amer. Nat., vol. 48, 1909, p. 553.

Leptasterias arctica VERRILL, Shallow-water Starfishes, 1914, p. 120, pl. 56, figs. 1, 2; pl. 71,
figs. 1, 2; pl. 72, fig. 1; pl. 83, figs. 2, 2a—Ctuark, Rep. Can. Arct. Exp., vol. 8, part C,
1920, p. 9¢ (Dolphin and Union Strait, Northwest Terr.)

This highly variable species is the equivalent of ZL. miilleri (Sars) to which it
is closely related. Its status with reference to miilleri is the same as that of Asterias
amurensis to A. rubens. In neither case is intergradation known, although very
likely it occurs. JL. arctica has two fairly well-marked formae. The northern or
typical form is much less like miilleri than the larger Bering Sea race. It has been
suggested * that L. hyperborea (Danielssen and Koren) is a high arctic form of miilleri.
It is noteworthy that the arctic phase of arctica (forma artica) is not like hyperborea.
If L. arctica intergrades with miilleri it does so by a series of forms which are less
like miilleri than its own southern form, beringensis. L. arctica forma arctica stands
geographically between beringensis and miilleri.

Diagnosis.—Forma arctica: Rays five; size small to medium. R=4 to 5+r in
fully grown examples, or 3.5 r in small specimens. Largest of type series, R 36 mm.,
r8mm.,R=4.5r. Disk small, rays slightly swollen at base, gradually tapered to
a blunt extremity. Rays well arched, with curved lateral face. Spines coarse,
unequal in diameter abactinally, subcapitate, striate, surrounded by a wreath (often
prominent) of crossed pedicellariae; carinal spines more crowded than the irregularly
spaced but uncrowded dorsolaterals, usually somewhat larger, and arranged in a
zigzag series distinguishable in the dried state at least. Superomarginal, infero-

"Includes possibly Asteracanthion mexicanum Liitken (Leptasterias mexicana, Verrill, West Indian Starfishes, Bull. Stat»

Univ. Iowa, Nat. Hist. vol. 7, no. 1, 1915, p. 24). Too little is known of this form to venture an opinion. As Verrill says, it may

be a young Asterias. In this work two other Species are referred to Leptasterias by Verrill, namely, Asterias fascicularis Perrier
which ts a Tarsaster and Leptasterias hartii Rathbun, an Allostichaster.
See von Hofsten, 1915, p. 62, for summary.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 25

marginal and actinal series normally quite regular; sometimes a short intermarginal
series of spines; adambulacral spines alternating irregularly one and two; unguiculate
straight pedicellariae in the intermarginal and lateral channels and also sometimes
abactinally and on proximal furrow spines.

Description —There are 42 specimens of Mr. Murdoch’s type series from 13.5
fathoms 10 miles west of Point Franklin, Aretic Ocean (No. 7625 U.S.N.M.), ranging
in size from R 4 mm. to R 36 mm. Forma beringensis attains R 73 mm., station
3442. The abactinal spines are coarse in relation to the size of the ray, not fine;
not uniform in size; very distinctly spaced, not crowded. Neither are they regularly
arranged in either transverse or longitudinal series, But there is a very distinct,
irregular, even zigzag, radial row of larger spines in most of the specimens and a
certain linear arrangement may be noted in the spinelets adjacent to the supero-
marginals. The spines are short and decidedly blunt; the larger, about as high as
broad, are subcapitate, with evenly rounded or subtruncate, finely striated tips;
the smaller are less capitate, cylindrical or slightly swollen with fewer striae and
less regular tips. The spines are surrounded at mid height, or near the tips, by a
wreath of crossed pedicellariae, the numbers of which are subject to individual and
locality variation. They are more numerous in forma beringensis. In some aleoholic
specimens the crowns of pedicellariae nearly or quite touch but not in the types.

The superomarginal spines are distinguishable by their regularity rather than
by their slightly greater length. They are clavate, finely striated and relatively
slenderer than the radial spines, as well as actually slightly longer. The series curves
upward to the interradial angle and the intermarginal furrow is here a bit wider,
and sometimes occupied by a short series of intermarginal spinelets which arise from
the descending lobe of the plate. The character is variable. Rarely there is also
a spine on the upper lobe of the plate, or a transverse series of three; station 3559.
There are no intermarginal plates.

The inferomarginal spines are still longer, and a little stouter than the supero-
marginals, sometimes a little bent upward, subterete to clavate in form (in variants
sometimes compressed, and much more robust than in the type variety). Both
series have rough-tipped spines, and cireumspinal wreaths of pedicellariae (or half-
wreaths on the inferomarginals of small specimens). There is one short series of
smaller, but similar, actinal spines. The length of the series varies with age; in the
largest Point Franklin specimen it is half the length of furrow. In the largest examples
the series reaches two-thirds to three-fourths the length of the ray.

The adambulacral spines are close-set, slender, terete, blunt, and stand one or
two to a plate in irregular alternation. The proximal 8 or 10 plates are usually
monacanthid, sporadically diplacanthid. About three post-oral adambulacral plates
are in contact. These apparently always have but one spine each. The inner of
the two adambulacral spines is generally slenderer and is bent over the furrow in some
examples. It seems to be the only one which regularly carries straight pedicellariae
(very variable in number, and absent in small specimens). Both spines have clusters
of crossed pedicellariae. Some of these differ in detail from the abactinal crossed
pedicellariae, as the jaw when viewed in profile possesses a longer terminal expanded
lip, with more teeth. (See pl. 10, figs. 2, 2a, 3, 3a.) The pedicellaria is rather more
tapered toward the end.

26 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The mouth plates are small and carry three spines, the innermost quite small and
at the inner furrow corner of the plate; the middle spinelet is a little longer and at the
inner suture corner, but not so close to the actionostome; the third is 50 per cent
longer and situated at about the middle of the actinal surface of plate. All bear
broadly lanceolate straight pedicellariae similar to those on the furrow spines. In
some large examples from other localities (as station 3543) the jaws may have two
or three conspicuous teeth, as has the adambulacral and other straight pedicellariae.

The papular areas are small with one to four papulae on the rays, or occasion-
ally slightly more. There is an intermarginal and one actinal series. (In the largest
specimens the papulae are more numerous.)

Straight pedicellariae: More or less markedly unguiculate, slender, somewhat
hand-shaped pedicellariae occur, in forma arctica, very sparingly on the abactinal
surface, in the interbrachial channels, and in the intermarginal channels. (PI. 9,
fig. 8.) Just external to the adambulacral spines and on the oral and proximal fur-
row spines, there are usually a few but they are not so numerous as in forma bering-
ensis. Ovate-lanceolate pedicellariae are common along the furrow face of the
adambularcal plates. These have the tips of the jaws crossed, or one curved tip
meets a notched tip. In southern Bering Sea specimens (forma beringensis) the
unguiculate pedicellariae (pl. 9, figs. 10, 11) are sometimes fairly common abactinally,
but not in forma arctica. The largest are in the axils of the rays, actinally, and in
beringensis these may have as many as five teeth. (Station 3543, pl. 9, fig. 9.)
While the dimensions vary a good deal, an axillary pedicellaria of a large specimen
is 0.8 to 0.9 mm. long.

Crossed pedicellariae occur in wreaths around the abactinal and marginal, and
as half wreaths on the actinal and adambulacral spines. In a specimen with R 36
mm., forma arctica, the abactinal are about 0.23 mm. long and the adambulacral
0.26 mm., most of the latter having a different shape from the former. (PI. 8, figs.
5, 5a-5h; pl. 10, figs. 1-3.)

Madreporic body small, but variable, about 0.6 r from center of disk, and with
three to six spines on the adcentral margin.

Variations—There are at least two recognizable intergrading formae.

Forma ARCTICA (Murdoch)

Plate 8, Figures 5, 5a-5d; Plate 9, Figures 2, 7, 8; Plate 10, Figures 2, 2a; Plate 14, Figures 1-4;
Plate 15, Figure 1; Plate 16, Figure 1

The typical variety of this species, as indicated by the type series of specimens,
is a fairly slender rayed, small sea star with rather coarse, unequal subecapitate abac-
tinal spinelets, among which a very irregular radial series is discernible by reason of
the greater size and usually more crowded arrangement of spines. In the type
series the crossed pedicellariae are not very numerous on the abactinal spines, but
this is a character which appears to vary with every station, and there are even con-
siderable individual differences at the same station. Examples from 15 to 20 fathoms,
Plover Bay, Siberia, and from East Cape, Siberia, are more plentifully supplied than
are those from the American coast, and therefore resemble forma beringensis. But
the crossed pedicellariae are more nearly like those of arctica, and the well-spaced

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 27

dorsolateral spinelets are much smaller than the carinals and superomarginals. This
may represent a ditinct forma ranging along the Arctic shores of Asia.

The straight pedicellariae are rather slenderer and have less well-developed teeth
than have some of the stout spined beringensis. They are very scare abactinally in
the type series; but in larger examples from Plover Bay and Cape Lisburne con-
spicuous straight pedicellariae of characteristic form are scattered here and there over
the dorsal surface. They are slender, incipiently toothed, and as long as the second-
ary spines. ,

The largest Arctic specimen, from Cape Lisburne, has R 50 mm., and coarse
well spaced, strongly capitate, striate, unequal dorsal spinelets.

The specimen from Nikolski, Bering Island, with R 39 mm., has heavy, capitate,
subtruncate, abactinal spinelets and closely resembles the typical Arctic form. The
marginal, actinal, and adambulacral spines are rather more robust than in any of
the Point Franklin examples. (Pl. 14, fig. 4.) It is of interest to find that forma
arctica extends so much farther south in the western part of Bering Sea than it does
in the east. In the latter region it does not range south of latitude 60°.

Forma BERINGENSIS, new forma

Plate 8, Figures 5f-5h; Plate 9, Figures 1, 3, 4-6, 9-12; Plate 10, Figures 1, 3, 4; Plate 14, Figures
5, 6; Plate 15, Figures 2-5; Plate 16, Figures 2-6

The bulk of the collection comes from the vicinity of the Pribilof Islands, from
Bristol Bay, from off the north coast of the adjacent Alaskan peninsula, and of
Unimak Island. Perhaps the most evident feature of these specimens is that they
average much larger in size, although of course different growth stages are abundantly
represented. Many of them have thick rays due usually to swollen gonads, although
in specimens from the same dredge haul there is much difference in the thickness of
the rays.

As a fair sample of the variations of specimens from the vicinity of the Pribilof
Islands, 20 examples from station 3536, 40 fathoms, may be taken. Two specimens,
each with R 58 mm., have r 11 and 12 mm., but the breadth of ray just beyond the
base is 16 or 17 mm., and 19 mm., respectively. The swelling in the latter fairly
extending toward the extremity. The abactinal spines are robust and have capitate,
finely striated tips. There is much variation in the robustness and degree of spacing
of the spines. In one variety the spines are fewer and widely spaced. Counting
across the abactinal surface between the superomarginal series of either side, there
may be only three series of major spines together with a few much smaller ones; or,
on another ray of the same specimen, seven series. In a more normal, slender-rayed
example of approximately the same size there are 10 or 12 spines across the base
of the ray, but there is scarcely any arrangement in longiseries with the exception
of the irregular carinals where the spines are usually distinguished by their slightly
to decidedly larger size, and more crowded position. The sparsely-spined variety
occurs at a number of stations, as, 3277, 3280, 3291, 3293, 3294, 3297, 3302, 3303,
3306, 3438, 3504, 3511.

The marginal and actinal spines are fairly typical and stand in regular series,
generally only one to a plate, but there is considerable variation in the details of

28 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

form, and the relative size of the supero and infero marginals. The latter are more
variable inter se than the former. They are usually a trifle to decidedly larger than
the superomarginals, varying from cylindrical to a subspatulate form, almost always
swollen toward the tip, which is rounded or truncate, or slightly bent upward. The
actinals are similar, but a little less heavy. The adambulacral spines ordinarily
alternate one and two, and are terete, much slenderer than the actinals, and less
variable as to form. ‘The inner or furrow spine is slenderer than the outer, while
the alternate single spine is about intermediate in size. One specimen, on the
proximal half of ray, has two spines on nearly all the plates. The largest specimen
in the collection, station 3442, with R 73 mm., has only one adambulacral spine to a
plate. Crossed pedicellariae are very numerous.

The crossed pedicellariae are more numerous and larger than in the forma
arctica (measuring about 0.27 mm. in length), but in this abundance there is consid-
erable latitude. The abactinal spines, however, are provided with rather thick
sheaths which usually reach nearly to the tip, and are there beset with one or, on
the dorsolateral regions, two circles of pedicellariae, while the marginals may have
three or four circles. On the inferomarginals there are often only half circles, and
the actinals regularly have only half circles on the outer side. For different varia-
tions see Plate 14, Figures 1, 2; Plate 15, Figures 3 to 5; Plate 16, Figures 2 to 6:

Unguiculate straight pedicellariae are scattered rather sparingly over the abac-
tinal surface but are fairly numerous and somewhat larger in the intermarginal
and actinal channels, and in the interbrachial channel. Considerably smaller
pedicellariae, with rather narrower jaws and smaller teeth are found sparingly on
the proximal furrow spines. But 27 examples from station 3559, 39 fathoms, gray
sand, vicinity of Pribilof Islands, while exhibiting individual variation, have these
straight pedicellariae of the furrow spines much more numerously represented,
even in small specimens. Straight pedicellariae are also more numerous on the
body surface. The same is true of station 3543. (See pl. 9, figs. 9, 10, 11.) While
the dimensions vary a good deal, an abactinal pedicellaria of a large specimen is about
0.9 mm. long, an axillary 1 mm., a suboral 0.6 mm., and a furrow 0.5 mm.

One of the specimens from station 3559 (pl. 15, fig. 1), intermediate with forma
arctica, R 56 mm., is remarkable for having the superomarginal spines in a vertical
arcuate series of generally three, or two on the distal third of the ray. The lowest
spine stands on the descending lobe of the plate and would be classed as one of an
intermarginal series which extends two-thirds or three-fourths the length of the
ray. This gives a very spiny aspect to the lateral face of the robust ray. The
spines of the radial region are coarse and abnormally crowded, the adambulacral
spines are heavier than usual and clavate, truncate, while the crossed pedicellariae
are unusually numerous. The other specimens have only a single superomarginal
spine and are slender rayed.

Station 3294, Bristol Bay, 30 fathoms, black gravel, July 18, 1890.—The greater
number of the 14 large specimens have the abactinal spines more widely spaced than is
usual although the appearance is accentuated by the turgid rays (swollen by large
gonads). The relatively few straight pedicellariae are slenderer and lack teeth as a
rule. These, however, are present on pedicellariae from specimens taken at 3295,
11: fathoms, fine gray sand, same general locality. The more sparsely spined

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 29

examples from station 3302 (fine gray sand, 30 fathoms) also have toothed straight
pedicellariae, but they are not very numerous. In fact the animals from the southern-
most part of the range seem to be poorly provided with straight pedicellariae (except
on the furrow face of the adambulacral plates) but there are exceptions to this, for a
large specimen, with R 70 mm., and well spaced spines (five to seven across the
dorsal area) from station 3277 (18 fathoms, gray sand and rocks, north of the end of
the Alaskan Peninsula) has numerous dorsal, lateral, and actinal straight pedicellariae
which either lack or have few short teeth. This specimen also has numerous furrow-
spine straight pedicellariae, and the largest rays recorded for this species. Three
other examples from the same station have much fewer pedicellariae and agree with
the generality of material from the region.

One of the examples from station 3536 has six rays.

Aleutian Island form.—Specimens from the Aleutian and Shumagin Islands
differ from typical beringensis in their smaller size, rather more terete and less obvi-
ously capitate dorsal spines. Straight pedicellariae are rather variable but in a
specimen from Atka they are well developed with three or four curved teeth to each
narrowly spatulate jaw. The largest specimens have R 35 mm. (Shumagin Island)
and 36 mm. (Atka). As compared to forma arctica, for example, from Point Frank-
lin, the Aleutian-Shumagin examples have more widely spaced, more uniform,
and longer spines, with much more conspicuous wreaths of pedicellariae.

Young.—The young forecast some of the more salient variations of the adults and
therefore differ among themselves, although less widely than do the mature examples.
There is a good series of growth stages from the type locality, the smallest specimen
having R 4 mm. In this there is a distinct lateral face to the ray formed by the
phanerozoniate marginals (about eight) and the flat dorsal surface has only an
irregular carinal series and a rudiment or two of the dorsolaterals. The dorsal plates
bear one or two spinelets and a few crossed pedicellariae have appeared. The
actinal plates have not yet appeared while the adambulacral plates are relatively
more prominent than in larger specimens, with one or two rather slender clavate
spines, relatively much longer than in the adult. A regular series of dorsalaterals
next appears (R 7 mm.), while the crossed pedicellariae increase in number. There
are none on the adambulacral spines which may be only one to a plate throughout
the ray. The first straight pedicellariae have appeared in the furrow, but the larger
type are found first in a specimen with R 9mm. Here there are one or two in the
interradial groove on the actinal surface where in adult life thay appear to guard the
gonoduct apertures. In this specimen also the first adambulacral crossed pedicel-
lariae occur on the proximal spines (in this specimen generally one to a plate). By
the time R reaches 12 or 15 mm. the animal resembles the adult but the spines are
slenderer, the pedicellariae much fewer, and the actinal plates either absent or rudi-
mentary. Naturally also the plates and spines are less numerous than on the larger
specimens.

Brooding habits—Specimens from the following stations are carrying young:

Station 3240, Bristol Bay, June 8, 1890, temperature not recorded; one specimen.

Station 3241, Bristol Bay, June 8, 1890, bottom temperature, 38° F.; one

specimen.
63160—30——-3

30 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Station 3251, north coast of Unimak Island, June 14, 1890, bottom temperature
37.5° F.; three specimens.

Station 3255, north coast of Unimak Island, June 14, 1890, bottom temperature
37° F.; two specimens.

The crucial temperatures for the development of the embryo are therefore in the
neighborhood of 37° and 38° F.

In size the specimens range from R 39 mm. to R 60 mm.

A brood pouch is formed by the strongly arched disk so that the basal portion of
all five rays is brought together and the adjacent marginal spines interdigitate. The
cavity thus formed is crammed with tiny sea stars which are attached to a common
central mass by a long slender strand of tissue proceeding from the actinal interradial
area just external to the mouth plates. This thread of tissue may be as long as the
diameter of the sea star (2.5 mm.) or considerably longer. It enters the central
mass which appears to consist largely of the twisted and tangled ends of these elongate
larval organs. The surface of the mass shows numerous soft lobes, some of which
appear to be the ends of larval organs, as one may tease the mass apart. It then
shows a fibrous structure. There seemed to be about six or seven of these centers of
attachment in the mass of tiny sea stars remeved from a specimen from station 3251.
In other words the mass was divisible into seven smaller clusters. The young were
closely crowded into the temporary brood cavity, and were pressed closely against
the actionstome. (Pl. 10, fig. 4.)

Anatomical notes, forma beringensis—Skeleton: There are three quite regular
series of plates on the ventro-lateral region of the ray, the superomarginals, infero-
marginals, and actinals, graduated in size from larger to smaller in the order named.
The superomarginals are prominently four lobed, the descending lobe overlapping the
ascending lobe of the inferomarginals, which tend to suppress their ventral lobe as
they are crowded by the similar actinal plates. There are fair-sized regular inter-
marginal papular areas, but the actinal series of papular areas may be suppressed in
slender-rayed specimens; or in thick-rayed examples two incomplete series are present,
the inner between the actinal and the adambulacral plates.

The carinal plates (pl. 9, figs. 1, 2, 3) with three to five short irregular lobes, are
larger than all but a few dorsolateral plates and are crowded into an irregular zigzag
series. The plates strongly overlap one another in the radial direction, without any
intermediate ossicles, and laterally overlap the dorsolateral plates which in shape are
wregularaly elongate or incipiently lobed and form a very uregularly meshed skeleton,
as in L. polaris katherinae, L. p. acervata, and other species. A variable number of
dorsolateral plates with three or four lobes, similar in form to the carinals are larger
than the others, and are, like them, the most external of the skeleton. A maximum of
three very irregular series can be traced on either side at the base of ray. The mesh
is completed by smaller plates without lobes, or with less obvious ones. Different
Specimens vary in respect to the robustness of the dorsal skeleton. (PI. 9, figs. 2, 3.)
If the plates average slightly thicker, the aggregate result is smaller papular areas.
The ambulacral ossicles are fairly large and typical of the genus, the two series of
pores on either side being fairly close, since the channel is relatively narrow. Cor-
responding to the first 10 inferomarginals there are 28 or 29 adambulacral plates.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 31

The actinostomial frame, composed of the combined first ambulacral ossicles
plus the five interradial odontophores internally, and the five pairs of mouth plates
externally, is stout and the actinostome is small—scarcely twice as broad as the suture
length of the first ambulacral element. The latter is prominent and the radial suture
is as long as the six succeeding plates measured on the radial line—a rather common
proportion in the less specialized Asteriidae. The odontophore (the interradial plate
to which the first ambulacrals, the orals, and the interbrachial septum are attached)
is considerably broader than the interracial dimension. (PI. 9, fig. 5.) The inter-
brachial septum consists of large irregular plates, sometimes completely filling the
membranous septum, sometimes leaving irregular uncalcified spaces.

There are no lateral pockets of the coelom formed by keeliike extensions of the
inner surface of the actinal and marginal plates. The inferomarginals have rather
long dorsal lobes, plane on the coelomic side, which underlie the descending lobe of
the superomarginal; the actinals have a similar but shorter dorsal lobe. There is a
rudimentary second series of actinal plates visible from the inside in large specimens,
even when not evident externally.

Internal anatomy.—The gonads when mature are large, the testes being larger
than the ovaries, and sometimes reaching nearly to the end of the ray. The ovaries,
from the very limited number of examples examined, appear to be more compact,
shorter, with a few divisions, the surface irregular from many lobules. The testes
consist of a long central axis, giving rise on the outer side to numerous short compact
very irregular lobes and on the inner or ambulacral side to six or eight long slender
subdivisions which are provided also with irregular lobules. The gonads are attached
to the wall of the ray on the inner side of a superomarginal plate, or just below it,
and exactly opposite the interbrachial angle. The short slender gonoduct turns
downward and passes outward very close to the interradial line, between the ascend-
ing lobes of the first two inferomarginal plates. In a large specimen from station 3442,
August 3, 1891, the prominent subspherical genital papillae occur in pairs in each
actinal interradial area, in line with the actinal plates. A swelling, probably the duct,
leads downward to each papilla, which has a conspicuous terminal aperture.

The stomach is spacious with strong muscular attachments to the ambulacral
ridge. The hepatic coeca reach nearly to the end of the ray but the intestinal coecum
is small, with two short blunt lobes, each usually subdivided at the end into two.

Type.—Cat. No. 7625 U.S.N.M. Type of forma beringensis, No. E1489 U.S.N.M.

Type locality —Ten miles west of Point Franklin, Alaska (Arctic Ocean), 13.5
fathoms, sand, August 31, 1883 (42 specimens). The type locality of forma bering-
ensis is station 3277, Bering Sea, latitude 55° 58’ 45’’ N., longitude 161° 46’ 40” W.,
18 fathoms, sand, gravel, rocks, bottom, temperature, 43.2° F.

Distribution —From southern Bering Sea and Aleutian Islands to Aretic Ocean,
eastward to Dolphin and Union Strait, Nortwest Territories; and from low tide to
43 fathoms.

Forma arctica. From Cape Franklin to Bering Straits in the Arctic Ocean, thence
south in the eastern part of Bering Sea to about latitude 60° where it very gradually
intergrades with forma beringensis. On the west side of Bering Sea, south to the Com-
mander Islands (lat. 55°). Vertical range 5 to 21 fathoms.

32 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Forma beringensis. Southern and eastern Bering Sea (south of about lat. 60°),
Shumagin Islands and Aleutian Islands, low tide to 43 fathoms; temperature range

35° to 46.2° F. Carrying very small young at 37.5° and 38° F.

Specimens ecamined.—Four hundred and ninety-three.

Specimens of forma arctica examined

Locality

Remarks

Point Franklin, Arctic Ocean_......-

Cape Lisburne, Arctic Ocean.

ATOR) (COED on napaneeeneens dane one

East Cape, Siberia
Plover Bay, Siberia '.

Off Port Clarence , Alaska...........

Bering Strait

Indian Point, Bering Strait..........
Bering Sea, 63° 37’ N., 165° 19’ W.-_.

Bering Sea, 61° 03’ N., 167° 55’ W_
Bering Sea, 60° 22’ N., 168° 45’ W_..

Bering Sea, 60° 15’ N., 167° 48’ W__.-
Vicinity Bering Strait, 65° 25’ 28” N., 171° 11’

26 W.
Station 3514, 59° 22’ N., 168° 21’ W..

Nikolski, Bering Island. ...........-.--------
Station 3518, vicinity St. Matthew Island,

60° 28’ N., 171° 42” W.

North coast Siberia, latitude 67° 07’ N., longi- | 13-15 Stones with clay_-

tude 173° 24’ E.

Num-
| Depth | Nature of bottom beret
| mens
|
Fathoms
paeeeee 155) | Saude eee e ens 42
1
cress 1
1
15-20 10
ene anaes 3
eee een ae 4
3
Sameera! 1
Seesancoy 1
1
6
Ree ees . 1
G0 Pee 1
1
1
1
2
1 Do.
1 Do.

Vega expedition.

1 The young specimens (No. 16584) from 10 to 15 fathoms recorded by Verrill, 1914

eggs in the stomach.

Specimens of forma beringensis examined

Type series No. 7625, John Murdoch.
No. 3628, W. H. Dall.

No. 3622 (1669), W. H. Dall.

No. 6031, Dr. Robt. White.

No. 3576 (1650).

No. 3558 (1635), W. H. Dall.

No. 7627, John Murdoch.

Nos. 1357, 1359, Wm. Stimpson.

No. 3582, W. H. Dall.

Geo. M. Stoney.

No. 15688, Geo. M. Stoney; intermediate.
Geo. M. Stoney.

U.S. S. Corwin.

Bottom temperature 40.8° F.; intermediate.

Bottom temperature 33.9° F.; intermediate.

p. 123, prove to be L. groenlandica, brooding

Sta-

tion Locality

1 Probably incorrect; too high.

Pebbles, stones.
Gravel, sand, shells
Fine black sand

Black mud

Bottom
Nature of bottom temper-
ature

Black Mit. 35 .es2e2--<4 38

ia same CON ee nameneecce ee eee eee
Pebbles, stones__..........-- 40.6
Gray sandy ss 2 2. es 146.2
Fine gray sand_.....-......- 37.5

Num-
ber of
speci-
mens

Remarks

Some intermediate.

Aberrant.
Do.
Do.
Do

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 33

Specimens of forma beringensis eramined—Continued

a Locality roa Dept n |
'

Pathoms |
3273 | Northeast of Unimak---...-....--..--..- 30
3277 | North of end of Alaska Peninsula. ie |
3278 47
3280 36
3285 35
3286 37
3287 30
3288 15
3290 16
3291 2A
3293 30
32M 30
3205 15
3296 4
3297 26
3302 30
3303 B
3304 | Mouth of Bristol Bay_-.-.....-.-.-.-.---- 2
GR ss. 0. sk. ee Aaah ectetgn dete acento 23
3306 |..... NER isaac Dead dqaptrens gph we pin 3B
S438 | Vicinity of Pribilof Islands, 57° 06’ 30” N., 20
170° 22’ 30” W.
3439 |.....do... 41
3442 |..--.! ieee las thee deg inate aah 47
M82 Vicinity of Pribilof Islands, 57° 18’ N., 42
170° 42° W.
3504 Vicinity of Pribilof Islands, 56° 27’ N., Mu
169° 27’ W.
3805 | Vicinity of Pribilof Islands, 57° 09’ N., 38.1
168° 17” W.
3507 | Bering Sea, 57° 43’ N., 164° 42” W_....---.. 31
3508 | Bering Sea, 58° 33’ N., 164° 49° W....-.... 3
3510 Vicinity of Pribilof Islands, 57° 12 30” N., 77
169° 51’ W.
39
38
3522 | Vicinity of Pribilof Islands, 57° 5S’ N., ai
170° oo W.
3536 | Vicinity of Pribilof Islands, 57° 05 N., 40
| 170° 38” W.
3843 Vicinity of Pribilof Islands, 56° 41’ N., 43
me 169° 39° W.
Vicinity of Pribilof Islands, 56° 56’ N.,| 39
3 169° 52 W.
North coast Unimak Islands. .........-..- 9
_ Round Island, Coal Harbor, Shumagin | Low
| Islands. | tide.
Sanborn Harbor,
| lands
| New Harbor, Unga, Shumagin Islands...
z 3-6
ti
10-16
| Chickagof Harbor, Attu................-- 57

| Bering Sea, 61° 03’ N.., 167° 85’ W.....

! Probably incorrect; too high.

| Gray sand, mud...

7 INGER, BEARER 2K Fos noone fonn wees saendeneesceesnanannene
Beach. | cist jusapatin td Wrage eisai ils

Nature of bottom

Gray sand, rocks.

Gray sand, black specks.....
Fine gray sand, shells,
gravel.

Fine black sand...........--
Black mud, shells
Green mud, fine gray sand__

Fine gray sand, black specks.

Fine ar green mud...
Coarse gray sand, gravel Lo

Green mud, fine sand.......
Black’sand, shelis...........
Gray sand, broken shells... .

Fine volcanic sand

BO astanee cial - <0 0 ~ game atfups

Stony and made

Num
Bottom
temper- ber of
ature

eal 1

specl- Remarks
mens

Intermediate, close
skeleton.

Typical.

Intermediate.

Variable.

Intermediate, with
J. aretica.
4) Resembles /. arctica.
1 | No. 6085, W. Hi. Dall.
| 2 No. 6012, W. H. Dall.
|
| 1 | No, 6052, W. H. Dall.
4 No, 6045, W. H. Dall.
4) (12-777).
339 (1061), W. H.
Dall.
1 0075, W. H. Dall.
1 Geo. M. Stoney.
|

34 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Specimens of Leptasterias miilleri examined

1 Stanford University.

Riksmuseum, Stockholm.
Do.

3 Riksmuseum, Stockholm
(Sandeberg expedition).

West coast of Sweden..-------------------------------------
Finmark (Grotsund) ---------------------------------------- 6
Bohusliin (west coast Sweden) ------------------------------- 9
Waideguba (40 fathoms) ------------------------------------

Iceland (Berufjord, 15 to 30 fathoms) ------------------------- 1 Do.
Iceland (Reikiavik, 8 fathoms) ------------------------------- 1 Do.
Northwest of Bergen (90 to 200 fathoms) - -------------------- 5 Do.
Southwest of Nidingen (west coast Sweden) - ------------------ 1 Do.
Specimens of Leplasterias hyperborea examined
Spitzbergen (Ice Fjord) -.----------------------------------- 1 Riksmuseum, Stockholm.
North of Spitzbergen (80° 65’ N., 14° 40’ E., 16 fathoms) ------- 4 Do.
Spitzbergen (1 from Cross Bay; 2 fathoms) -------------------- 2 Do.
Baffin Bay (67° 59’ N., 56° 33’ W., 98 fathoms) - -------------- 3 Do.

Remarks.—There is no sharp line of demarkation between the two formae.
There is a considerable number of specimens which may be referred to either. In
Bering Sea north of about 60° the specimens are more like the Arctic race, although
larger as a rule. Even as far south as the Pribilof Islands (as station 3559) large
specimens resembling forma artica are found, but the crossed pedicellariae are larger
than in the typical form from the Arctic Ocean. (Pl. 8, fig. 5e.) Occasionally (as
stations 3290, 3600) well within the range of forma beringensis, specimens are found
which closely resemble forma arctica abactinally. If there is any geographic signifi-
cance to the large form from the southern part of Bering Sea, it is likely that these
southern “intermediates” are simply variants of beringensis (especially as they have
the larger crossed pedicellariae), and not of the typical phase, found in the Arctic
Ocean.

The variety from the Aleutian and Shumagin Islands is rather variable and not
typical but is more like forma beringensis than arctica. A series of drawings of pedi-
cellariae on Plates 8 and 10 will show a real but slight difference between the northern
and southern forms. They are introduced also for comparison with the crossed
pedicellariae of L. hyperborea and L. miilleri.

Through the interest of Dr. T. Odhner, of the Naturhistoriska Riksmuseum,
Stockholm, I have received specimens of a number of Arctic species for comparison.
Especially useful have been the series of L. hyperborea and L. miilleri.

The question of whether these are distinct species or formae of the same species
has been frequently raised. Won Hofsten (1915, p. 62) has followed Grieg (1907,
p. 13) in considering them separate. The matter is of interest in determining the
nearest relative of arctica in the North Atlantic and adjacent Arctic Ocean. Grieg
(1907) writes:

If we compare Asterias hyperborea with Asterias miilleri and nearly allied forms, we shall find
that it has quite a different and coarser armature on the abactinal face of the disk. The most
characteristic difference, however, is in the pedicellariae. In Asterias miilleri there are a few
forficiform [straight] pedicellariae on the ambulacral papillae, or they are altogether wanting; in

Asterias hyperborea, on the other hand, the pedicellariae are forcipiform [crossed] and arranged in
groups.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——FISHER 35

The distinction is unfortunately not so simple. Crossed pedicellariae are present
on the adambulacral spines of all the specimens of milleri examined by me, and some-
times in fair numbers. They are also abundant on the same spines of hyperborea.
The abactinal spines do average larger on hyperborea and are often more numerous
than in milleri. There is, however, a small but distinguishable difference in the
crossed pedicellariae. (PI. 8, figs. 4, 4a, b; pl. 11, figs. 1, la-lc.) In hyperborea the
pedicellaria is wider (pl. 11, figs. 1, 1c) as seen in profile, with a larger terminal lip
which is broader than in miilleri. The large specimen of hyperborea from Ice Fjord,
Spitzbergen, has numerous conspicuous spatulate straight pedicellariae similar to
those of arctica. (PI. 11, figs. 1d, le.) This example has R 82 mm., and is much
bigger than average specimens of miilleri.

Dr. A. Djakonov, of the Zoological Museum, Academy of Sciences, Leningrad,
made comparisons of specimens of L. miilleri, L. hyperborea, and L. groenlandica at
my request and wrote me as follows:

My investigations have led me to the conclusion that: 1. Kalischewsky had an incorrect idea
of these species and what he held for L. hyperborea was really L. groenlandica in all cases. 2. L.
groenlandica is a good species and is clearly different from miilleri-hyperborea. 3. L. hyperborea can
only with great difficulty be distinguished from miilleri and is, as it seems, only its arctic form, there
being intermediate specimens between them. 4. ZL. miuilleri is not to be met with eastward of
middle Murman coast of northern Russia; records of muiilleri from the coast of northern Siberia
are referable to hyperborea.

I think the relationship of arctica is with miilleri rather than with hyperborea.
The crossed pedicellariae of arctica are more like those of miilleri than of hyperborea,
and the general appearance of arctica, especially of forma beringensis, is much like
that of miilleri. It is likely that the two species intergrade, probably through a
series of slightly differentiated geographic forms.

Verrill (1909, p. 553) has expressed the opinion that hyperborea and arctica are
the same, but in 1914 he recognized arctica as a full species without mentioning his
previous identification, nor pointing out the close similarity of the two.

Arctica seems to be the equivalent of milleri in Bering Sea and the Arctic Ocean
to the north. It differs from miileri in having either a closer, more capitate abactinal
spinulation or numerous spatulate straight pedicellariae or both in varying degrees.
The south Bering Sea form of arctica is larger than miilleri with heavier spines and
more numerous pedicellariae of both sorts.

LEPTASTERIAS HYLODES, new species
Plate 10, Figures 5-11; Plate 17; Plate 1S, Figures 1-3

Diagnosis.—Differing from L. arctica in having uniform, rather close-set, slender,
terete, often slightly tapered, striated but noncapitate, heavily sheathed abactinal
spinelets, the carinal series of which is not at all differentiated from the other spines.
The species occurs in a dwarfed form, sexually mature at R 15 to 17 mm., in which
the skeleton and spinelets are relatively more delicate than in the large specimens,
and more delicate than in corresponding sizes of L. arctica. Type, R. 39 mm., r 9
mm., R=4.3 r; breadth of ray, just beyond base 12 to 14 mm. Rays well arched
with rounded sides and plane actinal surface, slender to medium stout, gently taper-
ing from a slightly swollen base to a blunt extremity.

36 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Description.—The surface of the body has the appearance of being covered with
close-set. slender, subterete, uniform, small spinelets emerging, in alcoholic speci-
mens, from a pulpy sheath, carrying in the small specimens usually few crossed pedi-
cellariae. The spines are pretty uniformly spaced, but vary considerably in number.
In smaller examples (as with R 25 to 30 mm.) from station 2847 the spinelets appear
as a coarse nap, with very narrow interspaces between the sheaths of neighboring
spinelets; in others there are much wider interspaces with an evident arrangement of
spinelets in longiseries, although sometimes with no great regularity. The abactinal
spinelets are typically very much slenderer than in arctica, terete, slightly tapering,
terminally striated in large examples and bluntly pointed. In typical, small, deep-
water examples (as stations 2845, 2846, 2847, 2851, 2856) this character is accentu-
ated, the spinelets being very delicate and sometimes almost setalike—quite different
from the stout capitate spinelets of arctica. But in some localities the spinelets are
heavier, slightly clavate, with a rounded striated summit (as stations 4281, 3260,
3261, 3265). In hylodes there is no crowding of the carinal spines, nor are they dis-
tinguishable by greater size from the others, except in a few aberrant specimens,
where the discrepancy is slight. One of the most characteristic features of this
species is the uniform appearance of the abactinal area—entirely unlike that of
typical arctica. This is as well marked in the relatively gigantic type (and in an
equal sized specimen from station 2847) as it is in the general run of specimens which
are small, though sexually mature (as station 2856). In the type the dorsal spines
are 0.90 to 1.2 mm. long.

The supero and infero marginal spines form two usually very regular lateral
series, and in all but the smallest specimens there is an incomplete series of actinals,
the number varying from a few spines to a nearly complete series. There are thus
three ventrolateral rows of spines, which in typical specimens are similar to the
dorsals, though usually larger, and in the case of the inferomarginals and actinals
slightly heavier in small examples and markedly so in large ones. There is great
variability in the thickness of these marginal and actinal spines, just as there is in
the dorsal. A short series of smailer intermarginal spinelets is generally present.
The superomarginals have a complete wreath of pedicellariae, the inferomarginals
a complete or an incomplete wreath and the actinals a tuft on the outer side, the
number of pedicellariae variable but not high. The marginal and actinal spines in
the largest examples are less regular than in the small.

Adambulacral spines slender, subterete, slightly tapered, bluntly pointed, one
or two to a plate in fairly regular alternation, except on the first dozen plates which
may carry mostly one, or mostly two, spines. The adoral adambulacral plates
usually three in contact behind the oral plates (and the first rather the longest),
each carry but one spine. The large type (station 2851) has two spines on most of
the plates on the proximal half of ray.

Oral plates small, the spines two or three as in arctica. Where there are two,
which seems to be more often, the furrow or smaller marginal is suppressed. In the
type each plate has a short tapering blunt spine at the inner end and another, the

suboral, at the outer. The latter may carry a large unguiculate pedicellaria, or only
one or two crossed pedicellariae.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 37

Papulae usually not conspicuous, one or two to the abactinal areas (frequently
only one in small examples); two, three, or less often four intermarginally (one or
two in small examples); one to three in actinal areas. The skeleton is somewhat
more compact, less open, than in arctica and the papular areas are consequently
smaller.

Straight pedicellariae. (Pl. 10, fig. 9.) A few relatively large unguiculate
pedicellariae are present on the abactinal surface, especially near the interradii,
in the intermarginal and actinal channels, and in the actinal interradial channel.
Those of the abactinal surface may be nearly or quite as long as the slender spines
and considerably stouter. Similar but smaller pedicellariae occur on some of the
suboral and proximal adambulacral spines but not at all constantly. Smaller pedi-
cellariae without teeth but with a notch in the end of one jaw, and the tips of the
jaws crossed, are abundant, as in arctica, along the furrow face of the adambulacral
plates.

Crossed pedicellarie very variable in number but in small specimens generally
few, form a circle around the summit of the prominent sheath of the abactinal and
marginal spines, and tufts on the outer side of the actinal and adambulacral spines.
In most of the typical deep-water specimens the pedicellariw are few. In specimens
from station 2856, the dwarfed deep-water form, there are only two to five pedicel-
larie surrounding the dorsal spines, four to eight around the marginals, and one to
three on the adambulacral spines. Typical pedicellariw from station 2856 are fig-
ured (pl. 10, figs. 6 and 7). The abactinal are about 0.21 to 0.225 mm. long (in the
large type 0.20 to 0.25 mm.). The adambulacral may be slightly longer. Some of
the furrow spine pedicellarie differ from the abactinal, as in artica, in having a longer
terminal denticulate portion when seen in profile. These differentiated pedicellarie
seem to increase in numbers rather more rapidly than the others, as the animal
grows, and they appear to be less numerous in this species than in artica but it is
difficult to achieve any degree of accuracy since variation in the total number of
pedicellarie is so wide. (Pl. 10, fig. 7.) The crossed pedicellarie are more numer-
ous than usual in the large specimen from station 2847 (pl. 10, figs. 5, 5a), and the
sheaths carrying them are especially prominent, forming near the tip of spine a good-
sized flange of pulpy tissue carrying 6 to 12, or even more pedicellari#, which measure
about 0.27 mm. in length. The inferomarginals carry upward of 25. In contrast to
this, the type specimen from station 2851 (pl. 10, fig. 8) has slenderer abactinal
spines with only two to five pedicellarie each (0.20 to 0.25 mm. long), and upward
of 20 on the marginal spines.

Madreporie body small, variable, situated about 0.6 r from center and with a
circle or half cirele of spinelets guarding it.

Variations —The general appearance of this species varies less than that of
arctica; but there is nevertheless considerable difference in specimens from the same
station. Some of the differences, such as the spacing and robustness of spines, have
been stated in the description. It may be added that there is diversity in the robust-
ness of the rays as well as in the proportions of R to r.

The specimens from the greatest depth, 68 fathoms, station 2856, are all small,
although most of them are sexually mature, having well developed gonads. The
largest has R 24 mm.; the largest from station 2846, 44 fathoms, has R 35 mm.; that

63160—30——4

a

3 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

from station 2847, 48 fathoms, has R 39 mm.; from station 2845, 42 fathoms, 30 mm. ;
from station 2851, 35 fathoms, 39 mm.; from station 3259, 25.5 fathoms, 33 mm.;
from station 3265, 38 fathoms, 37 mm. Thus the maximum size ranges roughly
from 24 mm. to 39 mm., the smallest being from 68 fathoms. The specimens from
stations 3213, 3214, 3215, 3217, 3218 are also small, and are all within these extremes—
the majority having R less than 30 mm.

The specimens from Popoff Strait, the Shumagin Islands, and Agattu are not
typical. They are small and the spinelets are more widely spaced than in average
deep-water examples. Crossed pedicellariz are few on all spines.

Specimens from stations 3265, 3266, and 3291 have given considerable trouble
on account of their variability and general resemblance to ZL. arctica. Those from
3265 are referable to hylodes except one which if considered alone would be regarded
as L. arctica forma beringensis with rather slender spines, and is so recorded. Speci-
mens from 3266 are like this single example from station 3265 except one which is
referable to hylodes. Four have been recorded as aberrant arctica. Out of 28 speci-
mens from station 3291, one is referable to hylodes and the others to L. arctica forma
beringensis with unusually slender, but fairly well spaced spines. Other specimens
of arctica which approach hylodes are found at stations 3242 and 3262. Typical
hylodes is very distinct from arctica and ranges, through various small races, into
Japanese waters. It is found in its most typical form south of the Alaskan Penin-
sula, west of Afognak Island. Whether these aberrant specimens of arctica from the
region of Bristol Bay are intermediates, or the results of crossing with hylodes, speci-
mens of which were found in the same bottles (as stations 3265, 3266, 3291) must
remain undecided. It is probable that hylodes does not range far into southern
Bering Sea.

Anatomical notes—The skeleton is essentially like that of arctica, with sur-
prisingly little difference when allowance is made for the disparity in size. In
hylodes the abactinal skeleton consists almost entirely of primary, mostly 3-lobed
plates with relatively few intermediate ossicles such as multiply in large examples
of true arctica, or its southern Bering Sea form. Corresponding to the first 10
inferomarginals there are 29 or 30 adambulacrals. The series of actinal plates of
a specimen from station 2847 with R 22 mm. extends along the ray about two-
thirds R from center, or to the twelfth inferomarginal (a little over half length of
ray measured on side).

The gonads are fully developed in an example with R 17 mm. from station 2856.
They open on the ventral surface, close together, on either side of the interradial
furrow, and on a level with the actinal plates, or a trifle higher. The aperture is at
the summit of a conspicuous thick papilla, when the products are mature, and is
often guarded by a large straight pedicellaria. The testes extend fully half the
length of ray and consist of a number of relatively slender, lobulated lobes, while
the ovaries are shorter, more compact, with a few subspherical lobes, or without
lobes. In a specimen from station 3215, May 21, temperature 38.5° F., the gonads
are not mature, although they appear to be on the way to maturity. Those from
station 3265, June 25, temperature 39.8° F., have good-sized gonads, apparently
not yet mature, however. In examples from station 2847, July 31, temperature

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER

39

42° F., the eggs appear to be fully developed and relatively few and large; in speci-

mens from 2856, August 22, temperature 44° F., the gonads are also mature.

of the specimens has young.
Type.—Cat. No. E 1490, U.S.N.M.

Type locality.—Station 2851, vicinity of Shumagin Islands, 54° 55’ N., 159° 52’
W., 35 fathoms, gray sand, broken shells, bottom temperature 44.8° F,
Distribution —In a typical form, south of the Alaska Peninsula and Aleutian
Islands, 25 to 68 fathoms, but ranging into shallower water.

Specimens examined.—Two hundred and sixteen.

None

Specimens of Leplasterias hylodes examined

Locality

South of Unimak Island, 54° 05’ N,, 164°
oy W.

South of Unimak Island, 54° 08’ N., 162°
ww.

Vicinity of Shumagin Islands, 55° 01’ N.,
160° 12 W.

Vicinity of Shumagin Islands, 54° 52’ N,,
159° 40° W.

Vicinity of Shumagin Islands, 4° 55’ N.,
159° 52° W.

Off Afognak Island, 58° 07’ N., 151° 36’ W_

South of Unimak Island, 54° 10’ N., 162°
57’ 30” W.

South of Unimak Island, 54° 13’ N., 163°
ov W.

South of Unimak Island, 54° 14’ 40” N.,
163° 24’ W.

South of Unimak Island, 54° 14’ 50” N.,
164° 06" W.

South of Unimak Island, 54° 15’ 50” N.,
164° 21’ W.

South of Unimak Island, 54° 14’ N., 166°
as’ W.

South of Unimak Island, 54° 15’ N., 165°
ov Ww.

ERRNO I Netter toraelet Ais detgtps ecto aba cd aia

North coast of Unimak Island...

Popoff Strait, Alaska......................
Round Island, Coal Harbor, Shumagins_.
Captains Harbor, Unalaska

ARN eee ens ina eh ie de wp

+} Lowest water.

|
| Bottom} a
Depth Nature of bottom |temper-) ert Remarks
| Sture | mens
| | i ee —
Fathoms | F.
42 Coarse black sand_.......... 42 8} U.S.N.M
RR, OURO ane en eines 42 | 7 Do.
48 Fine gray sand.............- 42 17 Do.
21 | Broken shells....--.......-- 48.2 2 Do.
Gray sand, broken shells..... 44.8 1 Do.
|
cy Gray shells, black specks... 44 47 | Do.
AD Ak SENN esse 12} Do.
{
38 | Gray sand, gravel..........- soneese 2 | Do,
43 revels dis so hess S88 38.5 10 Do.
42° | Black gravel; _.......--..<.- EELS | nh Do.
} }
41 Black sand............:...-- 37.7 15 | Do.
OO ie. 10 chin ce kt 3 3 Do.
uM Gravel, broken shells........ eles 1 | Do.
10.5 | Pebbles, stones..............'..-..... 2 | Not typical.
5 | Fine gray sand_._. 37.5 26 | Do.
Fine black sand_....... 42 | 2)| U.S.N-Mi
41.2 ll
39.8 25
a 1 Do.
| Fine gray sand, shelis, 41.8 1 U.S.N.M. Small
gravel. | form.
Greet womd o5 Se. id 7 1 | Small form.
Green mud, sand, rocks..... Sista Pe 1) Do.
| Gray sand, broken shells..... 47.2 1} Do.
Re ey eat Te eee 1) W. H. Dall, 4191.
a pleased a nnccirglt mare anew 2) W. H. Dall, 0085.
POR PARURMO oh Sn dinsine mated ambensintl 1) W. A. Dall, 6075.
Knasnann deed daha heetie Ler Bats 2) Albatross, 1906.
Sean geu etn ainnatie skeen anaes <mteporae 3 | Albatross, L888.
3 Beach neap tide. ) Low tide.

40 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Remarks.—In the foregoing description all the comparisons have been with ZL.
arctica, but hylodes resembles, as closely, the arctic L. hyperborea. This species is
very rare, poorly understood, and usually confused with L. miilleri. L. hylodes re-
sembles hyperborea in having very numerous, close-set abactinal spines among which
the carinal series is not readily distinguishable. It differs in the form of the straight
pedicellariae (which are rarely present in hyperborea), and in the form of the crossed
pedicellariae. (Compare pl. 11, figs. 1, 1d, with pl. 10, figs. 5, 6, 9.) When compared
side by side with hyperborea, the rays of hylodes are seen to be thicker and shorter and
the abactinal spinelets much more numerous and shorter.

LEPTASTERIAS ORIENTALIS Djakonov

Plate 11, Figures 2, 2a-2h; Plate 19, Figures 2, 3; Plate 20, Figure 3

Leplasterias orientalis DsaxoNov, Neue Seesterne aus dem Ochotskischen Meer. II. Lep-
tasterias orientalis sp. n., Compt. Rend. Acad. Sci. U. R. S. S., 1929, p. 277.

Diagnosis.—Rays five, of medium stoutness, tapered, slightly swollen at base;
disk small; spinelets slender, tapered, with heavy wreaths of pedicellariae, rather uni-
formly and closely placed, but without longiserial order except on sides and actinal
surface. Differing from ZL. hylodes in having much more numerous crossed pedicel-
lariae, especially abactinally; less numerous abactinal spinelets; and in having more
numerous shank teeth on the jaws of the crossed pedicellariae. Differing from
L. hyperborea in the form of both crossed and straight pedicellariae and in the greater
abundance of the crossed pedicellariae. R 55 mm., r 9 mm. R=6 r; breadth of ray
at base, 13 mm.

Description.—Spinelets slender, tapering, pointed, less numerous, longer, and
with conspicuously larger circumspinal clusters of pedicellariae than in hylodes.
Across the abactinal area, at base of ray, 13 to 15 spinelets can be counted. The
wreaths touch one another and neither the carinals nor dorsolateral spines (except
adjacent to superomarginals) form longiseries. Papulae singly or in groups of two or
three emerge between the spinelets.

The marginal and actinal spinelets are longer than the dorsolateral and are clad
in heavy stoles of pedicellariae, which on the actinals are on the outer side of the spine.
One series of actinals extends nearly to end of ray while a second occupies the basal
third. Only one series is present in the specimen from station 5024.

First six or seven adambulacrals monacanthid; then irregularly monacanthid
and diplacanthid; then monacanthid on outer half of ray (mostly monacanthid
throughout on small example). The furrow spines of alternate plates are set a little
further furrowward, giving the appearance of two series of spinelets along the margin,
or even three on the basal half of the ray. Spines with heavy cluster of crossed pedi-
cellariae.

The apertures of the gonoducts are marked by two prominent papillae in each
actinal interradial area (near the sixth or seventh adambulacral plate), usually
guarded by a large straight pedicellaria. :

Mouth plates with one slightly curved, tapered, blunt actinostomial spinelet
about as long as median suture and carrying one or two lanceolate straight pedicel-
lariae; and one suboral, longer straight spinelet similar to the adambulacral spines.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 41

This usually carries, in addition to several crossed pedicellariae, one or two promi-
nent, narrow, straight pedicellariae with the end of the jaw notched or pointed.

Crossed pedicellariae (pl. 11, figs. 2c, 2d) measure 0.27 to 0.31 mm. long and have
rather numerous teeth especially in the outer of the vertical rows of shank teeth.
The specimen from station 4810 (pl. 11, fig. 2a) has smaller pedicellariae (length about
0.2 mm.) and is intermediate in this respect with that from off the Bay of Sendai
(station 5047) which represents another, closely related, race. In this form the pedi-
cellariae are still smaller (about 0.18 or 0.19 mm.) with numerous very small shank
teeth (pl. 11, fig. 2b), but are not nearly so numerous as in orientalis.

Straight pedicellariae 0.7 to 1 mm. long, sparingly present in intermarginal
channel and between proximal actinal spines; two are usually present, near the
apertures of gonoducts, in each actinal interradial area. The jaws may be narrowly
spatulate with a few teeth or tapered and without teeth. (PI. 11, fig. 2e.) Small
narrowly lanceolate ones are present abundantly along the furrow face of the adam-
bulacral plates; a few larger ones on the oral and proximal adambulacral spines.

Type—Cat. No. 5164, Museum of the Academy of Sciences, U. R. S. S.,
Leningrad.

Type locality —Okhotsk Sea, 57° 36’ N., 140° 34’ E., 106 meters, mud, bottom
temperature —1.4° C.

Distribution —Okhotsk Sea to Sea of Japan, cold area.

Specimens eramined.—Okhotsk Sea, 57° 36’ N., 140° 34’ E., 106 meters, mud (sta-
tion 9), hydrographic expedition, Pacific Ocean, August 14, 1918, from the Academy
of Sciences, U. R. S. S., Dr. A. Djakonov, one specimen (topotype).

Station 5024, off Cape Patience, Saghalien, 48° 42’ 10’” N., 144° 59’ 30’’ E., 67
fathoms, sand, pebbles, bottom temperature 30.9 F., one specimen (Albatross, 1906).

In addition to the above, two specimens with R 21 mm., from station 4810, off
Cape Sirakami (Hokkaido), west entrance to Tsugaru Strait, Japan, 195 fathoms, fine
gray sand, bottom temperature 44.7° F. (not typical).

Remarks.—The occurrence of this form in truly arctic temperatures of the Sea of
Okhotsk suggests that it may be a derivative of the high arctic hyperborea. It appears
to be more closely related to hyperborea than is hylodes; both resemble hyperborea,
however.

Orientalis has much heavier stoles of pedicellariae than hyperborea. For the form
of the crossed pedicellariae of hyperborea see Plate 11, Figures 1, la, 1b, lc. These
are relatively broader. The straight pedicellariae of hyperborea have wider, spatulate
jaws with more numerous teeth. (PI. 11, figs. 1d, le.)

The two specimens from the Sea of Japan are not at all typical. The crossed
pedicellariae are smaller and the straight pedicellariae are much slenderer, with more
delicate teeth (resembling the slenderest type of hylodes). They are probably ex-
amples of a small race, living in the cold water of the Japan Sea, intermediate be-
tween orientalis and an undescribed race found in the warmer waters off the east
coast of Hondo (station 5047). Station 4810 is at the southern limit of winter ice in
the Sea of Japan.

42 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

LEPTASTERIAS FISHERI Djakonoy
Plate 11, Figures 3, 3a—3g; Plate 20, Figures 1, 2; text Figure 1

Leplasterias fisheri DsaKonov, Neue Seesterne aus dem Ochotskischen Meer. I. Leptasterias
fisheri sp. n., Compt. Rend. Acad. Sci. U. R.S.5., 1929, p. 233.

Diagnosis.—A relatively large, 5-rayed species having all the spines arranged in
spaced longiseries. Rays slender, very gradually tapered, well arched; disk small;
R 85 mm.,r 15 mm.; R=5.66 r; breadth of ray at base, 15 mm. Carinal spines
blunt, one to several to a plate, in a prominent, irregular longiseries; on either side
two fairly regular well spaced dorsolateral series of smaller spinelets; then two
marginal series, successively larger and very regular, one spine to a plate; then two
actinal series, also regular and slightly larger; at base of ray a few small intermar-
ginal spinelets; all with neat stoles of normal-sized crossed pedicellariae; adambu-
lacrals two and one, alternating; large intermarginal and actinal, toothed, straight
pedicellariae. Differing from L. arctica, L. groenlandica, and allies in the longiserial
arrangement of the well spaced dorsolateral spines and in the conspicuously longer
rays and heavier bivalved pedicellariae.

Description —Carinal spines short (0.9 to 1.1 mm.), robust, round-tipped, one
to three to a plate, forming a conspicuous, irregular series in marked contrast to the
dorsolateral spinelets which are slenderer, shorter, and stand in two fairly regular
well-spaced series. Outside of these there is a very regular series of similar, but
larger, bluntly pointed superomarginals (1.5 to 1.8 mm.), one to a plate; then outside
of these and separated therefrom by about their own length is a very regular series of
pointed inferomarginals, one to a plate, about 0.2 to 0.23 mm. long. The supero-
marginals are rather high on the side of ray, whereas the inferomarginals define
ambitus as viewed from above, and may be easily mistaken for the superomarginals,
inasmuch as directly below them there is a series of exactly similar, but a trifle
longer, actinals which extend very nearly to the tip of ray. A second series of
similar actinals extends about half the length of ray. There are thus on either side
of the carinals, two series of small dorsolaterals, one series of larger superomraginals
and then three series of still larger subequal spines, the upper of which is inferomar-
ginal. These spines are surrounded at base by a neat wreath of crossed pedicellariae
(a tuft only on the inner actinals) and in the actinal and intermarginal channels are
fairly numerous bivalved pedicellariae about 1 to 1.2 mm. long. (Pl. 11, fig. 3a, b.)
On the disk the spinelets are unequal, irregularly spaced like the carinals.

Between these regular longiseries are longiseries of small groups of papulae, two
to four dorsally, one or two laterally and ventrally.

The carinal, marginal, and actinal plates are fairly definitely four lobed and are
directly imbricated in series, without secondary ossicles except sporadically, as shown
in text Figure 1. The exact form and relations of the dorsolateral plates are indicated
in the figure (dotted plates).

Generally speaking, the adambulacral spines, which are slender, tapered, pointed,
and heavily laden with crossed and straight pedicellariae (pl. 11, fig. 3¢) alternate
two and one along the furrow, except at the mouth angle, where the first few plates
carry one only. Adoral carina narrow, consisting of three pairs of contiguous plates,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 43

the fourth pair slightly separated. The first two plates are distinctly longer than the
third, and the two combined are about as long as the mouth plates.

At the inner end of the mouth angle are two blunt actinostomial spines, and on
the narrowed actinal surface the two suboral spines are crowded one behind the other.
Actinostome small, sunken, having the general appearance of Evasterias.

Crossed pedicellariae (pl. 11, fig. 3), rather numerous confined to spinal sheaths,
as noted above. The abactinal are 0.32 to 0.34 mm. long.

Straight pedicellariae of two sorts: (1) Rather generalized, lanceolate ones
(either robust or medium slender) of various sizes, on the adambulacral and oral
spines (somewhere around 0.7 mm. long, pl. 11, fig. 3c), and numerous smaller pedun-

Figure 1—LerrasteriasS FIsneR! DIAKONOV. PLAN OF THE ABACTINAL SKELETON BETWEEN
THE CARINAL AND INFEROMARGINAL PLATES, BASE OF RAY TO LEFT, TH& DORSOLATERAL PLATES
ARE DOTTED; X 8. CAR, CARINALS; J, INFEROMARGINALS; S, SUPEROMARGINALS; [-3, THE THREE
LONGISERIES OF DORSOLATERAL SKELETAL INTERVALS

culate ones at base of same spines; (2) larger toothed ones (pl. 11, figs. 3a, 36) in
intermarginal and actinal channels and very rarely on abactinal surface (1 to 1.2
mm. long). Two or three of these stand near the genital apertures on the very
narrow actinal interradial region.

Madreporic plate 2.5 mm. in diameter, situated at middle of r, at top of axil-
lary channel.

Gonads attached to body wall at base of ray between the superomarginal and
inferomarginal plates. In one interradius the two genital papillae can be demon-
strated just back of the adoral carina.

Type—Cat. No. 5139, Museum of the Academy of Sciences, U. R.S. S.,
Leningrad.

44 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

—

Type locality — Okhotsk Sea, 50° 03’ N., 144° 08’ E., 110 meters, mud.

Distribution. —Okhotsk Sea, Gulf of Tartary.

Specimen examined.—Okhotsk Sea, 50° 03’ N., 144° 08’ E., 110 meters, mud.

Remarks.—I am indebted to Dr. A. Djakonoy for the specimen upon which the
illustrations and the above account are based.

This species, like orientalis, lives in the very cold environment of the Okhotsk
Sea. It appears to belong in the hyperborea-groenlandica section of Leptasterias, from
which, as well as from all other 5-rayed north Pacific Leptasterias, the species can be
distinguished by the regular dorsolateral longiseries of spinelets. In a general way
fisheri resembles L. tenera forma compta of New England waters. In L. tenera, which
is a small species, the spaced spinules are heavily wreathed but the carinals are
similar to other abactinals, and no large bivalved pedicellariae are present in my
specimens. The crossed pedicellariae resemble those of fisheri in details, but are
only 0.2 to 0.22 mm. in length abactinally.

L. leptalea Verrill was based upon a very small specimen from southern Alaska.
Since the type is lost and no other specimen is known, it is doubtful if the species
can be surely identified, as no details are figured. The crossed pedicellariae are said
to be of “unusually large size, with strongly curved blades, numerous in groups
around the dorsal spines and on the papular areas.” This does not apply to fisheri.

LEPTASTERIAS LEPTALEA Verrill
Leptasterias leptalea VerRiLL, Shallow-water Starfishes, 1914, p. 119, pl. 18, fig. 3.

The 5-rayed type of this species, the ventral side of which is figured by Verrill
(pl. 18, fig. 3), is said to be from Sitka. The only locality mentioned in the
description is Virgin Bay, Prince William Sound, Alaska. The type can not now be
located at the Peabody Museum, Yale University. A small 6-rayed specimen from
Sitka (No. A. 275) collected by W. R. Coe, 1899, and labeled leptalea is obviously
not the type, nor this species, there being six rays, two or three superomarginal
spines, and small crossed pedicellariae. It is possible that the /abel may have been
attached to the wrong specimen, which resembles L. asteira.

Verrill’s small figure of the ventral surface lacks pertinent detail. Such figures
of Leptasterias without a dorsal view are of little value. It resembles L. groenlandica,
some specimens of arctica, and some of hylodes, and even hyperborea and orientalis.
From the standpoint of locality hylodes is the most likely, if the description fitted.
The dorsal spinelets of hylodes can hardly be described as “‘not numerous, conical,
relatively stout;’’ nor are the crossed pedicellariae ‘“‘of unusually large size with
strongly curved blades;’’ nor do they ever occur on the papular areas. The straight
pedicellariae are not ‘‘small, ovate.”

The following is the original description:

A small and very delicate 5-rayed species. Radii, 2.5 mm. and 15 mm.; ratio, 1:6. Rays
terete, evenly tapered. Dorsal ossicles relatively strong, thickened, especially those in the median
rows. Dorsal spines not numerous, conical, relatively stout, usually standing singly, one to an
ossicle, and forming three or five rather irregular rows. Ocular plate relatively large, bearing a
dense cluster of small spines. Minor pedicellariae of unusually large size and with strongly curved
blades are usually numerous in groups around the dorsal spines and on the papular areas. Adam-
bulacral spines mostly two to a plate, or alternately one and two distally, very slender, terete, often
slightly clavate. Lower marginal spines, which are much larger, acute, conical, like tle dorsals,

ASTERROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 45

but longer, form a single row next the adambulacrals, or there may be two to a plate, proximally, in
the larger specimens. The synactinal plates appear to be few and small, or altogether lacking in
our examples, but may occur in larger ones. Upper marginal spines form a single row and resemble
the dorsals in form and size. Major pedicellariae few, small, ovate.

The ambulacral feet are in four rows, but the adjacent rows form only a zigzag line, so that they

sometimes appear almost biserial.
LEPTASTERIAS GROENLANDICA (Litken)
Plate 8, Figures 1, la-le, 2, 2a, 2b, 3, 3a; Plates 21-23; Plate 24, Figures 1, 2

Asteracanthion groenlandicus LUTKEN, Vidensk. Meddel. N. Forening i Kjébenhavn, 1857,
p. 29.—STEENSTRUP, Forh. skand. Naturf., 1857, p. 228.—Duncan and SLADEN,
Echinodermata of the Arctic Sea, etc., 1881, pp. 27-29, pl. 2, figs. 9-12.

Asterias cribraria Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 270.—PeRRIER,
Rév. Stell., 1875, p. 65.—Lupwia, Zool. Jahrb., 1886, p. 288; Fauna Arctica, vol. 1,
p. 482.—BeE Lt, Proc. Zool. Soc., 1881, pp. 494, 505.

Asterias groenlandica Stimpson, Proc. Acad. Nat. Seci., Phila., 1864, p. 142.—VERRILL,
Amer. Nat., vol. 43, 1909, p. 553.—LigBERKIND, Vidensk. Medd. fra Dansk naturh.
Foren., vol. 72, 1920, p. 121, text figs. 1-4.

Asterias groenlandica var longimana Kauiscuewsky, Mem. Acad. Sci., St. Petersburg,
ser. 8, vol. 18, no. 4, 1907, p. 41.

Asterias hyperborea KaLiscHEWSKY, 1907; according to A. Djakonov, letter, 1929.

Asterias inermis Bevu. Proc. Zool. Soc., 1881, p. 512, pl. figs. 2, 2a. (Fisher, Ann. and
Mag. Nat. Hist., ser. 9, vol. 18, 1926, p. 198.)

Asterias spilzbergensis DANIELSSEN and Koren, Norwegian North Atlantic Exp., Asteroidea,
1884, p. 5-7, pl. 1, figs. 1-11.

Leptasterias groenlandica Vernritt, Proc. U. S. Nat. Mus., vol. 2, 1879, p. 151.

Asterias milleri groenlandica voN Horsten, Die Echinodermen des Eisfjords, Kungl. Svenska
Vet. Akad. Handl., vol. 54, no. 2, 1915, p. 60 (distribution).

Ctenasterias cribraria VERRILL, Shallow-water Starfishes, 1914, p. 148, pl. 25, figs. 3, 4, text
fig. 7.—A. H. Crank, Rep. Canadian Arctic Exp., 1913-1918, vol. 8, 1920, p. Se.

Leplasterias obtecta VERRILL, Shallow-water Starfishes, 1914, p. 144. (Fisher, Ann. Mag.
Nat. Hist., ser. 9, vol. 12, 1923, p. 599.)

Diagnosis.—Rays five, slender to robust, tapering, bluntly pointed. Recogniz-
able usually by the broad but short dorsolateral papular areas which sometimes
reach from the carinal to the superomarginal plates, these separated by irregular,
sometimes anastomosing transverse trabeculae bearing small spinelets, either spaced
singly or in spaced small groups. Superomarginal spines usually two or three;
inferomarginals usually two; actinals, one, all surrounded by complete, dense wreaths
of pedicellariae. Adambulacral plates diplacanthid proximally. Dimensions of a
large specimen: R 55 mm.,, r 1] mm, R=5 r; breadth of ray at base, 14 mm.; of
a small breeding specimen (Bering Strait) R 17 mm.,r4mm., R=4.2 r; breadth of
ray at base, 6mm. A very variable species.

Description —In order to render this variable species recognizable by descrip-
tion, it will be necessary to consider first the more prevalent forma, which has a
characteristic appearance. This is the form made known by Stimpson under the
name cribraria and later by Danielssen and Koren as spitzbergensis.

The rays vary from slender to rather stout and in small specimens may be
inflated and tumid. Plates of abactinal skeleton slender and arranged in numerous
transverse bars, fairly close together, with intervening transversely elongate papular
areas—especially adjacent to marginal plates and on outer part of ray. But it is

46 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

not uncommon to find these narrow areas all along the side of the abactinal area,
and some of them extend without interruption from the carinal to the superomarginal
plates. They are accentuated by drying. The abactinal spinelets are small (0.4
to 0.7 mm. long) of variable caliber, but generally slender, distinctly clavate to
subeylindrical on the one hand or subeapitate on the other. They are rather num-
erous and on account of the structure of the skeleton form many transverse lines.
They stand on the trabeculae in lines either singly or in linear groups of two or three,
and increase in thickness toward the carinals. The latter are generally the largest,
subeapitate or capitate, striate, and form a definable irregular longiseries. In most
specimens there is a swpramarginal channel, immediately above which the spinelets
form a fairly well defined longitudinal dorsolateral series which is more prominent
in forma groenlandica. . Even in forma eribraria there is considerable variation in
the caliber, shape, and number of the abactinal spinelets. In alcoholic specimens
the spinelets appear to stand in transversely oriented groups, surrounded by deep
narrow creases in the pulpy integument.

Superomarginal spines in a transverse series of two, three, or four, the median
(or second from the top) generally the largest. They vary from slender to stout,
and from terete, slightly tapering to cylindrical or clavate, depending upon the
robustness of the general armature. On the same specimen the spines increase in
thickness and become much shorter toward the end of the ray, and they are subequal
to or slightly larger than the abactinal spines. The series of combs is generally
quite regular. Below them is a well-defined, and in some specimens rather broad,
intermarginal channel (showing large papular areas when dried). There is great
variation due in part to the shrinkage of the specimen in alcohol and in part to the
size of the gonads. Below this are two series of inferomarginal spines and one of
actinal. The latter extends a third or half the length of ray. These three series,
or two distally, have spines similar in form to the superomarginals but are a little
stouter and longer. In some examples the first half dozen inferomarginal plates are
monacanthid. The terminal plate (quaintly termed the “eyelid” by Stimpson) is
closely covered with stout clavate or subcapitate spinelets stouter than any others
immediately adjacent.

Adambulacral plates typically diplacanthid, except distally where they are alter-
nately diplacanthid and monacanthid, or are mostly monacanthid. The spines are
slender terete, with bluntly rounded tips. The inner or marginal is likely to be
gently tapered, and slenderer than the outer, which is more often slightly clavate.
The robustness of both varies with that of the general spinulation and is subject to
considerable variation. There are two plates in contact on the interradial line ex-
ternal to the oral plates. The first if conspicuously larger than the second. About
23 or 24 plates correspond to the first 10 inferomarginals.

The oral plates are typical of Leptasterias, with one apical and one suboral spine
to each plate, the former about as long as the median suture, but neither especially
diagnostic. The apical spine is usually tapered. Verrill (1914, p. 149, fig. 7a) figures
the mouth plates with two apical spines each, one behind the other. This is an
artist's mistake, as I have been unable to find such an arrangement in any of my
specimens, including four from the same locality at which Verrill’s example was

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS FISHER 47

taken (between Icy Cape and Cape Lisburne). The spine marked a is the suboral,
while that labeled ¢ is the first subambulacral.

Abactinal papulae are arranged in irregular transverse rows between the trans-
versely arranged groups of spines. There is a longitudinal supramarginal series, an
intermarginal, and an actinal, the latter extending about half the length of the ray.
Although the marginal and abactinal areas are large there are relatively few papulae
to an area—one to three in the intermarginals.

Large straight pedicellariae occur very sparingly in the intermarginal channel
and in the actinal interradial channels, and occasionally on the lateral portions of
the abactinal area. These are compressed and thin, lanceolate, acute, as seen from
the side, and the jaws are narrow as viewed from the back. The tips are curved or
bent, and crossed; or one jaw is notched for the acute opposing jaw. This type is
found in large specimens, and, if at all in small ones, in the axillary channels only.
Rarely the jaw is tridentate. The largest of these pedicellariae measure 0.8 to 1.2
mm. Smaller but similarly formed ones are found in clusters on the oral, and less
often on the proximal furrow spines; and still smaller ones, on the furrow face of the
adambulacral plates.

Crossed pedicellariae of rather characteristic Leptasterias form are spaced, in
circles, around the abactinal spines, and form dense wreaths encircling the marginal
and actinal spines (not half-wreaths), and clusters on the outer side of the adam-
bulacral spines. In some specimens the furrow spines are practically without pedi-
cellariae (station 3252) while in others from the same locality small clusters are
present. In alcoholic specimens the pedicellariae are much more conspicuous than in
dried examples, and the marginal clusters are often in contact, especially distally.
When the abactinal spines are very small, as they sometimes are, especially in small
arctic examples, they are difficult to differentiate from the pedicellariae, so that the
abactinal surface has the appearance of being covered with a multitude of small
spinelets, as in Henricia. The dorsal pedicellariae of large examples measure 0.21
or 0.22 mm. (station 3252) to 0.25 mm. (station 3506).

Madreporic body fairly large, a little more than one-half r from center, and
partly or completely encircled by spinelets.

Variations —The extremes of variation include an arrangement of the abactinal
spinelets characteristic of Leptasterias cribraria on the one hand and of L. groenlandica
on the other. There are numerous specimens which if considered as isolated cases,
might be classified as aberrant examples of either cribraria or of groenlandica. These

are the intermediates.
Forma CRIBRARLIA (Stimpson)

Plate 8, Figures 1, la, 1d, 2, 2a; Plate 21, Figures 1,2, 4; Plate 22, Figures 3, 4, 5; Plate 24, Figure 2

The chief variations of forma cribraria (spitcbergensis) concern chiefly the size,
number, and spacing of the abactinal spinelets, and the relative numbers of them
which are slender, clavate, or subcapitate; the robustness of the abactinal plates,
and the spacing of the transverse ridges (for instance the number in the proximal
half of the ray); the number of superomarginal spines; the robustness of the marginal
and actinal spines; the abundance of pedicellariae, both crossed and straight.

48 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The conditions of the abactinal area affect the general appearance more pro-
foundly than variations of other parts of the body. A decrease in the number of
spinelets is generally accompanied by an increase in their size. A specimen from
station 3252 with R 54 mm. (pl. 21, figs. 1, 2), has very small spinelets of diverse
caliber, usually strongly clavate and 0.4 or 0.5 mm. long. Twenty or twenty-five can
be counted across the ray near the base. This example would be classed as typical
cribraria. A specimen of precisely the same size from station 3506 has the spinelets
about twice as robust—especially in the middle third of the transverse line—and there
are about 10 to 15 between the two superomarginal series, near the base of ray. This
might be classified as an aberrant groenlandica, but is rather more like cribraria,
although the spinelets are too thick. A cleaned specimen from this station shows that
the skeleton is much stouter than usual, and as a result the skeletal meshes are nar-
rower. In a somewhat similar specimen from station 3522 (pl. 22, fig. 3) the spine-
lets are upwards of 1 mm. long mostly, subcapitate or heavily clavate, and two or
three times thicker than in typical specimens. From station 3420, an example with
R 50 mm. has the spinelets mostly cylindrical, or else a trifle clavate in the carinal
series, about 0.75 mm. long, and well spaced, so that there are about 10 across the ray
near base. ‘These spinelets are in transverse series but they are not so small, so
numerous, nor so close together as in typical cribraria. There is also an indication
of longitudinal series; a considerable number of the supero and infero marginal plates
are monacanthid, and the adambulacral plates are irregularly monacanthid and
diplacanthid. This specimen, if considered alone, would be classified, hardly without
question, as L. groenlandica.

One feature which is common to slender spined and stout spined forma cribraria
and to the various modifications of forma groenlandica is the structure of the dorso-
lateral skeleton, in which the transverse trabeculae are dominant, and leave a series
of short but very wide papular areas. These areas are as well developed in a typical
specimen of groenlandica from the north of Asia (Stanford collection, No. 19, pl. 23,
fig. 1) as in typical cribraria from Bering Sea, many of them reaching from the
carinal to the superomarginal series of plates.

Forma GROENLANDICA (Ltitken)
Plate 8, Figure 1b, 1c, 2b, 3; Plate 21, Figure 3; Plate 22, Figure 1, 2; Plate 23; Plate 24, Figure 1

Forma groenlandica differs from cribraria chiefly in having fewer, larger, and
more widely spaced abactinal spinelets, which as a consequence do not show such an
obvious and definite arrangement in consecutive transverse combs. However, as
there are all sorts of intergrades it is difficult to lay down any definite rule. An
example of forma groenlandica from station 3485 has the abactinal spines in
definite longiseries about 9 or 10 at the base of ray, and also in definite transverse
series. There are usually only one or two superomarginal spines, while in cribraria
there are three, or even four, and in some but not all examples there are irregularly
one and two adambulacral spines. I can not find any constant difference in either
sort of pedicellariae. ;

I have no theory to offer in explanation of the wide variation in the species.
It is strange, in view of the fact that the young are carried by the mother, that there
have not been developed in various parts of the cireumpolar range—especially on the

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 49

southern margin—well marked local races. But the specimens from Bering Sea
exhibit pretty much the entire gamut of structural variations. Both formae are
found in the Arctic Ocean. Among the large number of specimens from station
3252, Bering Sea, all the largest examples are referable to forma cribraria—that is,
all having R over 30 mm., while about half of the small examples are forma groen-
landica or intermediate. In the list of specimens the forma found at each station is
marked—c for cribraria, g for groenlandica, and 7 for intermediate or “‘indifferent”’
forms.

FIGURE 2.—LEPTASTERIAS GROENLANDICA SHOWING THE STOMACH FULL OF YOUNG READY TO ESCAPE.
THE DORSAL WALL OF BODY AND PART OF THAT OF THE STOMACH HAVE BEEN REMOVED TO SHOW
THE VENTRAL STOMACH, WHICH HAS BEEN CONVERTED INTO A BROOD CHAMBER, THE FULLY
DEVELOPED YOUNG ARE SHOWN IN $iTU; ds, DORSAL STOMACH; g, GONADS; ™, MOUTH, SEEN
LOOKING VENTRALLY THROUGH STOMACH; ABOVE THE MOUTH 18 SHOWN THE FOLD SEPARATING THE
DORSAL FROM VENTRAL STOMACHS. X3

There_is a very characteristic feature of Bering Sea examples as compared with
those from,Greenland, Spitzbergen, and Kara Sea. The former generally have con-
spicuous clusters of pedicellariae on the outer adambulacral spines, and frequently
also smaller clusters on the furrow spines, although these may occur sporadically
when the outer spines are well armed. Rather often both sets are swell provided
with pedicellariae which do not differ essentially from the abactinal crossed pedicel-
lariae, though sometimes a little larger. This characteristic seems coyelated with
a somewhat greater abundance of pedicellariae elsewhere on the body.

50 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

In Greenland, Spitzbergen, and Kara Sea small examples the adambulacral
spines normally lack pedicellariae (where in Bering Sea specimens the outer spines
would have small clusters). In large specimens the outer spine of diplacanthid
plates may carry a conspicuous cluster of pedicellariae, but not the furrow spine.
Only a few specimens are large however; most of them lack adambulacral spine
pedicellariae, except sporadically.

A considerable number of dwarf specimens are among those recorded in the list
of localities, especially from Indian Point, Bering Strait (3582), Bering Strait (16591),
and Plover Bay, 10 to 25 fathoms (16584). Duplicate specimens of the last two
lots were examined by Verrill and are recorded (1914, p. 123) as L. arctica. The
Plover Bay specimens aré in poor condition. They are all small, with tumid rays
and numerous short spinelets. Some of them carry eggs in the stomach, a habit so
far as known, not characteristic of arctica. The adambulacral plates are mostly
diplacanthid, and the marginal spines have complete rings of pedicellariae. The
smallest specimen known to be carrying eggs has R only 12.5 mm., r 6 mm.

Young.—Very small specimens have that generalized appearance which renders
identification, in the absence of adults, extremely unreliable. It is difficult, in a
species which breeds when small (R 12.5 mm.) to differentiate between real young and
dwarfed specimens. There is a very real difference, in that the dwarfed specimens
have more spines. They remind one of dwarfed trees that have managed to add new
rings without much increase in size. Such are numerous small examples (some
carrying eggs) from Plover Bay, Siberia, and two examples from Indian Point, Bering
Strait, 17 fathoms (No. 3582), the largest of which has R only 13 mm., but which
resembles an adult of forma cribraria in miniature.

Undoubted young, with R 7 mm., have a 4-faced ray, with gently convex abacti-
nal surface. The abactinal spinelets, five or six across the abactinal area, already
are arranged in transverse rows, with a pretty definite longitudinal arrangement, in
about five series, in those specimens which are probably destined to become forma
groenlandica (station 3252). Of crossed pedicellariae there are a few abactinally and a
circle around the marginal spines; and of the straight a few along the edge of the fur-
row. There is one series of superomarginal, one of inferomarginal, and two of adam-
bulacral spines. The ebactinal plates, relative to width of ray, are very large and the
papular areas small. A specimen with R 14 mm. has eight or nine spinelets across
the ray near base, two inferomarginals on distal half of ray, one to five crossed pedi-
cellariae to each abactinal spine, and fairly well developed marginal wreaths. The
abactinal spines are quite regular, but I think the specimen was destined to become
forma cribraria. A breeding specimen of the same size (forma groenlandica) from
Bering Strait (No. 16591) differs in having complete circles of crossed pedicellariae
around the abactinal spinelets, mostly two superomarginal and two inferomarginal

spines with thick wreaths of pedicellariae, and large straight pedicellariae on the
axils of the rays.

Anatomical notes.

As mentioned under variations there is a good deal of indi-
vidual difference in the abactinal skeleton which is of the very open, irregularly reticu-
late type, gharacterized by slender plates and on the rays by unusually short but
broad meshes, one to three of which intervene between the carinal and superomarginal
series. The details are scarcely twice alike. The carinal plates are three or four

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 51

lobed and form a zigzag or sinuous series. The superomarginals and inferomarginals
are rather regularly four lobed, with longer dorsoventral than longitudinal lobes.
The dorsolateral skeleton is composed of irregular plates, rodlike, oblong, spatulate,
straight, bent, or irregularly three lobed, but with the long axis transverse to the ray,
except in the case of a few connecting plates between the consecutive costae. In the
thickness of these plates there is a difference of fully 100 per cent between specimens
from station 3252 and 3506, those from the latter having a heavy skeleton with
smaller meshes.

The skeleton is not basically unlike that of other species, and certainly not
greatly unlike that of L. miilleri the type of the genus. The development of large
dorsolateral papular areas or meshes in the skeletal reticulum must be regarded rather
as specific than generic, and is associated with the mostly transverse dorsolateral
ossicles, which Verrill believed to be of generic importance (Ctenasterias).

Internal anatomy.—The testes have the usual Leptasterias form, and when fully
developed fill most of the coelomic cavity of the disk and rays, impinging upon the
stomach, and occupying all available space between the dorsal and ventral divisions.
They press the hepatic coeca against the dorsal body wall, and reach three-fourths
the length of ray measured on side. Each is roughly spindle-shaped, and consists
of a central canal or axis, from which branch off a number of very irregularly lobulated
major divisions (usually toward the mesial line of ray) and numerous very irregular,
subdivided, small lobes which fill in pretty much all the available space. The duct
passes downward about opposite the first superomarginal plate, and opens by a pa-
pilla, on the actinal surface, close to the interradial line. There are usually, here,
one to three large lanceolate straight pedicellariae. The duct has ciliated epithelial
lining (Lieberkind).

The ovaries are very similar to those of other species of Leptasterias, as, arctica.
They were found to be full of large and small oocytes at the same time that the young,
nearly ready to shift for themselves, were still being carried by the mother.

Lieberkind found a specimen with nine ovaries, while the tip of the tenth was
hermaphroditic, the eggs lying along the wall and the spermatozoa in the middle.
(1920, fig. 4, p. 125.)

The stomach is of the usual form, and very spacious in the female by whom it is
used as a brood pouch. The intestinal coecum is rather triangular in general form
with three thick lobes radiating from the base, the whole about one half r in length.
The hepatic coeca are large and extend well toward the extremity of the ray.

Care of the young.\\—The eggs after emerging from the oviduects, which open
ventrally as in other species of Leptasterias, are probably kept for a short time under
the disk. It is usual for species of Leptasterias to form a temporary nursery by
drawing the rays together and raising the disk. Fertilization probably takes place
While the eggs are here. They are then taken into the stomach, where development
ensues. Specimens from Plover Bay, with short tumid rays and with R only 12.5
mm., are already carrying eggs in the stomach (No. 16584). These eggs are rela-
tively very lerge, measuring 0.8 to 1 mm. in diameter.

Lieberkind, On a Starfish (Aaterias groenlandica) Which Hatches its Young in its Stomach. Ssertryk af Vidensk. Medd
Fra Dansk. naturh. Foren., vol. 72, p. 122, fig. 1, November, 1920.

52 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

In all of the other specimens at my disposal (stations 3251, 3252, 3506, 3541,
Bering Strait, No. 6591) the young have reached an advanced stage. As Lieberkind
writes, it is the ventral, eversible stomach which is used as a brood pouch. The
number of young is in proportion to the size of the adult. In a specimen with R
43 mm., 42 young were counted but in the specimen figured (R 47 mm.) there were
probably 100, although they were not disturbed, or counted. As the egg develops
it naturally increases in size until the stomach becomes greatly distended, and the
walls in preserved specimens are quite thin. The young are crowded into the radial
pouches, and are found also in the central portion over the mouth. I did not find
them in the dorsal stomach, which confirms Lieberkind’s observations, although
“at times the number of the eggs or the young was so great that they protruded
through the opening between the lower and upper part, forming a plug in the opening
(perhaps, however, this may be due to contraction on preservation).!

The young are frequently oriented with the actinal side against the stomach wall
and in some cases they may be seen through the tissue. In all the specimens except
that from Bering Strait, the young are separated—that is, are not joined to a central
mass of tissue as figured for L. arctica. The larval organ, as a lappet of tissue, is
still conspicuous, and arises from an interradius back of the mouth plates. In the
Bering Strait specimen (No. 6591) however the voung are not quite so advanced, and
are anchored by a strand of tissue to a central mass, much as in L. arctica.

In this case nearly half the young are outside the mouth, and the disk seems to
be arched in a natural manner as if to form an external brood chamber. This speci-
men is small (R 16 mm.) and it seems likely that it swallowed more eggs than there
was subsequently room for after complete development. Most of the large speci-
mens, known to be carrying young, have a few around the actinostome, although
it is possible these were expelled by contraction of the rays during preservation,
especially as the stomach in a few cases is partly extruded.

I think however that during the later stages of development it is perfectly
normal for the young to be extruded from the stomach. This might well take place
for purposes of respiration, even if the actinal gastric space were sufficient to accom-
modate the increase in bulk due to growth. Possibly during the whole course of
development the eggs and embryos are given a periodic “airing.””

As stated above, specimens carrying young were taken at stations 3251, June 14,
1890, bottom temperature 37.5° F.; 3252, same date, bottom temperature 44.8° F.
(probably an error) ; 3506, July 29, 1893, bottom temperature, 32° F.; 3541, August 10,
1893, bottom temperature 36.1° F.; and at Bering Strait, July 14, 1879. The breed-
ing season extends probably from May through August. On June 14 the young are
large enough to leave the mother. The recorded temperature ranges from 32° to
44.8°. The first is usually low and the second unusually high for the locality,
judging by records of the preceding and succeeding soundings. (See list of stations.)
Thirty-six and thirty-seven degrees are probably near the optimum for successful
development.

My observations confirm those of Lieberkind in respect to the entire absence of
food from the stomachs of all specimens carrying young.

8 Saertryk af Vidensk. Medd. Fra Dansk. naturh. Foren., vol. 72, p. 122.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——-FISHER 53

What is there about the eggs and young that inhibits the secretion of digestive
fluids?

Parasites —Two specimens from station 3252 were found to be infected by the
curious ascothoracid cirripede Dendrogaster—possibly Dendrogaster astericola."© In one
specimen with R 40 mm. were three of the parasites, the most symmetrical of which,
as seen from the side toward the actinal surface of the starfish, is shown in text Figure
3a. The branches occupied two rays. The second parasite was crowded into one
ray, while the third occupied the disk and the two remaining rays. The gonads of
this sea star, a male, were very poorly developed although in normal examples, the

at

FIGURE 3.—A PARASITIC CIRRIPEDE, DENDROGASTER SPECIES, FROM A SPECIMEN OF LEPTASTERIAS GROENLANDICA
STATION 3252; DORSAL VIEW, X 2; 9, SIGNIFIES NORMALLY DEVELOPED GONAD; 9’, GONAD REDUCED IN SI7E; 9”,
GONAD ABSENT. A PORTION OF THE BODY WALL OF PARASITE HAS BEEN REMOVED TO SHOW CONTAINED EGGS,
3a. VENTRAL VIEW OF A DENDROGASTER FROM A SPECIMEN OF L. GROENLANDICA HAVING THREE PARASITES,
STAIION 3252, X 2

testes are very large. In the other specimen there was only one Dendrogaster, but
this was larger. (Text fig. 3.) The gonads in this starfish are partly normal, and
partly retarded. Seven testes are normal or nearly normal, as marked ‘‘g.”’ Those
in the unoccupied ray seem to be quite as large as in unparasitized specimens. Two
gonads (in positions marked ‘“‘g’’’) are reduced in size, while the tenth (g’’) is entirely
absent, apparently because of the pressure exerted by a large lobe of the Dendrogaster.
The stomach and hepatic coeca of this specimen are so crowded by the parasite that

4 N. Knipowitseh, Révue des sciences nat., d St. Petersbourg, No. §, 1890 (October), pp. 353-357. Summary p. 374. Beit. 2.
Kennt. Ascothoracida, Trans. Soc. Nat. Petershourg, vol. 23, 1892, p. 14,

54 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

it is difficult to understand how anything like normal feeding could have been car-
ried on.

Of the four specimens of Dendrogaster, three are full of eggs, or very early em-
brvos. The fourth (one of the three from a single host) contains myriads of Cypris
larvae, rather poorly preserved. These were about ready to emerge. Presumably
they reach the exterior through the papulae.

Type locality —Greenland.

Distribution.—An arctic, cireumpolar species. Von Hofsten (1915) gives a map,
and summarizes the distribution as follows (p. 60): Spitzbergen; Siberian Ice-Sea to
east of the New Siberian Islands; Kara Sea; Barents Sea; Murman Coast and White
Sea; south, southeast, west and northwest coasts of Greenland; Ellesmere Land;
Cumberland Gulf. East coast of North America: Labrador, Gulf of St. Lawrence,
banks off Nova Scotia, Newfoundland, Bay of Fundy. He gives the bathymetrical
distribution as from 5 to 150 meters.

A. H. Clark (1920) records the species from Bernard Harbor, Northwest Terri-
tories, 2 to 3 fathoms, and from west of Cockburn Point, Dolphin and Union Strait.

The specimens listed below range from the Arctic Ocean (Point Franklin and
between Cape Lisburne and Icey Cape) to the southern part of Bering Sea (Bristol
Bay and north of Unimak Island), and to Kamchatka. The bathymetrical range is
from 743 to 71 fathoms. Ludwig (1886) records the species from St. Matthew Island,
and St. Laurence Bay, Siberia.

Specimens examined—Three hundred and twenty-nine. The letters signify:
c, forma cribraria; g, forma groenlandica; i, intermediates.

Specimens of Leptasterias groenlandica examined °
Rea ; . Bottom Num: ‘
tion , Locality Depth Nature of bottom penne, spect: Remarks
mens
Fathoms is ok

3251 | Bering Sea, 57° 35’ 50” N., 164° 05’ W____- 25.5 | Fine gray sand_...-...-.-..- 37.5 ¢#3/ U.S.N.M.

3252 | Bering Sea, 57° 22’ 20’ N., 164° 24’ 40” W_ 29°6!)Blackimud -2-- == - nee 44.8 j¢ 94128 | U.S.N.M. (carrying
young).

3253 | Bering Sea, 57° 05’ 50’’ N., 164° 27’ 15” W_ 36 Mind Shr dees sewencee aes aaee 35 e¢2] U.S.N.M.

3254 | Bering Sea, 56° 50’ N., 164° 27’ 50” W____- 46 Green mud, sand ___- 36. 2 ‘1 Do.

3255 | Bering Sea, 56° 33’ 30’ N., 164° 31’ 40’ W_ 43 =e 37 co6 Do.

3256 | Bering Sea, 56° 18’ N., 164° 34’ 10” W____- 49 Green mud, black shells-___| 35 012 Do.

3309 | Bering Sea, 56° 56’ N., 172° 55’ W________- 71 Greenimud = 2. = <-==<-52 37.9 02 Do.

3485 | Bering Sea, 57° 18’ N., 172° 34” W________ 62, |z5.5 Oe ssh ae Se ssctes sc Se 37.1 ol Do.

3506 | Bering Sea, 57° 33’ N., 165° 55’ W________- 36 Gray sand, mud_____-______ 32 ¢9*12| U.S.N.M. (carrying
young).

3507 | Bering Sea, 57° 43’ W., 164° 42’ N_________ 31 Fine gray sand___-.____ -| 37.5 ¢1]U.S.N.M.

3520 | Bering Sea, 59° 28’ N., 170° 577 W ! 38 Green mud, fine sand__..___| 32.2 el; Do.

3522 | Bering Sea, 57° 58’ N., 170° 09’ W____ | 41 Coarse gray sand, gravel___-| 35.7 | ©2) Do.

3529 | Bering Sea, 58° 36’ N., 172° 24’ W___- 56 Greentmud S35 22 ee 36.1 e02| Do.

3530 | Bering Sea, 59° 39’ N., 173° 53’ W___. 59 Dark green mud, fine sand_.| 34.9 23 Do.

3531 | Bering Sea, 59° 55’ N., 174° 17’ W____ 59 ‘Greanimnd s-Soe- se eseeeee 35.1 62 Do.

3537 | Bering Sea, 54° 45’ N., 169° 08’ W 49 Fine gray sand_______-- =s 38 02) Do

3541 | Bering Sea, 56° 14’ N., 164° 0S’ W

36.1 | ¢*16 | U.S.N.M. (carrying

young).
3542 | Bering Sea, mouth Bristol Bay._.________ 49 Dark mud, fine sand________ 39. 2 ¢1| U.S.N.M.

Point Franklin, Arctic Ocean__...._._____ 18.'6:)| Sand 22s. Sscce sees ese eh ee | ¢12) Aug. 31, 83; part No.

| 7625, with ZL. arc-
tica.

49 Black mud, fine sand

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 55

Specimens of Leptasterias groenlandica examined—Continued

Between Icy Cape and Cape Lisburne__..

Northwest of Bering Strait, 66° 58’ N., 171°
ay W.
at Oh Pt oe |

67° 07’ N., 173° 24’ E., Vega’s winter har-
bor, Pitlekai.

Just north of Vega’s winter harbor ........

et Ae A) ll

Port Clarence, Alaska...

65° 14’ N., 168° 35” Ww...

Kara Sea, 71° 54’ N., 67° 37’ E............
Jugor Strait, off Kara Sea...............-.
Spitabergen, Treurenberg Bay. ..........
Preobraschenije

Island, mouth of Khat-
anga Bay, Siberia (113° 30’ E. long).

4 Recorded by Verrill (1914) as L. arctica, p. 123.

|
Depth
}

Fathoms
15-18

Stones with clay
Sand, a little clayish..

Greenland, Egeden Minde................ 15-30

Greenland, northwest of Cape a 15-15

Spitzbergen, Shoal Point.................. 25-50

«2

ee ws

be)

Remorks

W. H. Dall, 1213,
1214 (6125).

North Pacific expedi-
tion, Captain Rod-
gers, No, 1362.

North Pacific expedi-
tion, No. 1349.

No. 7628, Point Bar-
row expedition.

6005, U.S.R.8. Cor-
win, 1580,

W. H. Dall, 1850,
3582 (657).

Wm. Stimpson, 1357.

Dr. Robt. White,
July 14, 1879 (1659)
small, 2 with em-
bryos in stomach.

3576 (part).

W. H. Dall, 308
(1720) 16584.

No. 1328, North Paci

fic expedition, Cap-
tain Rodgers (1
arm only).

Stanford Coll.

Riksmuseum, Stock-
holm, Vega exp.
Do.

FFF ees

Riksmuseum, Stock-
holm, Amundsen.
Riksmuseum, Stock-
holm, 0, Tarrell.
Riksmuseum, Stock-
holm, Sofia exp.
Riksmuseum, Stock-

holm, Nova Zembia
expedition.
Riksmuseum, Stock-
holm, Vega exp.
Riksmuseum, Stock-
holm, Spitabergen
expedition, 1861,
Do.

it) Riksmuseum, Stock-

holm, Vega exp.

56 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Remarks.—I have not been successful in discovering any constant characters U%
by which Stimpson’s eribraria can be separated from the earlier groenlandica of
Liitken or the later spitzbergensis of Danielssen and Koren. The last two species are
considered the same by Déderlein," Grieg,’ and von Hofsten (1915, p. 59). Verrill
at first united groenlandica and cribraria (1909, p. 553) but later (1914) kept them
separate. In the latter work he compares cribraria with spitzbergensis and concludes
that they are ‘identical but variable.” Professor Verrill’s material was not abundant,
much of it consisting of depauperate and young specimens. With a large series at
his disposal I feel sure that he would have adhered to his first decision.

Mortensen at first considered miilleri and groenlandica to be varieties of the same
species. He says: “Though a typical A. groenlandica as to habitus differs very much
from a typical A. milleri, especially of the var. islandica Levinsen, there is evidently
not asingle character to be relied upon by which they may be distinguished. I think
we must then conclude that they can not be maintained as two distinct species. The
form groenlandica must be regarded as a more or less distinct variety of A. miillert.”
In his latest publication, however, he definitely separates the two species.”!

Von Hofsten (1915, p. 59) adopts a middle course and classifies groenlandica as
an arctic subspecies of the boreal miillert.

Light is thrown upon the question by a comparison of breeding habits. Accord-
ing to Sars,” LZ. miilleri broods its young under the disk in a cavity formed by the
rays. In groenlandica the eggs and developing young are carried in the stomach.
It is inconceivable that the same species would develop two such fundamentally
different modes of caring for the young.

L. arctica, the Bering Sea equivalent of miilleri, is perfectly distinct in every way
from L. groenlandica.

Leptasterias obtecta Verrill (1914, p. 144) is referable to groenlandica. I have
examined the type, No. 1208, Museum of Comparative Zoélogy, 12 miles east of
King’s Island, Bering Sea, 17 fathoms. The specimen is characterized by slender
spinelets very numerous crossed pedicellariae, and smaller than typical dorsolateral
papular areas. The skeleton of one ray was cleaned. It is not essentially different
from forma eribraria with the exception that on the proximal two-thirds of the ray
the transversely elongated papular areas adjacent to superomarginal plates are not so
broad as is usually the case in eribraria. Between the mesial ends of these meshes
and the irregular series of carinal plates are one or two irregular, very distinctly
smaller, papular areas. The supramarginal skeletal meshes are therefore larger than
the others and wider than long in most cases. The adambulacral plates of the
proximal half of ray are diplacanthid; those of the distal half alternately monacanthid
and diplacanthid. The Henricia-like appearance of the dorsal surface is due to the
numerous crossed pedicellariae which on account of the slenderness of the abactinal

" The only tangible difference is that of the adambulacral spine pedicellariae of both forma cribraria and forma groenlandica
of the Bering Sea region, as noted under Variations. In large Greenland examples these pedicellariae are present, and they are
sometimes very scanty in small Bering Sea examples.

1* Die Echinodermen der Olga Expedition. Wiss. Meeresunt, new ser., vol. 4, Abt. Helzol. 1900.

1 Report of the Second Norwegian Arctic Expedition in the ‘‘Fram”’ 1898-1902. No. 13 Echinodermata, 1907, p. 13.

* Echinoderms from East Greenland, Meddel. om Gr¢énland, vol. 29, 1903, p. 70.

" Echinoderms of the British Isles, 1927, p. 143.

% Fauna littoralis Norvegiae, 1 Lief., 1846, p. 58. Mortensen Echinoderms of the British Isles, 1927, p. 142,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——FISHER 57

spinelets appear to be relatively larger than they actually are when compared directly
with pedicellariae of typical forma cribraria. A pedicellaria attached to a slender
spinelet has the appearance of being larger than when attached to a spinelet 50 to 100
per cent thicker.
LEPTASTERIAS OCHOTENSIS (Brandt)
Plate 12; Plate 27; Plate 28

Asteracanthium ochotense Brannt, Middendorff’s Reise in den Aussersten Norden und Osten
Sibiriens, vol. 2, pt. 2, 1851, p. 28. (Also var. macrobrachia and var. brachybrachia.)

Asterias ochotensts Perrier, Pédicellaires, 1869.

Leptasterias ochotensis Fisuer, Bull. U. 8S. Nat. Mus., 76, pt. 2, 1928, p. 1.

The specimens upon which the following account is based are a part of Brandt’s
original material. Doctor Mortensen kindly loaned a dried example received by
the Copenhagen Museum in 1871 and furnished photographs of another specimen
representing Brandt's variety “‘macrobrachia.” (Pl. 28, figs. 2, 3.) Later, Dr. A.
Djakonov, of the Academy of Sciences, Leningrad, sent four specimens in alcohol
which are of particular value, since it is possible to determine that the gonads open
ventrally. The largest of these examples in size and detail is closely similar to
Plate 28, Figure 2. Doctor Djakonov writes that the museum possesses two jars
with many alcoholic specimens and some dried ones all labeled by Brandt himself.
No actual type appears to have been indicated. I am especially indebted to Doctor
Djakonov for the donation of this material, which is now in the United States
National Museum. Aside from its scientific value it possesses great historic interest
for anyone concerned with the zoology of the north Pacific.

Diagnosis —Rays five; R40 mm.,r 9mm. R=4.4r. (R=3% to 5r, Brandt);
breadth of ray at base, 10-11 mm. Abactinal spines small; well-spaced, cylindrical
or clavate (with a differentiated irregular carinal series) interspersed with abundant
triangular straight pedicellariae; ray normally rounded, the marginals close together
and lateral in position; a single series of superomarginal (proximally sometimes
two), inferomarginal, and actinal spines larger than the abactinals, and forming a
. bristling lateral and ventrolateral armature; superomarginal and inferomarginal
plates of adult specimens connected in series by a smaller intermediate ossicle;
adambulacral plates with proximally two, then one (or with alternately one and two)
spines; abundant lateral and actinal straight pedicellariae.

Description —The abactinal surface of the disk and rays is beset with small
cylindrical or clavate, minutely thorny and usually terminally grooved spinelets of
divers sizes, varying from about 0.5 to 0.7 mm. in length by 0.3 to 0.4 mm. in
breadth. Smaller spinelets measuring 0.35 to 0.5 mm. long by about 0.1 mm. in
thickness are much less numerous, and are found toward the end of the ray, or one
or two may accompany the coarse spinelets. (Pl. 12, figs. 3 and 4.) The spines
are fairly well spaced, varying in this respect as shown by the figures, and the carinal
series is differentiated as an irregular line. As in most forms of Leptasterias the
dorsal spines increase in length from the carinals toward the superomarginals.

Abactinal skeleton (pl. 12, fig. 9) an irregular, open, reticulum, the primary
irregularly three and four lobed plates being connected by elongate intermediate
ossicles.

58 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Superomarginal series turns upward near base of ray reaching the abactinal
end of the interbrachial or axillary sinus; beyond middle of ray intermerginal chan-
nel is narrow. Except sometimes at base of ray, most of the superomarginals alter-
nate with a secondary ossicle (pl. 12, fig. 9) which persists for about three-fourths
the length of ray. A similar ossicle is present, but less regularly, between the
inferomarginal plates. They are present probably only in fully adult animals since
none are present in a specimen with R 27 mm.

There is generally but one superomarginal spine to a plate, cylindrical and
slightly tapered, of the general form of the slenderer inferomarginal spines, but a
little shorter. There is a single series of stout inferomarginal spines, decidedly
more robust than the superomarginals, and a single series of similar actinal spines
which extends about half the length of the ray. The spines of these two series are
1.5 to 1.6 mm. long, robust, cylindrical, slightly tapered as a rule, and round-tipped.

The adambulacral plates are diplacanthid at the base of the ray, then mona-
canthid but with the spine alternately on the furrow margin and set slightly back.
In the largest specimen sent by Doctor Djakonov the spines alternate fairly regu-
larly one and two as described by Brandt (R 40 mm.). The spines are about 1.5
mm. long, slender, terete, round-tipped, sometimes slightly tapered. ‘The first and
second pairs of adambulacral plates are in contact interradially, or the second pair
simply touch; the first plate is conspicuously longer than the second. The first
three are monacanthid.

The mouth plates are rather short and broad, the length of the median suture
being less than that of the first two pairs of adambulacrals. There are two apical
and one suboral spine, the latter a little longer and stouter than the first few adam-
bulacral spines. (PI. 12, fig. 10.)

Straight pedicellariae of a characteristic subtriangular form are very abundant
among the abactinal spines, in the intermarginal furrow, actinally, and along the
furrow face of the adambulacral plates. They are less numerous on the oral and
adambulacral spines. The abactinal vary in size from 0.22 to 0.4 mm. in length
and oceur on the surface of the plates between the spines, the largest being in the
neighborhood of the marginal plates. The lateral (mtermarginal) are 0.5 or 0.6
mm. long, while the largest axial ones are 0.7 mm. long by 0.5 mm. broad at the
base. The adambulacral are 0.25 to 0.3 mm., and the largest oral 0.4 and 0.5 mm.
long. (Pl. 12, figs, 5, 6, 7.)

Crossed pedicellariae occur in circles eround the dorsal and superomarginal
spines, and in clusters on the outer side of the inferomarginal, actinal, and adam-
bulacral spines. They are apparently not very abundant abactinally. The abac-
tinal pedicellariae are 0.22 to 0.25 mm. long, the terminal toothed part being about
0.055 to 0.08 mm. long. The adambulacral are 0.26 to 0.28 mm. long, while the
terminal toothed part, in profile, is 0.12 to 0.15 mm. long. The difference in the
two sorts is shown on Plate 12, figures 1 and 2.

Madreporic plate convex, surrounded by a circle of spines, and located at
mid r. The striae are coarse and irregular.

Papulae few to an area, and similar to those of other small Leptasterias.

The ovaries of the specimen examined are compact, with subglobose lobules
containing relatively large ova. The duct is attached to the interbrachial septum

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 59

on a level with the nearest inferomarginal plate. It passes between two dorso-
laterally oriented plates of the septum, turns abruptly downward and can be followed
to the ventral integument. With the specimen under water and in sunlight the
external aperture is readily demonstrated. In each interradius two apertures lie
close together near the outer end of the third adambulacral, consequently in the
narrow angle just external to the adoral carina. Near the apertures are several
conspicuous straight pedicellariae, such as Plate 12, Figure 5.

Color in life-—The dorsal surface varies from dirty bluish red, or reddish brown
with orange to yellow-brown-red and sometimes even green to dull grass green.
Actinal surface always vermilion, the tube-feet flesh color (Brandt).

Type——Brandt’s material is in the Museum of the Academy of Sciences,
Leningrad.

Type locality—Okhotsk Sea (Shantar Islands, Tugur Bay, Achae Island;
Mamtscha Bay).

Distribution —Okhotsk Sea.

Specimens examined.—A dried specimen labeled Mamtscha Bay, Zoological
Museum, Copenhagen. (PI. 27.)

Four alcoholic specimens determined by Brandt, labeled ‘‘ Mare Ochotense, Coll.
Middendorf”’ (Museum Acad. Sci. U. R. S. S. Nos. 1327, 1354).

Remarks.—So far as known this species is confined to the Sea of Okhotsk. How-
ever, H. L. Clark has described a very similar form, Asterias similispinis ™ from
Taraku Island, near Nemuro, Hokkaido. I have examined one of the six original
specimens (No. 1964, Mus. Comp. Zoél.), the largest of which has R only 25 mm.
(R=5r). In the available specimen (R 22 mm.) the secondary marginal ossicles are
not present. Neither are they present in small ochotensis. The rays are a little more
robust than in ochotensis of similar size and the straight pedicellariae average smaller
but are of the same triangular shape. Most of the marginal and actinal plates carry
one spine. The gonads, although dried, indicate that the species is a Leptasterias.

Until a good series of Clark’s species can be examined and its status definitely
determined, I suggest calling it Leptasterias ochotensis similispinis.

Subgenus HEXASTERIAS, new

Leptasterias polaris group; symmetry hexamerous; size small or large. Type
Leptasterias polaris (Miiller and Troschel) Fisher. North Pacific, Arctic, North
Atlantic.

Includes the following species: aequalis (Stimpson), alaskensis (Verrill),
alaskensis asiatica Fisher, alaskensis multispina Fisher, aleutica Fisher, asteira Fisher,
camtschatica (Brandt), camtschatica dispar Verrill, coei Verrill, coei truculenta Fisher,
heractis (Stimpson), hexactis vancouveri (Perrier), leptodoma Fisher, polaris (Miller and
Troschel), polaris acervata (Stimpson), polaris katherinae (Gray), polaris borealis
(Perrier), pusilla Fisher.

* Bull, Mus, Comp. Zodl., vol. 61, No. 11, 1908, p, 288.

60 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

POLARIS SECTION
LEPTASTERIAS POLARIS (Miiller and Troschel)
Plate 30, Figures 1, la—Ic, 2, 2a-2d; Plate 32, Figures 3, 3a; Plate 35
Asteracanthion polaris MtuLpr and TroscHeEL, System der Asteriden, 1842, p. 16.
Asterias polaris Stimpson, Proce. Boston Soc. Nat. Hist., vol, 8, 1862, p. 271.
Leptasterias polaris FisHer, Ann. and Mag. Nat. Hist., ser. 8, vol. 12, 1923, p. 599.

On the Atlantic side of North America Leptasterias polaris ranges from Disco
Island, on the west coast of Greenland, south to the Gulf of St. Lawrence (Anticosti
Island and Gaspé Peninsula), and the fishing banks off Nova Scotia. It extends into
Hudson Bay. Although no specimens have been taken between Hudson Bay and a
point east of Cape Lisburne, Alaska, there can be little doubt regarding the continuity
of its range.

In the Arctic Ocean north of Bering Strait, in Bering Sea, and in the north Pacific
just south of the Alaska Peninsula, a polymorphic race of polaris is m an extreme
state of flux. It is represented by at least three well-marked forms, two of which have
been described as distinct species. This race is L. polaris acervata, which takes its
name from the earliest described of its formae. We appear to have here three species
in the process of evolution. LZ. katherinae Gray, from the mouth of the Columbia
and Straits of Georgia, is a more stabilized form apparently derived from the an-
cestors of the Bering Sea complex. It greatly resembles L. polaris borealis (Perrier)
of Labrador. These two races are not genetically so close as their similarity would
indicate, if they are each marginal differentiations, geographically widely separated,
of an arctic and sub-aretic species (polaris-acervata). A possible elternative expla-
nation would be that L. polaris katherinae and L. polaris borealis represent isolated
segments of a preglacial species having a continuous distribution by way of a formerly
warmer Arctic Ocean. With the advent of a colder era the arctic segment of the
species became changed, leaving less modified and nearer the ancestral form that
portion of the species which dwells in the warmer waters of either side of the continent.

L. katherinae is apparently nearest to forma aphelonota, the only one of the three
phases of L. polaris acervata to be found at the south of the Alaska Peninsula. Else-
where I have mentioned the possible duplicate origin of the specimens classified in
this forma. Typical aphelonota may represent a geographic race, an offshoot of
acervata to the south of the Alaskan Peninsula.

L. coei Verrillis probably an ancient derivative of polaris stock. Indeed, one may
go further and assign a common origin to all the North Pacific 6-rayed species of
Leptasterias. :

Since the type specimen of Asteracanthion polaris Miller and Troschel came from
Greenland, the Greenland race is naturally the substructure for taxonomic elaboration.
There are two fairly well-marked formae of this race: One, forma polaris, has the
spinelets fairly uniform in length, without evident aggregation into convex groups.
Sometimes the spinelets are fairly well spaced; sometimes crowded. In the other
and apparently less common variety, forma subacervata, new forma, there are a vari-
able number of convex groups of somewhat enlarged spines. This is the biological
equivalent, in polaris, of forma acervata in L. acervata. Similarly forma polaris is

an equivalent of aphelonota. Both subacervata and polaris are present among Disco
Island specimens.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 61

For purposes of geographic analysis, Perrier’s Asterias borealis from Caribou
Island, Labrador (pl. 35, fig. 3), can be retained to designate the southern race of
Atlantic polaris. Perrier’s specimen was derived from the Museum of Comparative
Zoélogy, which still possesses topotypes (No. 1307). It is doubtful, however, if this
race is of any practical value, for its differentiation is based upon slight differences
in pedicellariae. In the crossed pedicellariae as seen from the side (pl. 30, figs, 2, 2a—
2d) the terminal lip does not extend so far down on the shank in the southern as in
northern examples. A similar latitudinal variation is apparent in L. polaris acervata
and L. polaris katherinae. The specimens from the very cold area of the Grand Banks
are intermediate between polaris and borealis, but a giant specimen from Western
Bank (resembling the type of L. acervata forma aphelonota) has pedicellariae of bore-
alis. (Pl. 30, fig. 2.)

Among the specimens of L. polaris borealis, one from Caribou Island and one
from Gaspé, Quebec (M. C. Z., Nos.; 1307, 1308, pl. 35, figs. 1,2), are remarkable for
having longer and more widely spaced spines. In the Gaspé example (R 150 mm.)
the spines are conical, 3 to 4 mm. long, spaced their length or more apart, and along
the radial line sometimes aggregated in groups, of which the central is largest. Other
specimens have well spaced but shorter and subcapitate spines. It is these latter
that resemble L. katherinae; while the Gaspé example would correspond to a poorly
developed L. coei.

To summarize, the following divisions of L. polaris are recognizable:

L. polaris polaris, high arctic waters of eastern North America.

L. polaris borealis, a very slightly differentiated southern form of the above.

L. polaris acervata, Arctic Ocean, Bering Sea, and region immediately south of
Alaska Peninsula.

L. polaris katherinae, off Washington, probably intergrading with above.

Verrill discarded the name polaris since the combination Asterias polaris (Miiller
and Troschel), 1842, was antedated by Asterias polaris Sabine, 1824 (for Ctenodiscus
crispatus). Miller and Troschel, however, described the species as Asteracanthion
polaris. On account of the structure of its gonads, which open ventrally, polaris is
more naturally classified in Leptasterias. The specific name polaris is therefore ten-
able, so long as it is not combined with Asterias.

Specimens of Leptasterias polaris examined

i Num- |
Locality Depth Nature of bottom | eae Collection
| mens
Fathoms |
Godhavn, Disco Island, Greenland..........- Dantiaickae Jx-------0+eeeen enn 4 Riksmuseum, Stockholm.
Disco, Greenland. ..............-.-..---..-+-«| (of 21 Re oad 1
Davis Strait, 63° 47° N., 62° 26’ W... 35 | Shells... 1 Do.
West Greenland (Omenak)...............-... 30-40 | Stones............. Lien Mpa De
West Greenland (Rittenbenk)..............- BY NRT nas cece otians Ltd Do.
West Greenland (Gothaab)..........--..---- sf6o%on {SSE Stay see ‘ Do.
Godhavn, Disco Island, Greenland... ......|... 0...) .. 20. c ee eenseen enue 56 US.N.M,, 15815,
scene bdhipmeaannss lanannagaadnestsencpa 6 U.S.N.M,, 15816; Howgate expedition
| Doctor Pavy.
Devic Ariattr Prcepticass) dsc. dds. 10 | sateen sesh 1 | U.S.N.M., 38844, N. P. Soudder.
Hudson Bay (Cape Churchill) .......... 2...) ......00.- Neepasiaaaumanh: 1) U.S.N.M.,, 39012, E. A. Preble.
Hudson Bay (Richmond Gulf)............... 1B |..----222eeeeeeneeee | 1) U.S.N.M,, F, Johansen.
' 1 specimen.

63160—30———5

62

BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Specimens of Leplasterias polaris borealis examined

Num-
Locality Depth | Nature of bottom | Ber of Collection
mens
Fathoms

Battle Harbor, Labrador--__--..-------------- 1] U.S.N.M., 19627.
Henley Harbor, Labrador si 2] U.S.N.M., 8834, W. A. Stearns.
Egg Harbor, Labrador. -.-----.--------------- 1} U.S.N.M., 36964, Owen Bryant.
Bonne Esperance, Esquimaux Bay, Labrador- 2| U.S.N.M., 5835, W. A. Stearns.
Vabradors.-- 222-21 - 2222-522 ae 2| U.S.N.M., 15813, 1376.
Caribou Island, Labrador_---.--~------------- 2 | Mus. Comp. Zo@l., 1307; topotypes.
Gaspé, Quebec-......------------------------ 1 | Mus. Comp. Zo6l., 1308.
Godbout, Quebec. --...-.-------------------- 2| U.S:N.M., 31922.
Grand Banks, Newfoundland--_- 1] U.S.N.M., 15814.
Edge of Grand Banks------------------------ 1| U.S.N.M., 24503.
Wieatern: Bank sits tes eee ee ERE CR ES 2] U.S.N.M~, 14339.
Off Cape Race, Newfoundland--_------------ 80 |, Pebbly-<-=--: ---- | 1 | U.S.N.M., Owen Bryant.
Grand Banks, Newfoundland, 46° 13’ N., 59 | Stone and shells__-) 1 | Riksmuseum, Stockholm.

51° 46’ W.
Grand Banks, Newfoundland, 47° 20’ N., 91 | Small stones, mud_ 1 Do.

§2° 29’ W.
Grand Banks, Newfoundland, 45° 53’ N., 50 | Small stones, | 4juy. Do.

51° 56’ W. shells.
Station 2463, off Newfoundland, 45° 44’ N., 45 | Broken shells 30°_- 1} U.S.N.M., 11177.

54°27" W. |
Station 2495, off Nova Scotia, 44° 10’ N., 440 | Herds olen ee 1 | U.S.N.M., 11480.

59° 23’ 45” W.

LEPTASTERIAS POLARIS ACERVATA (Stimpson)

Plate 30, Figures 3-5; Plate 31; Plate 32, Figures 1, 2; Plate 33, Figure 7; Plate 36, Figures 2, 3;

Plates 37—40

Asterias acervaia Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 271. Bruu, Proc.
Zool. Soe. London, 1881, p. 494-—Murpocu, Rep. Int. Polar Exp. Point Barrow, 1885,
p. 158.—Lupwia, Zool. Jahrb. 1886, p. 287.—SLapEN, Challenger Asteroidea, 1889,
p. 818.—Verritt, Shallow-water Starfishes, 1914, p. 107, pl. 27, figs. 1, 2 (not pl. 106,
fig. 3=dispar.). .

A. polaris var. acervata VERRILL, Amer. Nat., vol. 43, 1909, p. 545.

Asterias polythela Verritu, Amer. Journ. Sci., vol. 28, 1909, p. 68; 1914, p. 104, pl. 55, figs.
1, 2; pl. 70, fig. 8; pl. 72, fig. 2; pl. 84, figs 3, 4; pl. 79, figs. 1-2a— A. H. Cuarx. Rep.
Canadian Arctic Exp., 1913-1918, 1920, p. 8c (Port Clarence).

Asterias philippii Breux, Proc. Zool. Soc., 1881, p. 511, pl. 47, figs. 1, 1a.
and Mag. Nat. Hist. Ser. 9, vol. 18, 1926, p. 198.

Stenasterias macropora, A. H. Cuark, Rep. Canadian Arctic Exp. 1920, p. 9c (Port Clarence).

Asterias acervata borealis, A. H. Cuarx, Rep. Canadian Arctic Exp., 1920, p. 8c (E. of Cape
Lisburne).

Leptasterias polaris acervata FisHpr, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 599.

See Fisher, Ann.

This name is intended to designate the polymorphic representative of L. polaris

which inhabits Bering Sea and adjacent parts of the Arctic Ocean and north Pacific.
A “composite diagnosis’’ covering all of its phases is of no practical value since the
race does not exist as a composite but in the guise of three major formae which inter-
grade. These formae are: aphelonota Fisher, acervata Stimpson, and polythela Verrill.

The race as a whole differs from L. polaris polaris of West Greenland waters in

this accentuated polymorphism (one phase of which, polythela, is not represented
east of Alaska) and in the larger size of the straight pedicellariae.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 63

In the order of their degree of differentiation from typical polaris, the formae
stand as follows:

1. Abactinal spines small, close-set, of nearly uniform size, without regularity of
arrangement and without any trace of aggregation into convex groups—aphelonota.

2. Abactinal surface with a variable number of convex groups of spines (often
in three i)l-defined longiseries), the subcentral spine of the group usually larger than
the others—acervata.

3. Abactinal surface with large, conical or acorn-shaped spines, typically soli-
tary, in twos or occasionally larger groups, arising abruptly from an irregularly
reticulate spinulation of small, capitate, unequal spinelets—polythela.

The type of Leptasterias polaris acervata is No. 1336, U.S. National Museum,
Bering Strait, clean gravelly bottoms, 5 to 15 fathoms. (Further details under forma
acervata.)

Forma APHELONOTA, new forma
Plate 31, Figures 2, 2a—2c, 3, 3a, 3b, 4; Plate 32, Figure 1, la—lh; Plate 33, Figures 7, 7a-7c;
Plate 36, Figures 2, 3

This forma is based upon a giant specimen from station 2850, off the Shumagin
Islands, 21 fathoms, which has the following characters: R 245 mm., r 41 mm.,
r=6r; breadth of ray at base 43 mm. Rays long, rather slender evenly tapered to
the pointed extremity; rays constricted at base; disk rather small; interbrachial
suleus hidden by contiguous sides of adjacent rays; abactinal surface of rays well
arched, closely studded with small, bluntly pointed or obtuse spines of pretty uni-
form length, arranged in loose groups of two or three to eight, corresponding to the
size of the underlying plate; papular areas small; abactinal plates very numerous
closely and irregularly reticulated; superomarginal spines in vertical combs or
groups of three to six, usually four, to a plate; inferomarginal spines two or three to
a plate, conspicuously longer and heavier than the superomarginals, which in turn
are slightly longer than the dorsolaterals; two series of actinals, similar to infero-
marginals; adambulacral plates diplacanthid; dermal straight pedicellariae sub-
conical to broadly wedge-shaped with five teeth similar in general appearance to
those of acervata.

Description.—The well-arched abactinal surface is closely beset with uniformly
small spines of nearly equal length but of unequal thickness, arranged without any
order, in loose groups, running into one another, and inclosing relatively very small
papular areas. Reference to a cleaned ray shows that these groups correspond to
the larger, primary, abactinal plates, which carry five to seven spines, while the
smaller intermediate plates carry one to three. The spines are therefore arranged
in a close reticulum but this is masked by the very numerous abactinal plates, the
smallness of the meshes of the skeleton, their irregularity, and by the fact that the
spines are in groups, not in single file. The spines vary considerably in form, but are
usually cylindrical at the base with a slightly swollen, tapered, coarsely sulcated,
bluntly pointed or round tipped end. Others, especially on the disk and sides of
the ray are rather more elongate and more pointed than the majority from the median
radial area of the ray; and on the outer third of the radial area the spines are blunter
than they are proximally. Here and there a spine is perhaps a third longer and
considerably thicker than the average which are 2 to 2.5 mm. long and 1 mm. thick.

64 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The smaller spines are about 1.5 mm. long by 0.5 or 0.6 mm. thick varying with their
position on disk and ray.

Superomarginal plates rather low on the side of ray. Superomarginal spines a
little longer than the dorsolaterals and intermediate between these and the infero-
marginals which are decidedly stouter and longer. The superomarginals form a
vertical comb, or sometimes a group of three to five (occasionally six) to a plate the
median generally the largest. Their general form is clavate with a tapered, blunt,
sometimes compressed end, and the distal spines are faintly suleated. The proximal
spines are sometimes ‘‘eroded”’ having a flattened area on the upper (and under)
side, converging, in some cases, into a sort of shallow groove on the distal part of
the spine. There is no well defined supramarginal channel although the series of
papular areas is here more regular on account of the regular series of superomarginal
plates. Intermarginal channel practically non-existent owing to the relatively small
size of the intermarginal papular areas and the heavy wreaths of pedicellariae which
pretty well fill up the channel.

Inferomarginal spines conspicuously longer and stouter than the superomarginals,
two or three to a plate, clavate, more or less compressed, blunt to subtruncate and
frequently with definite eroded or sunken areas on the upper side of the spine. The
spines stand in an oblique series of two or a group of three.

There are two series of actinal plates with a rudimentary third series at the base
of some of the rays. The spines, one or two to a plate, are very similar to the infero-
marginals, even a little heavier near the base of ray and some are quite definitely
subspatulate and truncate. The outer side is slightly concave as in the case of the
inferomarginals, this taking the form of a groove or of a sunken area. The whole
actinal surface has a very bristling appearance, and one may count from the upper
edge of the superomarginals to the adambulacral spines eight or nine spines, or in
exceptional instances one or two more.

Adambulacral plates diplacanthid, and as is usual in such cases the slender furrow
spine of alternate plates stands further into the furrow than the stouter inner spine
of the intervening plate. In effect, the armature of alternate plates is set a little
more toward the outer edge of the plate. The spines are mostly terete, slender,
round-tipped, slightly tapered, or untapered, and many of them have a slightly
sunken area on the outer side near the tip, as in the case of the actinals. There
are four pairs of contiguous plates making up the adoral carina. On a few proximal
plates there are three spines. The spines lengthen toward the mouth plates, the
longest on the adoral carina (one to a plate) being about as long as the first five
adambulacral plates (6.5 mm.).

The actinostome is very small. The oral armature consists of two heavy spatu-
late, round-tipped or subtruncate apical spines to each plate, and a cylindrical sub-
oral spine similar to the adambulacral spines. The larger, more median, of the two
apicals is nearly twice as long (4 mm.) as as the short plate, and its breadth varies
on the different plates.

Papular areas smaller than the primary plates and with the distribution common

to the genus. The papulae are abundant and fill the spaces between the groups of
spines.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 65

Straight pedicellariae. (Pl. 31, fig. 4; pl. 32, figs. 1, la-1h.) Conspicuous
subconical, heavy, round-tipped pedicellariae with three or four, occasionally five,
terminal teeth are abundant abactinally, especially toward the sides of the area, on
the marginal and actinal plates, and also sometimes on the bases of the spines. Some
have broad, heavy jaws with three or four teeth and the back of the jaw sometimes
grooved, while in others the jaw rapidly tapers and the end is pointed with only two
or three teeth, or even without teeth. The heavier kind might be described as broadly
wedge-shaped. The largest measure 1.15 mm. long by 0.8 mm. broad at base (side
view), but many lateral and abactinal ones are only 0.7 mm. long. Smaller, broadly
lanceolate, compressed pedicellariae of various sizes down to that of the crossed sort,
occur on the adambulacral spines (especially those bordering furrow) and oral spines;
and others, also small and lanceolate, or lanceolate-ovate, occur on the actinal and
furrow face of the adambulacral plates.

Crossed pedicellariae (pl. 31, figs. 2, 2a—2c, 3, 3a-3b) are very abundant, forming
heavy wreaths around all the abactinal and superomarginal spines, and three-quarter
or half wreaths around the inferomarginal and actinal spines, filling up practically all
the space between the spines. The adambulacral spines are provided with large
clusters, and the pedicellariae are of different form from the abactinal, the jaw having
a longer cutting edge. These are a little narrower than the corresponding pedicellar-
iae of acervata and polythela. As compared with typical polaris, northern acervata,
and polythela, the side view of the abactinal pedicellariae shows the lip extending less
far down on the side of the jaw. Abactinal pedicellariae measure about 0.27 mm.
long, while the adambulacral are about 0.33 to 0.45 mm. long, the larger being less
numerous.

The madreporic body is circular, slightly convex, and situated about mid r.

Anatomical notes —The abactinal plates are very numerous, irregularly lobed,
and form a very close, irregular reticulum, the irregular spaces of which are usually
smaller than the primary plates. The supramarginal spaces are a little larger than
the plates. About 15 to 18 of the spaces can be counted across the base of the ray
between the two superomarginal series of plates, and about the same number of
larger plates, although the plates are so unequal in size and irregular in distribution
that there is no accurate method of computation. The superomarginals are a little
larger than the largest abactinals, are four lobed, and imbricate strongly, all being
spiniferous. The inferomarginals and actinals are smaller, decreasing in size toward
the furrow and are strongly united by their internally carinate dorsoventral lobes.
The ambulacral ossicles are moderately compressed, there being 10 in the space of
10 mm. (at the base of the ray).

The ovaries of the type are rather large spindle-shaped organs subdivided into
many lobules. They taper to a point and are therefore similar in form to those of
ZL. acervata and unlike the somewhat globular or sausage-shaped ovaries of L. camts-
chatica and allies. The ovary is attached rather low on the septum and the duct
turns downward, as in other members of the genus, to open ventrally, although the
apertures are not visible on this particular specimen.

Variations —This forma is divisible into at least two groups which may have a
different origin. The typical form has very numerous, close-set abactinal spines,
several superomarginal spines to each plate, and large bivalved straight pedicellariae.

66 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The bristling armature of the sides of ray, due to the vertical combs of marginal
spines is noticeable. (Stations 2850, 3214.) The less acervate examples of forma
acervata are near this subforma. ;

The other subforma has less closely placed abactinal spines, there are only one
or two superomarginal spines to a plate, and the straight pedicellariae are less heavy.
This variation resembles polythela with suppressed major spines. A specimen from
Kodiak has one or two spinelets of the carinal series enlarged. Old Harbor, Kodiak;
Port Clarence.

Type. —Cat. No. E 1491, U.S.N.M. ne

Specimens examined.—Type, station 2850, vicinity of Shumagin Islands, Alaska
(54° 52’ N., 159° 46’ W.), 21 fathoms, broken shells, bottom temperature 48.2° F.,
one specimen. (Pl. 25, figs. 2, 3%)

Station 3214, South of Alaska Peninsula (54° 13’ N., 163° 06’ W.), 38 fathoms,
gray sand, gravel, one specimen.

Old Harbor, Kodiak Island, August 11, 1888 (Albatross), one specimen.

Port Clarence Bay, Alaska, 2 to 3 fathoms, Canadian arctic expedition.

Forma ACERVATA (Stimpson)
Plate 30, Figures 3, 3a-3/, 4, 4a; Plate 32, Figure 2, 2a—2c; Plate 37; Plate 38, Figure 1

Asterias acervata Stimpson, 1862, p. 271.

This is the forma to which the type specimen of Asterias acervata belongs (No.
1366, U.S.N.M.). The type, however, is an extreme rather than a mean of the forma,
being on the border toward polythela. Forma acervata represents a rather hetero-
geneous series of subformae connecting aphelonota with polythela and its type is
intermediate between average acervata (pl. 37, fig. 4) and typical polythela. The
forma may be characterized as follows:

A large 6-rayed Leptasterias extremely variable in detail, but characterized by
having certain of the abactinal plates (often in three irregular series) larger than the
rest and armed with convex groups of coarse, capitate spines of which the central
spine is usually the largest; the ends of the spines subglobose to subconical, rather
finely striated; other abactinal spines well spaced, smaller, but similarly formed;
superomarginal spines one, two or even three to a plate, longer and slenderer, clavate
at times; inferomarginals similar, but heavier, one or two to a plate; one or two series
of actinal spines (one to a plate), similar to inferomarginals; adambulacral spines
frequently alternating one and two; furrow spine of alternate plates slenderer, tapering,
more advanced into furrow; consecutive plates with sometimes only one spine or
sometimes two; straight pedicellariae medium to large size, very broadly lanceolate
in profile, R=upward of 6r. Largest specimen, R 154 mm., station 3496.

Description —Forma acervata is difficult to describe in other than rather general
terms because it is so unstable. The abactinal surface is especially inconstant to
stereotyped detail, although maintaining a recognizable facies. The spinulation of
the marginal plates is scarcely more uniform. All specimens which show a tendency
to have convex groups of slightly to conspicuously enlarged but subequal spines are
included herein, the extremes being well shown on Plate 37, Figures 1 to 4; Plate 38,
Figures 1, 2. As mentioned above the actual type is intermediate between typical

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—-FISHER 67

or average acervata and those specimens of polythela with several prominent spiky
spines in a group, or a single spine surrounded by enlarged secondary spines, as shown
on Plate 38, Figures 2,3. A specimen with R 82 mm., r 18 mm, (R=4.5 r) has the
following features:

Abactinal spines, as the plates, are of two very diverse sizes. There are about
three longitudinal series, quite irregular, but in other specimens sometimes fairly
regular, of spaced heaps of capitate spines with swollen oblate hemispherical, sub-
globose, ovate (or in other specimens even acorn-shaped) vertically striated tips.
The central spine is much the largest of any of the group, while the subsidiary mem-
bers, unequal in size, form a circle or irregular group around it. The diameter of the
largest spines is about 1.5 mm. and that of the smaller 0.4 to 0.75 mm. Scattered
closely over the abactinal surface in irregular lines, rather vaguely defining a reti-
culum, are numerous usually unequal small spinelets (0.4 to 0.6 mm. diameter)
which duplicate in miniature the form and sculpturing of the acervate spines. Just
above the superomarginals is a zone variable in width, rather free from spinelets and
filled with numerous lanceolate, blunt, straight pedicellariae about 0.5 mm. long.
These are sometimes incipiently toothed. Smaller ones are scattered over the dorsal
surface. The spines are surrounded by small crossed pedicellariae in one or two
circles, the interstices between the spines of the groups being filled with them. In
alcoholic specimens the pedicellariae seem to arise from cushions of tissue surrounding
the spines and spinelets, and have the appearance of being everywhere numerous.
In large specimens there may be three tiers around the spines.

Superomarginal spines very variable in length and caliber, but slenderer and
sometimes larger than the major dorsal spines. In the type subforma each plate has
one spine except the distals which sometines have two. The spines are blunt, cylin-
drical-clavate, faintly striated longitudinally and the tips are often compressed.
Other variations have two and even three spines (in such cases smaller than when
one is present) on the proximal plates and three and four on the distal (for example
another larger specimen from station 3247, one from stations 3235, 3290, 3274, ete.;
see variations). In such specimens the abactinal acervate spines form usually
more rounded heaps, without a very prominent central member, but this is not
invariable (as stations 3233, 3274).

The type subforma has a row of inferomarginal spines similar to but longer and
heavier than the superomarginals, and a series of actinals similar to but a little
smaller than the inferomarginals. Other varieties have two equal or unequal infero-
marginals, and two series of actinals, the same specimens having also two or even
three superomarginals proximally. (See variations.) The superomarginal spines
have a dense wreath of about three tiers of crossed pedicellariae, while the infero-
marginal and actinal spines have thick clusters on the outer face. In the type
subforma the intermarginal channel, owing to the single row of superomarginal and
inferomarginal spines, is conspicuous (but it is practically obliterated in varieties with
multiple rows of marginal spines) and is armed with lanceolate blunt pedicellariae.

As might be expected, when there is little stability in the number of marginal
spines, the adambulacrals are variable. In the type subforma the spines most
commonly alternate two and one, the furrow spine of alternate plates being more
advanced on the furrow face of the plate, as in typical Asterias (although not so far).

68 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

This furrow spine is slenderer than the subambulacrals, tapering, bluntly pointed
or obtuse, while the other spines are heavier, subcylindrical, round-tipped, some-
times a trifle clavate or compressed. A considerable number of consecutive plates
may have but one spine, or may have two. The spines bear thick, subterminal
clusters of broadly lanceolate, unequal, straight pedicellariae, together with smaller
blunt crossed pedicellariae, the latter being more numerous on the subambulacral
than on the furrow spines, where the straight pedicellariae predominate and are of
larger size. Some of the specimens with numerous marginal spines have two and
three adambulacral spines on a portion of the proximal part of the ray. The third
(and outer) spine is shortec than the others. The armature of the furrow margin
appears crowded by reason of the more numerous spines and their heavy masses of
pedicellaviae; station 3274.

Four pairs of adambulacral plates are contingent behind the mouth plates
(fewer in the small specimens). The mouth plates are small, not especially sunken,
scarcely wider than the first pair of combined adoral adambulacral plates, and
their greatest length equals that of the first two or .wo and a half adambulacrals.
There is a very short, stumpy, pedicella-ia-clad spine divectcd across the mouth of
furrow (and reaching about to the middle); another longer spine, tapering and blunt,
directed over actinostome, and a third, suboral, still longer, slightly tapering, and
standing erect like the following adambulacral spines.

The madreporic body is prominent, convex, situated at about the middle of r,
and has irregular radiating striae.

Papulae numerous in small irregular groups dorsally and irregularly distributed
as the meshes of the skeleton; prominent just above, between, and just below
marginal plates, and present also between the actinal plates (when there is more than
one series), and adiacent to adambulacrals.

Straight pedicellariae (pl. 32, figs. 2 to 2e) are usually abundant on all surfaces
and on the adambulacral spines and vary considerably in size, the largest usually
occurring in the axillary channels. (See variations.) The larger straight pedicellariae
of acervata is a distinguishing feature from polaris. Arctic specimens have propor-
tionately more numerous straight and fewer crossed pedicellariae on the adambu-
lacral spines than have southern examples.

Crossed pedicellariae (pl. 30, figs. 3, 4) vary greatly in size in different parts
of the range. The abactinal measure from 0.22 to 0.8 mm. in length. The arctic
specimens have pedicellariae more closely resembling those of polaris than have
the southern Bering Sea forms. For measurements see explanation of plates.
Numerous crossed pedicellariae occur between the abactinal spines as well as imme-
diately surrounding them.

Color in life: Above, clouded with very dark brown; madreporic plate cream-
colored. Sides of rays and actinal surface, yellowish cream color (Stimpson).

Anatomical notes——The cleaned skeleton of the ray consists of a rather close
regular network of very unequal, irregular plates, there being a variable number
of large convex, irregularly lobed plates corresponding to the groups of spines.
These seem to have no order of occurrence except that the majority are near or on
the radial line. The intermediate and smaller primary plates are oblong, and of
various polyhedral forms with rounded corners. Twelve to fourteen plates can

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 69

be counted across ray between the superomarginals. The latter as well as the
inferomarginals, and actinals are arcuate, broad lozenge-shape or incipiently four-
lobed, and imbricate to form transverse as well as longitudinal series, lower and
adoral ends outermost.

The interbrachial septa are stout and strongly calcified, and their interradial
dimension is about one-half r. The actinostomial ring is stout, decidedly narrow,
so that the actinostome is small, and the odontophore is not conspicuous as in Asterias
amurensis. The ambulacral furrow is rather narrow and the ambulacral plates are
not crowded.

The intestinal coecum is of the irregular tubular form with a common saccular
portion adjacent to the intestine. The hepatic coeca are very large and extend to
the end of the ray. The eversible stomach has large radial diverticula, the central
portion being defined by the inner edges of the interbrachial septa. The septa as
mentioned above are about one-half rin breadth. The sides of the ray are divided
into a series of lateral pockets by means of dorsoventral partitions similar to those
of Asterias amurensis. Into these the lateral diverticula of the hepatic coeca extend.
The gonads are attached to the interbrachial septa about midway between the dorsal
and ventral surface and about one-fourth of r inward from the apex of the interbrachial
angle. The gonoduct passes outward and downward in the septum, and opens on
the ventral surface about one-fourth r from the interbrachial angle, between the
two rows of marginal plates, at the end of a low papilla often guarded by a major
pedicellaria. The papillae are very close to the interradial line, one on either side
for each interbrachium, and are difficult to find. The gonad is in form spindle-
shaped, a little over one-fourth to one-half R in length and consists of numerous
short lobulated branches surrounding a central duct. The eggs are large.

Variations.—As an example of the variations which may occur at one locality,
10 specimens from station 3233, 7.25 fathoms, Bristol Bay, are very informing. R
varies from 31 to 93 mm., seven examples being large. The range of difference
in abactinal armature is shown by Plate 37, Figure 1, Plate 38, Figures 1 and 2.
The first two are from station 3233; the last is from the Arctic Ocean, but a specimen
from station 3233 is practically the same. Of the 10 specimens: (a) One is Plate 37,
Figure 1; (5) two are between this and Plate 38, Figure 1; (c) five fall in a class with
the latter or have slightly more prominent heaps of spines. For instance, in three
of the five, the central spine is more accentuated. (d) One is like Plate 38, Figure 2.
The variation in superomarginal spines is as follows: (a) Usually three; (6) young,
one or two; (c) four specimens have generally two spines; two specimens (one of
which is young) have one spine or sometimes two on distal plates; (d) one spine
(typical of polythela). Of inferomarginal spines a has usually two, the others one.

Specimens from stations 3231 and 3232 exhibit the same class of variations,
ranging through the same amount of difference with the exception that there is no
example so near polythela as that in class d above.

Specimens from stations 3496, 3525, 3557, and No. 39984, off St. George Island,
have very large bivalved straight pedicellariae like those of the typical aphelonota,
All are from the region of the Pribilof Islands. Inasmuch as some of the specimens
of polythela from the same locality show a similar variation it would be interesting to
know the cause.

63160—30——6

70 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Of the above, two large specimens, No. 39984 and station 3496 are unlike the
usual run of acervata. They have much fewer, more widely spaced, larger spines.
No. 39984 abactinally resembles L. katherinae, but has the crossed pedicellariae of
acervata: station 3496 has a few larger spines here and there and one series of supero-
marginals. It may perhaps be a variant of polythela. Both have larger and more
widely spaced spines than f. aphelonota.

Young.—The smallest specimen has R 8.5 mm. (station 3233) and could not be
accurately identified if it were not accompanied by adults. The rays are moderately
stout, depressed, and the abactinal spines unequal, comparatively coarse, and rela-
tively few. Even at this early age a few spines here and there in the radial region
overtop the others and are usually distinguishable by having one to three crossed
pedicellariae on their bases. There is one superomarginal spine to a plate and one
inferomarginal, a little longer, each with several minor pedicellariae. Adambulacral
spines either one or two, rather irregularly; a few straight pedicellariae scattered
along the furrow face. One pair of adambulacral plates meet behind the mouth
plates. First two or three tube-feet biserial, then quadriserial in quincunx. Papulae
solitary, none actinal, but an intermarginal series present.

Another example with R 10.5 mm. is essentially the same.

Small examples with R about 30 mm. show most of the varietal differences of
the adult, but in a less accentuated form, since the spines are fewer. There are usually
one inferomarginal, one actinal, and either one or two superomarginal series of spines.
Sometimes a second inferomarginal spine of smaller size. Crossed pedicellariae are
plentiful. These and the absence of large bivalved denticulate intermarginal and
supramarginal straight pedicellariae will serve to distinguish young acervata from the
closely similar shore form, Leptasterias alaskensis, found along the Aleutian Chain
from Unalaska to Attu and Agattu.

The specimen figured by Verrill (1914, pl. 106, fig. 3), Nazan Bay, Atka, as young
acervata is referable to Leptasterias camtschatica dispar.

Type.—Cat. No. 1366, U.S.N.M., William Stimpson. The specimen figured by
Verrill (1914, pl. 27, figs. 1, 2) is not the type, as stated, but is Cat. No. 1367,
U.S.N.M., Arctic Ocean, Captain Rogers. It is, however, practically identical with
the type.

Type locality — Bering Strait, clean gravelly bottom, 5 to 15 fathoms (Stimpson).

Distribution —Forma acervata is found from the Arctic Ocean to Bristol Bay,
Bering Sea, Attu Island, and on the Siberian coast to Indian Point. Its known
temperature range is slightly higher than that of polythela, but the temperatures of
arctic stations are not known.

Specimens eramined.—Kighty-nine.

tion Locality

3231 | Bristol Bay, Bering Sea_..................
Seb es init tee te tee he
3233 |..... otk ee a eee |
3236 }..... Ie nr renee cimsc oroesetoe
3235 |..... “0 OS. a eee
3247 | Mouth of Bristol Bay_....................
3274 | North end Alaska Peninsula..............

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER

71

Specimens of Leptasterias polaris acervata forma acervata examined

Off Avatcha Bay, Kamchatka, 52° 46’ 50”
N., 158° 44’ 30” E.

Piover. Day, Siverie: - J...
Off Indian Point, Siberia... .............-
Head of Norton Sound, Bering Sea__...-..
6t. Michaels, Alaska. -...............-.-.
Chickagoff Harbor, Attu Island...........
Off St. George Island, Pribilof Islands_.._.

1006 | North coast Siberia, 67° 7’ N., 173° 24’ BE...

ef I MMII Boonie ceetnnwaumacwase

Depth Nature of bottom
Fathoms|
12 FENN eseretisniageetaamarterie
10.5 | Pebbles, stones..............
7.25 | Sand, pebbles...............
5 CET OUI os cerca here
bP Blagk on... sccee= ons!
17_—_| Pebbles, stones......-.....--
19 | Black sand, shells...........
18 Gravel, sand, rocks
16 AIO, BEIM ea cine emeeinere
16 A Re re a
15 OLIN era ee
41 Gray sand, stones, green
mud.
2 Black sand, sbells_.........-
26 RNIN acount eine canes dl
39 Gray sand, broken shells___-
6948 Green sand, pebbles......_.-
5-15 RINE VON Sk dnb tl cece
57 Rend, RAVES. . occnacoapea-
15-26 TINO Ws acetate aaa aioe
17 GrAWOls sdgsabs~ dance ces
I ae thea ele ae
I hb | Poker taersatoiapa peel epson nc be
Bt Aid cd couse cdtbddad am ah ppiepitein
MB) pieces eg hia hs dias imate
16 CTS, CRUE oo nnn econ
RT Reena wah Danan ae temrerd anes aearerts

Remarks

U.8.N.M.
Do.
Do.
Do.
Do.
Do.
U.S.N.M. (interme-
diate).
U.8.N.M.
1 Do.

Do.
U.8.N.M. (interme
diate).
U.S.N.M.

Do.

Do,

U.8.N.M. (interme
diate).

Albatross, 1006.

Wm. Stimpson, 1364,

type.
Captain Rogers, 1363;
1367.
W. H. Dall, 1880;
6111.
15714,
gale.
6112, W. H. Dall.
6086, W. H. Dall.
7622, John Murdoch.
15842, L. M. Turner.

beach after

39084, G. D. Hanna.

1 Riksmuseum, Stock-
bolm, Vega Expe-
dition.

1 Do.

Forma POLYTHELA (Verrill)
Plate 30, Figure 5, 5a, Plate 31, Figure 1, la—-1f; Plate 38, Figure 3; Plates 39, 40

Asterias polythela VERRILL, 1909, p. 68.

This forma comports itself more like a conventional species than does acervata.
When approximately typical (pl. 39, and 40) it is very easily recognized by reason of
the few large rather widely separated upright abactinal spines.
forma acervata by numerous intermediate forms of which the type of acervata is one.

Diagnosis.—A large to medium-sized, 6-rayed form, with rather slender to moder-
ately stout, depressed to well arched rays and small disk; abactinal surface of rays
with 5 to 30 large, conical or acorn-shaped spines, typically solitary, or in twos, or
occasionally more, borne on convex plates much larger than the others, and arising

It is connected with

72 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

from an irregularly recticulate spinulation of small capitate, unequal spinelets;
typically one or two superomarginal spines proximally, three to five short tubercles
distally; usually one inferomarginal, and one or two series of actinal spines; adam-
bulacral spines proximally two, distally one and two alternating; crossed pedicellariae
abundant; marginal and actinal straight pedicellariae frequently with broadened
toothed jaws. Differing from acervata in having larger conical dorsal spines with no,
or only a very few, smaller accessory tubercles. R=upward of 6.25 r, and upward
of 200 mm.

Description.—The type form as described and figured by Verrill, and abundantly
represented in the collection at my disposal is about the most easily recognized asteriid
of Bering Sea and the adjacent parts of the Arctic Ocean. It exists in nature, however,
in various guises which depart from the characteristic form above alluded to, and
bridge pretty completely the gap separating the type subforma from that of acervata.

The largest specimens, upward of 200 mm. major radius, are considerably bigger
than any acervata that I have examined.

R 200 mm., r 32 mm., R=6.25 r; breadth of the ray at widest part 40 mm.,
at base 33 mm. Rays swollen at base, very gradually tapered to a blunt extremity;
rays and disk usually well arched and sometimes inflated.

Abactinal surface beset with numerous, unequal, spaced capitate spinelets,
constricted above the base, then enlarged into a subglobose, acorn-shaped, or swollen,
subtruncate extremity, usually with weak longitudinal striations. These spinelets
are similar to those of acervata, and define rather irregular small sunken areas occu-
pied by papulae and pedicellariae. Arising from the general level are relatively few
much larger convex plates, bearing stout striated subconical or acorn-shaped blunt
or subacute tubercles, which in large specimens attain a length of 3 mm., or 5 mm.,
including the convexity of the plate. These tubercles vary in number and position
but are more frequent along the radial line than on the dorsolateral regions. Occa-
sionally they form three irregular series, but with more spines in the radial than in
either lateral row. They stand singly or in groups of two to five (in which case one
is usually larger than the others); and in some specimens certain of the minor spines
may form intermediate tubercles by enlargement. The disk may entirely lack the
large spines (type variety) or may carry upward of 20; station 3251. The number
of large spines on the rays varies greatly, ranging in general from 5 to 30 to each ray.
A very large example from station 3440 (pl. 40, fig. 2) has relatively few, mostly on
the outer half of the ray, while a medium-sized specimen from station 3251 (R. 78 mm.)
has about the maximum number in three series. As soon as the large primary spines
are accompanied by smaller ones so as to form an acervate grouping, the dorsal
surface approaches in appearance exaggerated cases of the type variety of acervata.
(PL. 38, fig. 2.) In definitely intermediate examples which can not be assigned to
either polythela or acervata the enlarged spines occur in groups as in acervata, but the
central spine greatly overtops the others and is conical in form. The superomar-
ginal spines may or may not stand singly on a plate.

Crossed pedicellariae are very numerous on the abactinal surface, densely cover-
ing the slopes of the plates leading up to the base of the large spines and surrounding
the minor spines in from one to three circles. Much less numerous straight pedi-
cellariae are scattered over the abactinal surface usually on the edge of papular

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——FISHER 73

groups. They rather increase in number and size toward the marginal spines,
reaching a length of from 1 to 1.25 mm. in the biggest specimens, and 1 mm. in an
example with R.170 mm. The form is broadly lanceolate to broadly oval, or even
subelliptical, more or less compressed, and the end of the jaws are either smooth
(station 3251) or are provided with one to five short, interlocking teeth (also station
3251), both being very large specimens. In a slender rayed variety from station
3530 and station 3518 the straight pedicellariae are fewer than normal.

What might be called the typical arrangement of marginal and actinal spines,
if there were not so many exceptions, is one series for each of the marginal rows, and
a single series of actinal spines—three in all; or two series of actinal spines (proxi-
mally at least) in the largest examples—four in all. In three very large specimens
from station 3251 two have a single superomarginal spine (except sporadically two)
on the proximal two-thirds or three-fourths of ray, then a transverse series of two or
three short tubercular spines; the third has mostly two spines except at the very base
of the ray where there is one, and on the outer fourth or fifth where there is a trans-
verse series of four or five short tubercles; of the first two, one has frequently two
inferomarginal spines and but one actinal series, the second has one inferomarginal
spine to a plate and two series of actinal spines and plates of nearly equal size; the
third has occasionally two inferomarginal spines, and two series of actinals, the
inner much the smaller. A second superomarginal spine, smaller than the primary
is not uncommon, and even a third may be present in an otherwise typical specimen;
station 3441. The distal plates very often have three to five tubercles, but may
have only one. Irregularity in the number of inferomarginal spines is less common
but occurs. The superomarginal spines are decidedly smaller than the dorsals,
subconical, sometimes compressed, with pointed, blunt, or irregular tips, and fre-
quently are acorn-shaped distally. The inferomarginals (and actinals also) are
abruptly larger if the superomarginals are small, or subequal if they are large, of
much the same shape, more or less clavate and slightly bent, the tips being blunt.
The superomarginal spines are provided with thick wreaths, and the inferomarginals
and actinals with half-wreaths of crossed pedicellariae, with much larger broadly
lanceolate to triangular-ovate straight pedicellariae scattered among them. The
intermarginal and actinal channels are proximally (and also usually distally) armed
with stout, straight pedicellariae of the general form of the dorsals but larger as a
rule, with typically broadened jaws ending in several small curved teeth. Some of
the pedicellariae (as station 3520) have the jaws nearly as broad as high, viewed
from the back, with parallel sides and gently curved margin armed with about five
or six teeth. They resemble somewhat the bivalved pedicellariae of Goniasteridae.

The adambulacral spines stand two on a plate on the proximal half or two-thirds
of ray, then they alternate one and two. The spines resemble those of acervata.
Alternate furrow spines are more advanced into the furrow, and are slenderer than
the subambulacrals which in turn are much slenderer than the actinals. The furrow
spines taper slightly, while the others are usually untapered, subcylindrical (or even
clavate), sometimes a trifle flattened. All have large subterminal groups of tapered,
unequal straight pedicellariae. The straight pedicellariae are toothless and have
narrow jaws viewed from the back; rather acute-ovate from the side. In large
specimens there are four pairs of contiguous adambulacral plates behind the mouth

74 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

plates, the width of this adoral ridge varying somewhat. The first two to four

plates have one spine each. j (sae
The mouth plates resemble those of acervata and each has three spines in similar

fashion.

The papulae are numerous in small not very crowded groups of 10, 15, or even
more, dorsally, intermarginally, between the inferomarginals and first actinals, and
between the first and second where present. The papulae alternate with the spines of
the second actinal row, or when this is not present stand, at least in large, specimens,
between the first actinals and the adambulacral plates. There is often only one, at
most only a few, papulae to each group actinally.

Madreporic body prominent large, slightly convex, with radiating striae. It is
situated about at the middle of r.

Straight pedicellariae are similar to those of acervata and vary in size but are
consistently large; only a very few specimens have them of inferior size. They
are generally very numerous, and have either smooth or notched edges.

Crossed pedicellariae (pl. 31, figs. 11; pl. 30, fig. 5) are very numerous and vary in
full sized specimens from about 0.25 to 0.29 mm. in length, 0.27 mm. being the usual
size of the abactinal. The adambulacral spine pedicellariae which have the usual
different form are about 0.3 mm. In arctic specimens the general appearance of the
pedicellaria is close to that of polaris, and as in forma acervata, the lip extends farther
down on the side of jaw than in southern examples. (Compare figs. 1 and 16, pl. 31.)

Anatomical notes—The main features of the anatomy are similar to those of
acervata. The intestinal and hepatic coeca are similar. The latter are relatively
very large, especially in the large specimens and reach to the end of the ray. They
fill the coelom of the ray and extend into the shallow lateral pockets of the coelom
formed by internal partitions. The latter are due to internal dorsal processes of the
marginal and actinal plates overlaid by membrane. The gonads of the largest
specimens open more in the axils, than on the ventral surface, but this opening is
below the superomarginal plates and is distinctly not at all dorsal as in Asterias, ss.

An examination of the skeleton reveals no important points of difference. The
abactinal plates in large specimens are quite irregular and are crowded in the median
region, the meshes being smaller than the skeletal intervals. The supramarginal and
intermarginal meshes are the largest. The marginal and actinal plates are regularly
four-lobed, the dorsal (internal) lobes being the longest and carinate, so as to support
a membranous partition, the series of which, along the side of the ray, act as a support
and occasion shallow lateral pockets of the coelom. These elongate dorsal lobes are
longest at the base of the ray, and they overlap very broadly the descending lobe of the
plate above, the latter being external. In the large specimen from station 3251,
there is a third series of small actinal plates which can not be seen from the outside.
As mentioned elsewhere the large tubercles are borne on very convex plates much
larger than the others. These plates seem to be among the first formed in the young.

Variations—The principal variations have been stated in the description.
It will be necessary only to give a brief summary here.

The number of rays is six, but there is one specimen with five rays from station
3518. The stoutness of the rays varies independently of the size of the specimen, the
condition depending partly on the maturity of the gonads, also somewhat upon the

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 75

amount of deflation at the time of death. Aside from these factors there is an actual
individual difference in ray breadth. The breadth of ray at widest part goes into
R 4.5 to 6+ times, in the largest examples; and 4 to 5.5 times in the medium-sized
specimens; in small examples there is not so much variability. Of course in the above
comparisons there is allowance to be made for the increase of ray length with age.

There are no two specimens alike in respect to the distribution, number, and size
of the abactinal tubercles, while their form ranges from that of an elongate cone to
that of a short acorn, or at the tip of ray to a depressed spheroid, and the tip may be
fairly sharp, to rounded. Not only is there variation in number between specimens,
but between the rays of the same example. The tubercles are perfectly solitary,
surrounded only on the plate by a dense felting of small minor pedicellariae, or there
may be 1, 2, 3, 4, or 5 accessory tubercles of various, generally smaller, sizes arising
from the same plate. There is scarcely ever more than two large subequal tubercles
arising from the same plate. Two extremes, that of acervate tubercles and that of
mostly solitary spines, are encountered at the same station, 3518, and at this station
also is a specimen with the disk bristling with spines (pl. 39, fig. 2) and others without
a single spine. About the same extremes are found at station 3251, at which were
taken the largest specimens (mostly with few or no enlarged tubercles on disk).
(Pl. 39, fig. 1.)

A curious variation is found in a specimen from the Arctic Ocean, north of
Bering Straits (steamer Corwin, No. 15819); R96mm.,r19mm. The dorsal surface
is typical, but the actinal spines are lacking opposite the first 8 to 12 inferomarginals.
From this point on there is a single series which extends nearly to the tip of the ray.
There is a single series of very short superomarginal spines and proximally a second
accessory series of shorter tubercles. The inferomarginals are abruptly much larger
and dominate the ambitus. The straight pedicellariae of the dorsolateral and mar-
ginal regions are large, with broad, toothed extremities. This is not a variety char-
acteristic of the Arctic Ocean for another specimen collected by the Corwin, also
No. 15819, Arctic Ocean, north of Bering Strait, has much shorter rays, a conspicuous
intermarginal channel, and a relatively broad area between the inferomarginal and
adambulacral plates occupied by a series of stout actinal spines.

Specimens not typical and variously diverging toward acervata were taken at
the following stations, no two of these specimens being closely similar: Station 3236,
one intermediate; 3252, one; 3253, one; 3254, two; 3277, three; 3506, one; 3520, two
intermediate, the proximal part of the ray of one example being acervata and the
tip, polythela; 4795, one; from this station a typical polythela. Arctic Ocean, north
of Bering Strait, one. (PI. 38, fig. 2.)

Young—From station 3254 along with adults and undoubted young is a tiny
specimen having six rays and R 4.5 mm. This is chiefly interesting in having the
abactinal surface covered with thin flat subcircular, overlapping plates from some of
which arises an erect spinelet; no papulae; two series of tube feet; mouth plates large,
like those of the Solasteridae, with two actinostomial spinelets and at the outer end
of the plate a larger suboral; one pair of adambulacrals touching behind the mouth
plates; one large adambulacral spinelet; actinostome big; a brief series of marginal
spineletsin each row. This specimen is of course quite unlike the adult and resembles
one of the Asterinidae more than an Asteriidae.

76 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Very small specimens have short, broad, depressed rays. The permanent spines
make their appearance early, a specimen with R 9.5 mm. having about 12 in three
series on each ray, each surrounded by a circle of relatively large minor pedicellariae.
The big plates and spines are among the first to appear and their early differentiation
shows that the character is deep-seated. The smaller plates of the adult are second-
aries, the prominent plates being the first that are acquired by the young sea star.
There are about five series of single dorsal papulae, one intermarginal, and none
actinal; no actinal plates; adambulacral spinelets alternating one and two; tube feet
in two zigzag series or in four, arranged in quincunx.

The young of acervata of the same general size (station 3232, 3233) have much
slenderer rays. An example of acervata with R 10.5 mm. has several of the radial
plates with two or three spines, the character showing itself early in this case, also.
The specimen with R 8.5 mm. shows the same tendency.

In specimens of polythela with R 15 to 20 mm., the adult characters are readily
recognizable, although the actinal plates are wanting and the pedicellariae are still
relatively few.

There is no evidence of an acervata stage in the development of polythela, nor
particularly, of the converse.

Type —Cat. No. 15820, U.S.N.M.

Type locality —Arctic coast of Alaska, U.S. R. S. Corwin.

Distribution—Bering Sea (Bristol Bay, vicinity of Unimak Island, Pribilofs,
St. Matthew Island, Petropavlovsk, Port Clarence) and the Arctic Ocean north of
Bering Strait, in from 2 to 59 fathoms, on gray, green, and black mud, or sandy mud,
exceptionally on coarse sand or gravel. Temperature range, 31.1° to 44.8° F. This
form appears to live on softer bottom and in colder water than does acervata. The
typical phase of the forma is found on mud or very fine sand, in water under 38°.
It is possible that the higher readings of 43° and 44° may be erroneous.

Specimens ecamined.—One hundred and thirty.

Specimens of Leptasterias polaris acervata forma polythela examined

Num- |
Sta- ; Bottom
Mon Locality Depth Nature of bottom ltemper-| Der of Collection
atures | SPecl
mens
Fathoms [fe sogtlte

$236 | Bristol Bay, Bering Sea____.........___ __ 14.75 | Gravel, sand, shells._.______| 39 1| U.S.N.M.

3244 | Mouth of Bristol Bay -_.______-_.________ 4.5 | Fine gray sand ied 1 Do.

3245 |---.. leet eee een ee | 11.5 | Sand, pebbles_____ 3 Do.

3251 | North coast Unimak Island__.._._________ | 25.5 | Fine gray sand 8| U. S. N. M. (very
. 8. N. M.
| large).

44.8 14 | U.S.N.M.

35 3 Do.
36. 2 13 Do.
37 19 Do.
3256 Senan DRE fe scte= Seen ase ate 49 Green mud, black shells__._- 35 2 Do.
3277 | North of end of Alaskan Peninsula________ 18 Aland, rocks: = 24 =. 43.2 3 Do.
3439 | West of Pribilof Islands, 57° 06’ N., 170° 41 Fine gray sand, shells____ 44 1 Do
35’ W. re j
$440 | West of Pribilof Islands, 57° 05’ N., 170° 48 Black mud, shells_ 1 Do.
4V' W. [RRR COAML ETS CSTE Aer aI De Finn ihe Beas) 5,

3441 | West of Pribilof Islands, 57° 04’ 20’ N.,} 51 |... do 9

170° 52’ 30” Ww. Vi oa a e ° oo

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 77

Specimens of Leptasterias polaris acervata forma polythela eramined—Continued

| Num-
Bottom
om Locality Depth Nature of bottom ftioas — Collection
atures | hens
}
|
. Fathoms °F,
3506 | Northeast of Pribilof Islands, 57° 33°.N.,| 44 | Fine gray sand... m1) 9) 0.8.NM
165° 65’ W.
3512 | North of Pribilof Islands, 57° 49’ 30” N., 38 Fine, sand, green mud_..... | 36.6 | 1
109° 27 W. |
3518 | Bering Sea, southeast of St. Matthew | 36 | Green mud........-...--- | 3.0 8 Do.
Island. |
Pere emer! TEE Or eal 37 | Black mud, fine sand___.__. lista] e Do.
3520 |...... OOo SoS E eed naasdinned 38 | Green mud, fine sand....... | 322) 2 Do.
$522 | Northeast of Pribilof Islands, 57° 58’ N., 41 Coarse gray sand, gravel..... 35.7 | 10 Do.
170° 09° W.
3529 | Bering Sea, 58° 36’ N., 172° 24’ W.......-. 1 Do.
3530 | Bering Sea, 59° 39’ N., 173° 53’ W......-.. MN Do.
3531 | Bering Sea, 59° 55’ N., 174° 17’ W.._...... 9 Do.
3611 | Bering Sea, east of Pribilof Islands-.. Q 1 Do.
47% | Off Avatchba Bay, Kamchatka-_ - 1 | Albatross, 1906.
4795 |..... UO ockas cane sae eorannnannaienaae~- nie 5 | Albatross, 1906 (with
acercata).
4796 | Off Avatcha Bay, Kamchatka._.......... 4 | Albatross, 1906.
re A NG) er a Wet ees ewcten 1/ Lieut. Geo. M.
Stoney, 15835.
Aretie Ocean, north of Bering Strait oc) co - c-| ened enpernnnnenenendeyecbowsloweenact 6| U. 8. R. 8. Corwin,
15818, 15819.
Uarchic coget Ofislaske ss Sei seot 15 iors tes otek ehh tak: 3... RSE See... 1} U. S, R. S. Corwin,
15820.
ea” 6 10 NN. 166" 08" da” W....--..---<-- WN le eaecarsioletvatesinrtactaptasia tea esa ited mec atae 1/ U. 8. R. 8. Corwin,
18823.
GON Sr Rte 106s Oe Tn i cehewond nade a Al chia ies Beetle 1| Lieut. Geo. M.
| Stoney, 158M.
1068 | 62° 39’ N., 177° 05’ W......-.......----.--4 BBs, b Chay nc rea AER RWS AKO wn Ses } 1 | Riksmuseum, Stock-
holm, Vega expe-
} dition.
1049 | Port Clarence, Alaska_._........- peas 4°16) |) Stones. cseceb ceo. ca ae. a} i Do.
pine a Lor est viel B75", (a ae ae SOT | Rectan lerahantia lit tink wali ls ileal | 1 | Riksmuseum, Stock-
° } holm, Vega expe
dition, ecersate
from this station.
1046 | Mouth of St. Laurence Bay, Siberia. ...__- 4-6 Pe, CNG dcdeteneca= eel iscan ne | 1 | Riksmuseum, Stock-
| holm, Vega ex-
pedition, interme-
| diate with acerrata.

LEPTASTERIAS POLARIS KATHERINAE (Gray)
Plate 33, Figures 1-6; Plate 34; Plate 36, Figure 1; Plate 41

Asterias katherinae Gray, Ann. and Mag. Nat. Hist., ser. 1, vol. 6, 1840, p. 179; Synop.
Starfishes Brit. Mus., 1866, p. 2—Perrrier, Révision des Stellérides, 1875, p. 67.—
VerriLt, Shallow-water Starfishes, 1914, p. 112, pl. 51, figs. 1, 2; pl. 52, fig. 1; pl. 83,
fig. 1.

Asteracanthion katherinae Mtuter and Troscuer, System der Asteriden, 1842, p. 19.—
Dvusarpin and Hupé, Hist. nat. Zooph., Echinodermes, 1862, p. 339.

Asterias douglasi Perrier, Révision des Stellérides, 1875, p. 69.

Asterias dubia Verritt, Amer. Nat., vol. 43, 1909, p. 545.

Pisaster grayi Verritt, Shallow-water Starfishes, 1914, p. 97. See Fisher, Ann. and Mag.
Nat. Hist., ser. 9, vol. 17, 1926, p. 556.

Leptasterias polaris katherinae Fisuen, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, 1923,
p. 599.

78 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Asterias Ikatherinae was very briefly described by Gray from specimens stated to
have been collected at the mouth of the Columbia River. The original specimens
are eight in number and were presented to the British Museum in 1838 by Lady
Katherine Douglas. They were entered in a preliminary register as coming from
Labrador. Doctor Gray, however, gives ‘North America, mouth of the Columbia
River” as the locality and, since another specimen has been taken in the Gulf of
Georgia, there can be little doubt that the entry made in 1838 was incorrect and
that Doctor Gray was right.

Through the courtesy of Sir S. F. Harmer, formerly director of the British
Museum (Natural History), and of Dr. R. Kirkpatrick, then in charge of the collec-
tions of Echinoderms, I received for detailed study four of Gray’s types. Doctor
Kirkpatrick had photographs made of the others, and forwarded me notes on the
specimens. Subsequently I visited the British Museum and examined all of the
types. At his request, one of the specimens having the original register number
attached, namely, 38.7.15.107, has been formally designated as type of the species.

It is necessary to correct an erroneous impression, due to a statement by Perrier,
that the actual types of Gray are not known with certainty. Perrier (1875, p. 67)
says: “Il est assez difficile de savoir avec précision ce que Gray a voulu designer
sous le nom d’Asterias Katherinae.”’

There is no shadow of doubt concerning the eight original specimens. They are
all identifiable with certainty. Five have the original register numbers glued to
them. Of the other three, two without either register numbers or Gray’s original
small letter, had labels, ‘As. katherinae”’ in Gray’s writing. The remaining one
alone without number, letter, or original name label of Gray was labeled (sub-
sequently, possibly by Professor Bell) A. katherinae, mouth of Columbia River
(No. 38.7.15.111). So it too must be one of the original eight.

In a manuscript list Gray entered the eight specimens as follows; the asterisk
denotes specimens which were sent to me.

a. Five-rayed 38.7.15.108 (a true katherinae).

b. * Six-rayed 38.7.15.110 (smallest specimen; type of A. douglasi Perrier).

c. Six-rayed 38.7.15.104.

d. Six-rayed 38.7.15.105.

e. Six-rayed 38.7.15.106 (“‘As. katherinae Gray”’).

f. * Six-rayed 38.7.15.107 (type).

g. * Six-rayed “variety” (38.7.15.109) (‘‘As. katherinae Gray”’).

h. * Six-rayed “variety” (38.7.15.111) (‘cotype’’ of following description).

The numbers of e, g, h, were lost and were replaced in 1920 by Doctor Kirk-
patrick; h was without any label but by a process of elimination is almost certainly
one of the “variety’’ specimens. I have examined all of the specimens; the last is
certainly a true katherinae and is referred to as a ‘‘cotype”’ in the following descrip-
tion.

Although No. 108 is a 5-rayed specimen and was labelled ‘‘ Asterias troscheli”’
by Bell or Perrier, it is a true katherinae.

All the specimens were dried without having been cleaned, so that the spines to
some extent are obscured by dried slime, sand, and sawdust, while the tube-feet are
more or less plastered over the oral and adambulacral armature. It was a simple

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 79

process to clean the type and specimens 6 and h. One ray of h was detached at the
base for a study of the skeleton and to determine the position of the gonads.

Although Perrier (1875, pp. 67, 68) redescribed A. katherinae, apparently from
the authentic specimens of Gray, he mentioned two 5-rayed examples. One of the
6-rayed specimens he described as Asterias douglasi, stating that there were also
several specimens without locality in the Museum d'Histoire Naturelle (p. 69). The
British Museum specimen (No. 110 of the foregoing list) was considered the type.
The dimensions (R 70 mm., r 17 mm.) and description given by Perrier apply to it.
It seems to me to be typical katherinae and certainly one of the original series since
there is clinging to it the same kind of sand that is attached to the other examples.
Asterias douglasi is therefore a synonym of katherinae. (PI. 33, figs. 3, 5.)

In his description of the specimens of katherinae Perrier (p. 68) writes that the
adambulacral spines are in a single series along the furrow margin. This is simply
an error of observation, for his description was not taken from one but from several
specimens, since he says: ‘Voici la description des individus auxquels doit étre
réservé, selon nous, le nom d’ Asterias katherinae, la description de Gray ayant
évidemment été faite d’aprés eux.” In none of the authentic examples are the spine-
lets in a single series. This mistakes was unfortunate since it led Verrill to infer the
existence of a second set of 6-rayed asterids having monacanthid adambulacrals,
which he named Asterias dubia (1909, p. 545), later Pisaster grayi (1914, p. 97). No
such second set exists, however, so that these names must be regarded as synonyms of
katherinae.

Diagnosis.—Rays six; exceptionally five; size rather large. Closely resembling a
thick-rayed L. polaris having irregularly and well spaced abactinal spines. Differing
from L. polaris acervata in lacking convex groups of abactinal spines, and from forma
aphelonota in having smaller straight pedicellariae and more widely spaced abactinal
spines. Abactinal spines very irregularly arranged, sparser than in acervata, clavate,
subcapitate, with the ends finely grooved; carinal spines not clearly differentiated; a
single row of superomarginal, one of inferomarginal, and two of actinal spines, sub-
cylindrical, often slightly clavate, robust; adambulacral spines normally one and
two alternating, the furrow spines of diplacanthid plates slenderer than the others,
and more advanced into furrow than that of monacanthid plates; very numerous
small ovate or broadly lanceolate straight pedicellariae. R=4.5 to 5.3 r.

Description —The measurements of the type, No. 38.7.15.107, British Museum
Register, are as follows: R 145 mm., longest ray and 137 mm. shortest ray; r 27 or
28 mm.; R=4.9 to 5.3 r; breadth of ray at widest part, just beyond base, 28 to 31
mm.; height of disk, 20 mm. Measurements of cotype No. 38.7.15.111: R 148 to 155
mm.;r33 mm.; R=4.5 to 4.7 r; breadth of ray at widest part near base (ray depressed
in drying) 33 to 35 mm.

The abactinal spines are numerous, not large, unequal, and arranged irregularly
on the irregular reticulum of dorsal plates. They are shortest on the radial area,
where the primary or principal spines are about 1 to 1.8 mm. long, increasing gradu-
ally to 2 or 2.5 mm. on the dorsolateral surface of ray. The irregularly arranged
radial plates, and to a less extent the adradial plates, carry, in addition to the principal
spine, which is thick-set, clavate or capitate, and longitudinally striated at the globose
or subtruncate tip, three to seven smaller but usually relatively slenderer accessory

80 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

spines, not uniform as to size or arrangement. These are, when perfect, also some-
what grooved at the top and beset with unequal points. Some of the radial plates
have eight or nine subequal spines of the minor size, in a group, or in transverse lines,
and connecting plates also bear a few spines of secondary size. The dorsolateral
plates, however, usually have but one spine, or at best only one or two small acces-
sories. In the cotype the abactinal spines are slightly stouter and rather less numer-
ous than in the type specimen. The maximum number on the radial plates is nine,
but is usually only five or six. It is only on plates where the spines are numerous
that there is any suggestion of the acervate or heaped arrangement characteristic of
acervata. Even this suggestion is not at all striking or conspicuous. Just above the
superomarginal plates there is a noticeable zone free from spines on the proximal half
of the ray.

The superomarginal spines, proximally one to a plate, and separated for about
the length of the spines, form a regular lateral row, which rises to the level of the
abactinal surface at the interbrachial angle so that here the intermarginal zone widens
abruptly. The spines are stout, subcylindrical, either slightly tapered or slightly
clavate, blunt, about 3 mm. long, and a little weaker than the nearest dorsolateral
spines. There are about 50 superomarginal plates, and those near the end of the
ray usually bear two or three stubby tubercles similar to the abactinal, while sporadi-
cally a proximal plate carries two spines.

The inferomarginal spines are very similar to the superomarginals though a
shade longer, and a few of the proximal plates quite irregularly carry two or even
three spines; the distalmost plates usually bear but one. The intermarginal zone is
conspicuous and at the broadest part of the ray is about twice as wide as the length
of the superomarginal spines. Just below the inferomarginal spines which are on
the actinolateral angle or curvature of the ray, is another narrower, spineless zone,
and then follow two series of actinal spines, somewhat stouter than the inferomarginals
the outer of which reaches the end of the arm, while the inner extends about three-
fourths the length, measured on side.

The adambulacral plates follow closely after the inner actinals and are rather
crowded, there being about 33 to 10 inferomarginal plates at the base of the ray.
The normal arrangement is one and two spines to a plate alternating, the single spine
standing in line with the interval between the two of adjacent plates. The inner
spine of these alternate plates is slenderer than the others, tapered and is carried on
the angle between the actinal and furrow surfaces of the plate. The furrow spine is
2.8 to 3 mm. long, the others, which are untapered to very slightly so, a trifle flat-
tened, and blunt, are 3.2 to 3.8 mm. long. The first 10 or dozen plates beyond the
adoral carina of contiguous adambulacrals usually carry two spines each, after which
the regular arrangement becomes fixed.

The mouth angle is formed of the mouth plates followed by four pairs of con-
tiguous adambulacrals, each of the latter carrying a single spine. This oral carina
is rather narrow, and its length about equals the width of actinostome. The entrance
to the furrow is narrow. The mouth plates bear each three spines, a somewhat
flattened truncate or round-tipped one directed over actinostome, a small one guard-
ing the mouth of furrow and carried on the inner furrow angle of plate, and a much
longer tapered suboral 3.5 to 4.5 mm. in length. (Pl. 34, fig. 3.)

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 81

The papulae could be determined by softening a ray of the cotype. Abactinally
at the base and middle or ray there are 15 to 25 in the larger skeletal meshes, and 10
in the small; in the intermarginal meshes, 6 to 10; in areas between the inferomarginal
and first actinal plates, five or six; between the two series of actinal plates, three or
four; and adjacent to the adambulacral plates, proximally two or three.

Two kinds of pedicellariae, straight and crossed, are present. The straight
pedicellariae are numerous, small, and practically all of one form, ovate or broadly
lanceolate, obtuse, as seen in profile. They are thickly scattered all over the abactinal
surface, intermarginally, actinally, and, with small crossed pedicellariae, form half
wreaths on the inferomarginal, actinal, and subambulacral spines, and conspicuous
clusters on the furrow spines (of alternate plates). A few quite small ones are found
on the furrow face of the adambulacral plates, and clusters are present on all of the
mouth spines. The largest are found in the intermarginal channel where they are
commonly 0.5 to 0.65 mm. long at the base of ray. Those on the actinal plates and
spines are a little smaller, while the numerous adambulacral ones are 0.3 to 0.375 mm.
long. The abactinal straight pedicellariae are 0.3 to 0.5 mm. long as a rule. As
shown by the figures, these pedicellariae are of a simple generalized form with rather
irregular minutely denticulate edges to the jaws. No trace of specialized unguiculate
pedicellariae could be discovered.

The crossed pedicellariae are quite small, mostly between 0.2 and 0.3 mm. in
length and form clusters or wreaths around the base of the abactinal and superomargi-
nal spines, and are scattered also on the surface of plates at a distance from spines.
Mingled with the straight pedicellariae, they form half-wreaths or bouquets on the
outer side of the inferomarginal and actinal spines, and a few are found in the adam-
bulacral clusters, where the straight pedicellariae greatly predominate. The exact
form of these pedicellariae is best appreciated from figures as there is some variation.
(Compare with pedicellariae of Z. polaris, L. polaris acervata and formae.)

The madreporic body is conspicuous, 5.5 mm. in diameter, very slightly convex,
with rather fine branched, radiating grooves or striae. The center of the madreporite
is about 0.4 r from the center of disk.

Anatomical notes.—The cotype was softened for cleaning and one ray removed.
The gonads although shrunken appear to have been voluminous. They are attached
to the interbrachial septum about midway between the dorsal and ventral surfaces
of the body, considerably below the level of the upturned line of superomarginal plates,
The gonoduct appears to turn downward but whether it opens intermarginally or
actinally can not be determined.

The dorsal skeleton, that is, all the plates above the superomarginals, is very
irregularly reticulate. The primary spine-bearing plates are usually irregularly
three or four lobed. These are rather outnumbered by elongate or faintly lobate
plates of diverse sizes which might roughly be classed as connectives. A portion of
the abactinal skeleton from the radial region of the cotype is figured on Plate 34
Figure 2. This is from the coelomic side. Just above the superomarginal plates
(pl. 34 fig. 1s) is a row of elongate transversely oriented connectives, between which
are large papular areas generally well supplied with straight pedicellariae. The
superomarginal, inferomarginal, and both series of actinal plates are regularly four
lobed and are imbricated in transverse as well as longitudinal series. Internally,

82 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

slender, compressed plates bind the superomarginals to the plates above (7) and to the
inferomarginals, either directly or by a second similar plate, closely joined in vertical
series; while one or two similar, smaller plates may continue the series across the
actinal plates, attached to their dorso-ventral lobes, which are themselves somewhat
carinate on the coelomic side. These plates thus form ridges separating shallow
pockets along the ventrolateral and lateral wall of the ray, into which the viscera are
normally crowded.

The ambulacral ossicles are not especially crowded, there being 12 in the space
of 10 mm. counting from the outer end of the contiguous oral adambulacral plates
(oral carina). The form of the pores is shown by the figure. (PI. 34, fig. 1, am.)

Variations —The smallest specimen of Gray’s type series is ‘‘376,” or No. 110,
which has R 70 mm. (longest ray), r 16.5 mm. (Perrier gives 17 mm.), breadth of ray
at base, 15 mm. It differs from the type and cotype in having a more closely knit
skeleton, there being fewer and shorter secondary connecting plates. As a result,
the spiniferous primary plates are less widely separated, the papular areas are rela-
tively much smaller than in the type, and the spines are very much closer together.
The median radial plates are larger than the other dorsal plates and the series is
clearly distinguishable both by reason of the larger size of the plates and by their
more numerous, close-set, capitate spinelets. In form these might be described as
very depressed globose with numerous, fine, rather deep vertical, or meridional,
striae, and there are 12 or even more to the larger plates, but only five or six are of
major size, the others being crowded on the periphery. This number is greater than
is to be found on similar plates of the type. The proximal superomarginal plates
carry one or two, sometimes three, longer, clavate spines, while the plates of the outer
half of the ray generally have three or four spines subequal to the abactinals, in a
transverse, somewhat arcuate, series. The inferomarginal spines, one or two to a
plate, are slightly longer and heavier than the superomarginals (especially distally is
the difference conspicuous) while the outer actinal spines are subequal to the infero-
marginals. The inner series is only starting, and consists of a few smaller spines at the
base of the ray. There are generally two adambulacral spines to each plate, or
distally two and one alternating. The pedicellariae (pl. 33, figs. 3, 5) are essentially
like those of the larger specimens. The subambulacral crossed pedicellariae have the
characteristic longer serrulate straight-edged jaw. (Fig. 3a.)

As previously mentioned, this specimen presents a much more compact appearance
than does the type. Apparently increase in stature is due primarily to an increase
in the size and number of secondary connecting plates, resulting in a separation of
the primary spine-bearing plates and a generally more open skeleton. The Gulf of
Georgia specimen is less compact than No. 110, but more closely knit than the type.

This is the specimen described by Perrier (1875, p. 69) as Asterias douglasi. I
have seen a very similar specimen of L. polaris borealis Perrier from Labrador (U. S.
Nat. Mus 1376). The resemblance is uncanny.

Professor Verrill (1914, p. 112) has described and figured the specimen from the
Gulf of Georgia (Museum of Comparative Zoélogy, No. 1181). Through the courtesy
of Dr. H : L. Clark I have received this specimen for study. I think it is referable to
Leptasterias p. katherinae, although as might be expected from its smaller size and

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 83

different habitat, it is not precisely like the type. Its dimensions are: R 85 mm.,
r18mm., R =4.7 r; breadth of ray at the widest part, just beyond base, 19mm. The
rays are not so inflated as in either the type or specimen /, and there is no zone above
the superomarginals of conspicuous width at the base of ray free from spines. The
abactinal spines are typical in form and arrangement and are relatively about the
size of those of the cotype, and a trifle larger than those of the type. The supero-
marginal spines are stouter and a little longer than the adjacent abactinal spines,
whereas in the type and cotype the adjacent abactinal spines are a trifle heavier.
The distal superomarginals bear a transverse series of three or two spines, which is
typical, and occasionally a proximal plate carries an extra spine, or a group of three
smaller spines. The intermarginal channel is narrower, as is also that below the
inferomarginal spines. The actinal plates are in two series and the spines are a little
thicker than the inferomarginals, clavate and obtuse or pinched at the tip. A few
of the plates bear a group of three or four smaller spines in place of the single normal
spine.

Professor Verrill writes: ‘‘The adambulacral spines are unusually numerous,
crowded in three or more rows. They stand mostly two on a plate, but often there
are three on part of the plates, or alternately two and three’’ (1914, p. 113). This
is misleading since plates with three spines occur very rarely. After considerable
search I found one on the outer third of the ray, and the third spine is small and
interpolated between the two regular spines. The mistake probably arose from
counting the spines of two consecutive plates, since the armature is crowded. Taking
at random one furrow margin and commencing with the first plate, the spines per
plate are as follows: five with one spine, then eight with two, then 1212212121
22:122:212:2:21221 222121212121212222121 2; then 32 plates
with alternately 2 and 1; then 312122111 2, ete.

The adambulacral spines are therefore normal for the species. The mouth
plates may have a second small spine on the actinostomial margin adjacent to median
suture.

The pedicellariae, both straight and crossed, are like those of the type in form
and are similarly distributed. The abactinal crossed pedicellariae are about 0.225
mm. long while the straight are usually 2 or 2.5 times as long. The latter are very
numerous, and the largest about 0.65 mm. in length are found laterally near the base
of the ray. The abactinal pedicellariae are figured. (PI. 33, figs. 2,6.) The crossed
pedicellariae seem to have a few more teeth than in the typical form.

Leptasterias p. katherinae seems destined to cause confusion. Verrill (1914, p.
112) describes and figures this specimen, No. 1181, as A. katherinae, but later in the
same work, p. 178, records it as Orthasterias merriami and figures (pl. 75, figs. 1-1d)
the dorsal, marginal, and adambulacral spines, as of merriami. The specimen is not
merriami, however. I suspect there is a mistake in this plate. Figure 1, a, 6, and ¢,
and Figure le do not represent typical ordinary dorsal spines of No. 1181 and would
certainly mislead anyone attempting to use the figure for katherinae. The typical
dorsal spines are similar to those figured in this work (pl. 33, fig. 1/), being stubby,
clavate to subcapitate and grooved at the tip. They do not carry straight pedicel-
lariae on the sides, as shown in Verrill’s Plate 75, Figures 1, a, 6.

84 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Type.—No. 38.7.15.107, British Museum (Natural History). Cotypes 38.7.
15.104, 105, 106, 109, 110, 111. The last, kindly presented by the authorities of
the British Museum will be deposited in the United States National Museum.

Type locality—Mouth of the Columbia River.

Distribution.—Mouth of the Columbia River; Gulf of Georgia, Washington.

Specimens examined.—Nine, as follows: Mouth of the Columbia River, the type
series from the British Museum (Natural History), Lady Katherine Douglas, eight
specimens.* Gulf of Georgia, Washington, No. 1181, Museum of Comparative
Zodlogy, one specimen.

LEPTASTERIAS COEI Verrill

Plate 29, Figures 3, 3a, 5, 5a

Leptasterias coei VerrtLL, Shallow-water Starfishes, p. 123, pl. 9, fig. 1; pl 17, figs.’ 1, (2
(young).

Orthasterias merriami VerriLL, Shallow-water Starfishes, 1914; p. 177, pl. 18, figs. 4, 5;
pl. 19, figs. 1, 2; (pl. 75, figs. 1-1d refer to L. katherinae).

Diagnosis.—Resembling a large 6-rayed L. tenera (Stimpson) with fewer, coarser,
and shorter spines. R112 mm., r 17 mm.; R=6.5 1; rays slender gradually tapered,
well arched; abactinal spines very well spaced, coarse, cylindrical, blunt, surrounded
by broad mats of pedicellariae; superomarginals conical, inferomarginals and one
or two series of actinals, longer, subcylindrical, terminally compressed, blunt or sub-
truncate; adambulacrals proximally monacanthid or diplacanthid; over most of ray
alternately monacanthid and diplacanthid.

Description.—Dorsal ossicles broad, strong, convex, and firmly united, with small papular
spaces intervening; spines well spaced, conspicuous, stout, but not very long, cylindrical or a little
tapered, obtuse. They stand singly on the ossicles and are surrounded at base by large wreaths of
pedicellariae. They form an irregular radial row and two or three indistinct or imperfect rows
each side of it, with smaller spines interpolated. The upper and lower marginal spines form equal
regular rows, one spine on each plate. These spines are larger and longer than the dorsals, mostly
conical and subacute; those of the lower series are distinctly longer, especially near the base of the
rays. Those in the upper row bear large dense, complete wreaths of pedicellariae; those in the
lower row have them only on the upper side. The two rows are separated by a naked channel which
becomes wide proximally and bears large, ovate major [straight] pedicellariae, clusters of minor
{erossed] pedicellariae, and a row of papular pores. At the base of the rays are two close rows of
actinal spines, similar to the lower marginals. Of these the outer row is close to the marginal, but
with small intervening papular pores, and it extends nearly to the end of the ray. Its ossicles are
rounded, convex, and nearly as large as the marginals. The inner row extends about half the
eee of the ray. Many of the actinal spines are flattened or acuminate at the tip; others are
obtuse.

; The adambulacral spines on the middle and distal plates stand one or two to a plate, alternating
irregularly, but on the proximal fourth of the ray they are mostly one to a plate. They are rather
stout, shaped like the actinal spines, but smaller and shorter, varying in size, mostly obtuse, but
the smaller ones often acute. Toward the mouth they become longer, more slender and subacute.
The peroral [actinostomial] spines are large and strong and nearly meet over the mouth. The
two apical peroral spines, which are much stouter than the adorals, but not so long, are straight,
tapered, a little flattened, obtuse; the smaller side-spine is about half as long, but of the same
shape. They bear small, ovate, forficulate [straight] pedicellariae. The epiorals and adorals are
similar in form, distinctly longer than those farther out, slender, terete, and evenly tapered. The

xu ane types have considerable sand adhering which might be described as dark gray, as it consists of blackish and translucent
(quartz ?), somewhat water-worn grains. Possibly it is only beach sand.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 85

adoral carina consists of three pairs of closely contingent plates besides the orals. Large ovate,
major pedicellariae occur between the interactinal spines. (Verrill, pp. 177, 178, O. merriami.)

I have examined the type of Orthasterias merriami figured by Verrill on Plate 18,
Figures 4 and 5, and the cotype, Plate 19, Figures 1 and 2 (Verrill, 1914). The
pedicellariae and spines figured on Plate 75, Figures 1 to 1d, do not belong to a speci-
men of this species but to L. katherinae. The specimen in question, No. 1181,
Museum of Comparative Zodlogy, I have examined. It appears to be true katherinae
and it is so listed (under Asterias) by Professor Verrill (p. 112). The record of
merriami from the Gulf of Georgia, based upon the same specimen, is therefore an
error (Verrill, p. 178).

I have quoted Verrill’s description of the type of Orthasterias merriami because
my three specimens from Shelikof Strait are smaller, and were taken at some distance
from the type locality, Glacier Bay.

The superomarginal spines of the type are slender-conical (1.75 mm. long) and
are noticeably smaller than adjacent dorsolateral spines, while the inferomarginals
(about 3 mm. long) are not at all conical but rather compressed-cylindrical, only
slightly tapered, with a blunt or subtruncate tip. The actinal spines have the tips
more compressed and often chisel-shaped or slightly gouge-shaped.

In the cotype (Berg Bay, Glacier Bay) the proximal adambulacral plates are
nearly all diplacanthid, the furrow spine of alternate plates being slenderer, and set
a little farther into the furrow. Near the end of the ray the plates are alternately
monacanthid and displacanthid. The abactinal spines show a slight fluting, well
marked on the proximal carinals. It is only faintly visible on some of the spines of
the type, which, however, is not so well preserved.

Crossed pedicellariae are very numerous and occur in broad bands surrounding
the abactinal and superomarginal spines as well as in thick clusters on the outer
side of the inferomarginal and actinal spines. Mixed with straight lanceolate pedi-
cellariae they form small clusters on the adambulacral spines. Between the primary
abactinal spines there are numerous small clusters attached to smaller plates while
secondary abactinal spinules, smaller than the conspicuous primaries, carry corre-
spondingly small wreaths. The abactinal pedicellariae measure 0.30 or 0.31 mm.
long. (Pl. 29, fig. 3.) Those of the adambulacral plates are of about the same size
but somewhat different form, as is usual. (PI. 29, fig. 3a.)

The straight pedicellariae (pl. 29, figs. 5, 5a) are small (about 0.5 mm. long)
compressed, ovate, or broad-lanceolate. They may have even more compressed
jaws than those figured and are very diminutive as compared to the broad-toothed
ones of L. coei truculenta. The largest specimen from Shelikof Strait (station
4292) measures R 63 mm., r 12 mm., the rays being slender, tapered, and of about
the same form as those of the type of merriami figured by Verrill.

The pedicellariae have the characteristic form, but the straight are slightly
longer (0.7 mm. in the intermarginal channel). Some of them have two or three
teeth, although the jaws are narrow. None of the intermarginal or actinal straight
pedicellariae approach the large size and breadth of valve characteristic of truculenta.
The abactinal straight pedicellariae are quite typical, of a compressed broad-lanceo-
late or ovate form, with only a slightly crenulated distal margin. The abactinal

6 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

CO

straight pedicellariae, like those of the type, measure about 0.28 mm., while the
adambulacral measure 0.25 mm.

These specimens have the characteristic heavy apical oral spine, each angle
pair with a small companion at each side. The first 9 or 10 adambulacrals are
mostly monacanthid, the rest alternating monacanthid and diplacanthid. There is a
single row of actinal spines extending about half the length of the rays and the carinal
spines are conspicuously heavier than the dorsolaterals. Both sorts are subeylindri-
cal, not long, with rounded very finely striate tips. The coarsely striated madre-
porite is slightly adcentral to the middle of r.

The type of L. coei was taken in Berg Bay, 10 fathoms, and is figured by Verrill
(1914) on Plate 17, Figure 1; the cotype of Orthasterias merriami was taken in 10
fathoms, Berg Bay (it is labeled ‘Berg Bay, Glacier Bay’’). Another smaller speci-
men of coei is figured as the type on Plate 9, Figure 1, and as cotype on Plate
17, Figure 2. Unfortunately, neither specimen of coei can be found in the Yale
Museum.

I think ZL. coei, the type of which measured only R 30 mm., is undoubtedly a
young form of the cotype of O. merriami. The cotype of merriami seems not dis-
tinguishable specifically from the type (from off Juneau, Alaska, 20 fathoms). If the
types of coei are really lost, as seems probable, the species will have to rest upon the
type of merriami. The name coei has page priority over merriami.

Type.—Y ale Museum (probably lost).

Type locality —Berg Bay (Glacier Bay), Alaska, 10 fathoms (Harriman Alaska
expedition).

Distribution.—Juneau to Shelikof Strait, Alaska, 10 to 102 fathoms.

Specimens examined.—Off Juneau, 20 fathoms (type of merriami). Berg Bay,
Glacier Bay, 10 fathoms (paratype of merriami). Station 4292, Shelikof Strait, 94
to 102 fathoms, blue mud, fine sand, bottom temperature 39.8° F. (three specimens).

LEPTASTERIAS COEI TRUCULENTA, new subspecies

Plate 24, Figure 3; Plate 25; Plate 26; Plate 29, Figures 1, 2, 4

Diagnosis.—Rays six; size large; in general appearance similar to L. coei (adult)
but with more numerous abactinal spines, on the average longer; without conspicuous
interspinal clusters of crossed pedicellariae, either alone or surrounding small second-
ary spinelets (these having developed into primary spines); with numerous very large
abactinal, intermarginal, and actinal straight pedicellariae, having oblong to squarish
coarsely toothed jaws, in addition to smaller compressed ovate ones; abactinal spines
finely to coarsely fluted. Rays long, rather slender, arched in alcoholic specimens;
disk small; interbrachial angles very sharp; skeleton well covered by opaque skin,
and abactinal spines with a thick pad of pedicellariae which in alcoholic specimens
frequently touches that of neighboring spines; spines upright bristling, rather uni-
formly spaced, robust, the abactinal without definite serial order, but the marginals
and actinals (one to three series) in regular, rather close series. R 220 to 225 mm.,
r 30 mm., R=7.3 r; breadth of ray at base 30 mm. (type). R 121 mm., r19mm.,
R=6.3 r; breadth of ray at base 20 mm.; station 4777.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS--FISHER 87

Description.—Abactinal spines well spaced, prominent, cylindrical at the base,
with a slightly swollen, longitudinally striated conical or acorn-shaped round tipped
to subacute distal half. The carinals are usually distinguishable by being a little
closer together rather than by superior size; but often they are not distinguishable
from the other spines. The relative thickness and form of the spines is decidedly
variable. Sometimes they are fairly short and robust, the acorn-shaped portion
occupying almost the entire length; or the spine is distinctly styliform. The large
type from Captains Bay has coarsely grooved subacute, unusually widely spaced
spines which in the dried condition appear slender-conical. The plates are distinctly
convex, which adds to the apparent length of the spines of this species. The dorso-
lateral spines are not in regular longiseries, not in quincunx, but for short distances
sometimes stand in longiseries or oblique series. There are the equivalent of about
three to five longiseries in grown specimens.

Superomarginals cylindrical, tapered, bluntly pointed, less often obtuse, some-
times asymmetrical or even a trifle bent at the tip, which is also usually a trifle
swollen. The spines stand regularly one to a plate, are a little longer than the
abactinals, and the series curves upward to the abactinal surface at the interbrachial
angle. Intermarginal channel narrow, definite, very well marked.

Inferomarginal spines still longer than the superomarginals (3 to 5 mm. in
medium-sized to large specimens), cylindrical at base, tapering on outer half, and
finally at the tip sometimes decidedly compressed, blunt or subtruncate. Most of
the terete spines are bluntly pointed.

Actinal spines in one to three series (according to size), similar to the inferomar-
ginals with which they also form transverse combs, the narrow intervals between the
series being subequal. In average sized specimens, the outer row of actinal plates
extends to within a short distance of the ray-tip while the second series reaches up-
ward of two-thirds the length of ray (R=124 mm.). A specimen from station 4777,
with R 114 mm., has only a single series of actinal spines, which is exceptional. In
the very large type from Captains Bay there are three series of actinals to the middle
of ray, and two nearly to the end. (PI. 26, a,' a,? a.)

The abactinal skeleton is irregular, the plates overlapping broadly by their
three or four usually unequal asymmetrical lobes. The marginals and actinals, on
the contrary, build very regular series and imbricate longitudinally and transversely
very strongly. The entire width occupied by the actinal and marginal plates is much
less (about two-thirds) than the breadth of the dorsolateral area. The marginals
and actinals are proximally asymmetrically four lobed, and beyond the proximal
fourth of ray become longer than broad, and appear superficially more often three
than four lobed. The central portion of the plates is strongly convex.

Papular areas relatively small (those just above superomarginals largest), three
to six, or more papulae to an area abactinally where they are not conspicuous owing
to the thick mats of crossed pedicellariae surrounding spines. There is an intermar-
ginal series of areas, and one to four actinal series, proximally, to one to three distally
according to size of specimen.

Adambulacral plates compressed, about 33 to 10 inferomarginals at middle of
ray. The armature is ideally alternately monacanthid and diplacanthid, but the
regularity is disturbed at base of ray where the plates of adoral carina and a few

88 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

following are usually monacanthid. In the very large type most of the plates are
diplacanthid. The furrow spine of diplacanthid plates is shorter, slenderer, and
more obviously tapered than the others which are terete, slightly tapered near the
blunt tip, sometimes slightly compressed. In nicely killed specimens the spines
appear to form three longiseries at margin of furrow since the slenderer furrow spine
of diplacanthid plates is advanced into the furrow; its companion spine forms the
outer series, the middle being composed of the single spines of alternate monacanthid
plates. Adoral carina composed of three or four contiguous postoral adambulacrals,
the single spines of which are appreciably longer than those following.

Mouth plates with a conspicuous apical, tapered, blunt or subacute spine (with
a small lateral companion) longer than the median suture, and a long slender tapered
suboral spine almost identical with the first few adambulacral spines, though a
trifle longer.

Crossed pedicellariae (pl. 29, figs. 1, la—le) are very numerous, 0.29 to 0.81 mm.
long, in conspicuous wreaths (one to four tiers of pedicellariae broad) surrounding
abactinal and superomarginal spines, and scattered singly or in groups of two or
three among the papulae (type); in fairly large clusters on the outer side of infero-
marginal and actinal spines (along with straight pedicellariae in largest examples);
mixed with compressed broadly lanceolate straight ones, in clusters, on the outer
side of adambulacral spines. The last differ in form from the abactinal, being slightly
smaller (0.27 mm.) with a larger terminal portion to jaw, as seen in profile. (Pl. 29,
figs. la, 1d.)

Straight pedicellariae numerous, large. (Pl. 29, figs. 4, 4a-4f.) The abactinal
vary greatly in size and form, some having broad squarish dentate jaws, others
narrower dentate jaws with two or three teeth; still others are small, compressed,
ovate, edentate, as in coei. The intermarginal and axillary channels are conspicuously
provided with broad-valved toothed pedicellariae (upward of 1.25 mm. broad and
1.5 mm. high), veritable giants compared with the same armature of coei. Similar
but somewhat smaller ones are found among the actinal spines, where also there are
still smaller, compressed, ovate or broad-lanceolate pedicellariae without obvious
teeth, together with intermediate stages between the two. Some of the latter migrate
upon the inferomarginal and actinal spines in the largest example (Captains Bay).
The largest pedicellariae are much broader than the spines in small and medium-sized
examples and vary somewhat in number, but are always numerous. Ovate com-
pressed pedicellariae occur abundantly on the adambulacral and oral spines, and very
small ones singly or in clusters on the furrow face of the plates. The difference in
size between these and the intermarginals is enormous.

Madreporic body conspicuous with fairly coarse striae, situated at nearly mid r.

Tube feet fairly large, crowded in four series. Actinostome small, rather sunken.

Young.—A specimen with R 28 mm., was taken with adults at station 4777,
and resembles the larger examples, but has only a few actinal spines at base of ray.
The characteristic large pedicellariae are well developed. The next size with R 67
mm. is adult, in respect to spines and pedicellariae.

Anatomical notes—The gonads (not mature in any example examined except
the large one from Captains Bay) are attached to the interbrachial septum on a level
with the inferomarginal plates, just a few millimeters adcentral to the interbrachial

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—-FISHER 89

angle. The duct passes between the dorsal lobes of two inferomarginal plates, turns
distad and slightly downwards, to open in the axillary channel on a level with the
inferomarginal plates, or slightly below. There are two pulpy prominences close
to the innermost (reduced) inferomarginal spine and usually guarded by large straight
pedicellariae, which mark the opening. They are situated on the actinal surface, as
viewed from below, but close to the margin. (Pl. 26, g.)

The interbrachial septum is firm and thick (calcified) and extends about two-
thirds of r toward the center of disk.

Type.—Cat. No. E 1492, U.S.N.M.

Type locality —Ridge, Captains Bay, Unalaska, 1880. W.H. Dall, No. 1749
(1 specimen).

Distribution —From Unalaska to Semisopochnoi Island, Aleutian Islands.

Specimens exramined.—In addition to type, 17 from: Station 4777, Petrel Bank,
Bering Sea, northeast of Semisopochnoi Island, Aleutians, latitude 52° 11’ N., longi-
tude 179° 49’ E., 52 to 43 fathoms, fine gravel, three specimens.

Station 4779, same locality, 54 to 56 fathoms, broken shells, pebbles, sand, 14
specimens.

CAMTSCHATICA SECTION

The species of this section are, so far as known, restricted to the coasts bordering
the north Pacific and southern Bering Sea. On the Asiatic side they have not been
recorded south of the Kuril Islands and on the American coast south of Santa Cata-
lina Island, Calif.

All the forms are very variable. Species lines are extremely indistinct, some-
times hardly recognizable. For this reason descriptive terms have to be handled in
a relative manner, and figures are indispensable. Most of the species are a composite
of small varieties or formae. Some of these forms perhaps represent incipient species.
Most of the species have paucispinous and multispinous phases, as well as coarse-
spined and slender-spined formae; rays slender, medium, or robust. Various com-
binations of these extremes and their seemingly endless intermediate stages occur.
Even with a large series it is not possible to satisfy oneself of the systematic position
of every specimen. It is not expected therefore that this treatise will suffice to
allocate every isolated specimen of which the name is required. Nor do I admit that
a few apparent exceptions will invalidate the whole systematic arrangement. The
careful investigator will find many of his perplexities anticipated in the following
pages. I have found this group more difficult even than the polaris section.

The following is a list of the names which have been applied to or intended for
species of this group, in order of their priority.

Asterias camtschatica Brandt, Prodromus, 1835, page 270. Kamchatka. This
appears clearly to apply to the common 6-rayed Kamchatkan Leptasterias. Leptas-
terias camtschatica.

Asterias hexactis Stimpson, 1862, page 272. Puget Sound. Types in the
United States National Museum; studied. Leptasterias heractis forma heractis.

Asterias aequalis Stimpson, 1862, page 273. Monterey, Calif. Type in United
States National Museum; studied. Leptasterias aequalis.

Asterias vancouveri Perrier, 1875, page 64. Puget Sound. Type in British
Museum; studied. L. hexactis vancouveri.

90 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Asterias macropora Verrill, 1909a, page 65, Figure 10. Alaska (see below).
Name has status only by bare technicality. Stenastervas macropora.

Asterias epichlora Verrill (not Brandt), 19096, pages 544, 549. See below.

Asterias epichlora variety alaskensis Verrill, 1909), page 549. No type locality;
no type. See below.

Asterias multiclava Verrill, 1914, page 114. Bering Island, Commander Islands.
Type in United States National Museum; topotypes studied. Synonym of L.
camtschatica.

Leptasterias aequalis variety compacta Verrill, 1914, page 130. Pacific Grove,
Monterey Bay, Calif. Type in Yale Museum. Synonym of L. aequalis forma
aequalis.

Leptasterias aequalis variety nana Verrill, 1914, page 132. No type or type
locality. ZL. aequalis forma nana.

Leptasterias aequalis variety concinna Verrill, 1914, page 132. Monterey Bay,
Calif. Type in Yale Museum; studied. Synonym of L. aequalis forma aequalis.

Leptasterias epichlora Verrill (not Brandt), 1914, page 132. Sitka. Verrill’s
epichlora covers part of L. hexactis and L. alaskensis. See below.

Leptasterias epichlora alaskensis Verrill, 1914, page 136. Dutch Harbor,
Unalaska. (Pl. 85.) Type missing from Yale Museum, 1927. Name restricted to
L. alaskensis; used by Verrill for alaskensis, multispina, and hexactis, part.

Leptasterias epichlora alaskensis variety carinella Verrill, 1914, page 137. Sitka
and Dutch Harbor; Dutch Harbor herewith designated as type locality. No speci-
mens in Yale Museum, 1927. Believed to be young of L. alaskensis.

Leptasterias epichlora alaskensis variety siderea Verrill, 1914, page 137. Several
localities, Sitka to Dutch Harbor. Yakutat herewith designated as type locality.
Type figured, but neither it nor any other example in Yale Museum, 1927. Lept-
asterias hexactis forma siderea.

Leptasterias epichlora miliaris Verrill, 1914, page 138. Cape Fox, Sitka, Alaska;
Queen Charlotte Islands. No figure; no specimens in Yale Museum, 1927. A young
form, said to resemble L. aequalis; possibly the young of L. hexactis forma plena.

Leptasterias epichlora miliaris variety regularis Verrill, 1914, page 139. Cape
Fox, southern Alaska. Authentic specimen seen at Yale Museum. L. hexactis forma
regularis.

Leptasterias epichlora miliaris variety subregularis Verrill, 1914, page 139. Sitka,
Alaska. Not figured; no specimen in Yale Museum, 1927. Possibly the same as
regularis; or a 6-rayed Evasterias troschelii.

Leptasterias epichlora miliaris variety subnodulosa Verrill, 1914, page 139.
Wrangel, Alaska. No specimen in Yale Museum, 1927. This is almost certainly
young vasterias troschelit.

Leptasterias epichlora plena Verrill, 1914, page 140. Vancouver Island. Type
in Victoria Memorial Museum, Ottawa; studied. L. heaxactis forma plena.

Leptasterias epichlora pugetana Verrill, 1914, page 142. Puget Sound. Type
from University of California. This is a variation of L. hexactis forma regularis.
Leptasterias (?) dispar Verrill, 1914, page 142. Dutch Harbor, Unalaska. Type
in Yale Museum; studied. Leptasterias camtschatica dispar. ;

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 91

Stenasterias macropora Verrill (Asterias macropora 1909), 1914, page 145. Sitka;
Queen Charlotte Islands. Stated, 1909, to be 5-rayed, and 1914, 6-rayed; R15 mm.
No specimens in Yale Museum, 1927. Probably does not belong in this section.

It is extremely unfortunate that Verrill resuscitated Brandt's Asterias epichlora
for any 6-rayed sea star. Brandt distinctly says that his species is 5 rayed, and to
assume that he had a 5-rayed variant of a normally 6-rayed species is to stretch
probability too far. After examining literally hundreds of specimens of L. alaskensis,
L. camtschatica, L. heractis, and L. aequalis and their various formae, I have found
exactly one 5-rayed L. heractis forma aspera (Kodiak).

It is probable that Brandt had a young Lvasterias troschelii forma alveolata.
Verrill himself described small examples of this form as variety subnodulosa under
Leptasterias epichlora. Many years ago Doctor Stimpson restricted epichlora to
what I have called Evasterias troschelii {. alveolata. 1 think he was right. He knew
that young alveolata is sometimes green.

All of the examples of Verrill’s “epichlora” which I was able to find at the
Yale Museum in 1921 and 1925 (Sitka, Wrangell, Cape Fox) are referable to L. her-
actis forma regularis and aspera. One specimen of L. alaskensis labeled Sitka,
is certainly n¥t from there, but probably from west of Kodiak Island. The straight
pedicellariae are unmistakable.

In view of all this uncertainty it is best to discard the name epichlora. If it
were referable to a 6-rayed Leptasterias, the probabilities would be in favor of L. her-

actis forma regularis.
LEPTASTERIAS CAMTSCHATICA (Brandt)

Plate 42, Figures 1, la-ld; Plate 49, Figure 1

Asterias camischatica Branpt, Prodromus, 1835, p. 270 (Kamchatka).—Stapen, Challenger
Asteroidea, 1889, p. 820.

Asteracanthium camtschaticum Branpt, in Middendorff’s Reise, vol. 2, 1851, p. 32.

? Asteracanthion camtschaticus Groupe, Nova Acta Acad. Caes. Leop. Carol. Nat. Cur.,
vol, 27, 1857, p. 23.

Asterias multiclava Verritt, Shallow-water Starfishes, 1914, p. 114, pl. 58, fig. 2; pl. 59,
fig. 1; pl. 69, fig. 1; pl. 84, figs. 2, 2a.

Leptasterias camtschatica Fisner, Ann. Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 599.

Diagnosis.—Rays six, of medium width and length, tapered rather evenly.
R 62 mm., r 14 mm., r=4.4 r; breadth of ray at base, about 15 mm. A small species
differing from L. alaskensis Verrill in the presence of a nearly complete series of
actinal plates and spines, together with a short second series at base of ray; in the
more regular arrangement of the spines and papulae of the lateral parts of the abac-
tinal area; in the size and form of the straight pedicellariae, which are small, com-
pressed, and lanceolate or ovate, not large with broad obtuse denticulate jaws.

Description —Abactinal spines short, fairly robust and of nearly uniform length
(0.45 to 0.55 mm.), with denticulate capitate tips, the denticles arranged more or
less in definite rows, or without serial arrangement. Spines typically in rather poorly
defined longiseries, except for those bordering a conspicuous supramarginal channel
which usually form a very regular longiseries. The carinal plates each carry several
spines so that the midradial line is occupied by a series of small crowded groups
of spines, somewhat larger than the others. Between these and the regular lateral

92 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

series there is sometimes one zigzag series, or two, or the space may be filled with
spines not regularly arranged. Those of disk form small groups, or stand singly,
but are not typically acervate.

The papular areas are in more or less distinguishable longiseries abactinally.
Those occupying the supramarginal channel form a fairly straight row. ‘Two others
between this series and the carinal plates can be made out in some specimens—
six for the abactinal area (Petropavlovsk, Kuril Islands). In two specimens from
Medni, Commander Islands, the central area of ray is broken up and irregular.
One series of intermarginal areas; proximally two series and distally one of actinal.
Areas small, two to five papulae each.

Superomarginal spines two, capitate, striate, sometimes a trifle larger than the
dorsals, forming a regular duplicate longiseries rather high on side of ray, with the
conspicuous supramarginal channel above and an intermarginal channel below.
A specimen from Petropavlovsk has an arcuate vertical series of three spines on
most of the plates. The marginal series of plates bend upward at the base of the
ray, and the upper series of spines, which border the abactinal surface, passes inward
on the interradial line, adjacent to the same series of the neighboring ray.

The inferomarginal spines are usually one to a plate proximally and two dis-
tally. They are cylindrical, or slightly clavate, round-tipped, and are distinctly
heavier and longer than the superomarginals on the proximal half of the ray, but
only a trifle larger distally, where the lower of the two spines is sometimes longer
than the upper. In this case the upper is subequal to the superomarginals. There
are two series of actinal spines (and two series of plates), the outer extending three-
fourths to nearly the whole length of the ray, while the inner reaches to the middle
in large specimens (R 47 mm.).

The first five to eight adambulacral plates carry a single, slender, rather long,
terete slightly tapered blunt spine, followed by a very variable number (usually
few) which have mostly two spines. Then the plates have one and two spines
in fairly regular alternation; or less often mostly one spine with diplacanthid plates
irregularly scattered. Beyond the first half dozen adoral plates the spines become
shorter and thicker, often clavate, with the exception of the alternate furrow spine
of diplacanthid plates, which is generally decidedly slenderer than the others. It
is, however, frequently of the same clavate, or club-shaped, form as the others.
The first three pairs of adoral adambulacral plates are in contact interradially;
the first two plates are longer than the succeeding plates by about 30 to 50 per
cent.

Mouth plates very similar to those of alaskensis and typically there are three
spines, two at the inner apex and one at the actinal surface. Sometimes the shorter
of the apical spines, which corresponds to the furrow spine of diplacanthid adambul-
acral plates, is lacking. The longer, that adjacent to median suture, is about as
long as this suture, or a little shorter. The two spines of a pair are frequently dis-
similar in length. The suboral is slender, tapering, and upwards of twice the length
of the apical spine.

Straight pedicellariae (pl. 42 fig. 1c) inconspicuous and few abactinally, outside the
supramarginal channels. They are broadly lanceolate to ovate, sharp to blunt, and
measure 0.36-0.46 mm. They are very inconsiderable in size as compared to the

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 93

spines and are so much smaller and different in form from the corresponding ones of
alaskensis that they furnish a convenient character for identification. There is the
usual cordon of small pedicellariae along the furrow face of the adambulacrals, as
well as on the oral and first six or eight adambulacral spines.

Crossed pedicellariae (pl. 42 fig. 1b) are small, blunt, with a rather extensive
terminal lip. They measure about 0.2 to 0.26 mm. in length abactinally, where
they form a sparse series around the spinelets or occur only here and there on the
integument between (Medni). They form wreaths on the superomarginals and
clusters on the outer (upper) side of inferomarginals, actinal and adambulacral
spines.

Madreporic body circular, rather small, situated about mid r, and surrounded
by a circle of spines.

Color in life: ‘‘Above, deep olive green with the tips of the spines whitish; those
of the edges very pink.”” (L. Stejneger, note on specimen from Bering Island).

Variations —The specimens from the Commander Islands have somewhat less
regularly arranged abactinal spines than have the Kamchatkan examples. The
latter have somewhat broader, more evenly tapered rays, and on either side two
pretty regular dorsolateral series of spines.

The three specimens from Simushir, Kuril Islands, are small (R 7, 21, and 25
mm.) yet the actinal plates and spines are well developed in the two larger. These
and the small straight pedicellariae will serve to differentiate the species from L.
aaskensis asiatica, with which it is associated. The abactinal papulae are in six
longiseries, but the intervening spines are not regular. Many of the dorsolateral
plates carry two or three spines, while in Petropavlovsk specimens there is more
often only one spine, although plates with two or three do occur here and there.
The superomarginal, inferomarginal, and actinal spines of the Simushir example are
relatively longer and a little slenderer than in either Bering Island or Petropavlovsk
specimens, and there is but one apical oral spine. Straight pedicellariae are fairly
numerous, compressed, lanceolate ovate, obtuse to fairly acute.

Type locality —Kamchatka. .

Distribution —Commander Islands, Kamtschatka, and the Kuril Islands.

Specimens examined.—Fourteen.

Bering Island, Commander Islands, L. Stejneger, 1883; one specimen. Nikolski,
Bering Island, Commander Islands, shore rocks, June 15, 1906, Albatross, two speci-
mens. Medni Island, Commander Islands, shore rocks, June 13, 1906, Albatross,
three specimens. Petropavlovsk, Kamchatka, harbor, August 13, 1896, June 18,
1906, Albatross, five specimens. Milne Bay, Simushir, Kuril Islands, shore, June
23, 1906, Albatross, three specimens.

Remarks.—Brandt’s diagnosis reads: ‘‘Disci diameter circiter pollicaris. Radii
sex conici acuminati, longitudine vix subequales, pollicis unius vel 14 longitudine,
anti basin multo latiores. Totum dorsum umbrinum vel ochraceum papillisque
pedicellatis, truncato-capitatis, frequentibus, albis vel e fuscente albidis, eminentiis
reticulatis impositis, in radiis series subregulares formantibus obsessum. In mari
Camtschatico.” On page 32 of the Reise, he says: ‘‘ Unterscheidet sich durch das
Vorkommen von 6 armen * * * und auf dem Ricken 6 bis 7 Reihen geképfter
Stacheln tragen, so wie durch dreireihige diinne Ambulacral stacheln.”

63160—30——-7

94 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

It is apparent that Brandt was describing a small Kamchatkan sea star with
six tapered pointed rays in which the small, numerous, truncate capitate spines were
arranged in six or seven sub-regular longiseries. This diagnosis agrees with the
species under discussion. On some specimens the adambulacral spines appear as
if three ranked owing to the posture of the spines. The furrow spine of diplacan-
thid plates is directed obliquely over furrow, the other obliquely toward the actinal
spines. These form the inner and outer series, while the middle series is formed by
the upright spine of the alternate monacanthid plates.

Verrill in commenting upon Brandt’s description, which he considers indefinite
and vague, renders ‘‘series subregulares,” ‘‘subangular series.” This error obscures
Brandt’s meaning. It seems to me that the description is definite enough. Moreover
there is no Kamchatkan species, other than the one under discussion, which has six
rays and the abactinal spinelets in somewhat regular longiseries. From Brandt’s
comments in Middendorff’s Reise (1851, p. 32) he apparently had specimens which
had been dried without much prior preservation. Grube in fact says (1857, p. 24)
that his description seemed to have been made from only dry material.

LEPTASTERIAS CAMTSCHATICA DISPAR (Verrill)
Plate 42, Figures 2, 3, 4; Plate 49, Figures 2-6; Plate 50, Figure 1

Leptasterias (?) dispar Verritu, Shallow-water Starfishes, 1914, p. 142, pl. 16, fig. 7, type;
pl. 106, fig. 3 (as acervata).

This polymorphic race is the representative of Leptasterias camtschatica on the
Aleutian chain, whence it extends as far as Kodiak Island. It is found in Bering
Sea at the Pribilof Islands. The name of the race is derived from one of its formae,
originally described as a distinct species.

Three phases or formae have been distinguished—nitida, nesiotis, and dispar, of
which the first is the equivalent of the common form of L. camtschatica camtschatica.
In the Asiatic race only one form has been observed.

These three phases are morphologically equivalent to formae found in widely
dissimilar species. For example they correspond, in the order named, to Pisaster
ochraceus {. ochraceus, {. confertas, {. nodiferus; or to Evasterias troschelii f. alveolata,
f. acanthostoma, f. troschelii.

The small type variety of f. dispar does not resemble f. nitida (which is the
equivalent of L. camtschatica camtschatica). Therefore only forma nitida is useful
for direct comparison with true camtschatica, since no equivalent of dispar is known
for the latter species.

However distinct dispar may appear to be when only extreme forms are compared,
there exists a pretty complete series of intermediate specimens connecting it with
f. nitida. There is available the alternative of regarding dispar as a distinct species
which has freely crossed with nitida. Dispar has been taken on the outer part of
the Aleutian chain where nitida has not yet been found (but where it may never-
theless occur). There seems to be no necessity of invoking hybridism, for quite as

dissimilar formae are known in other species of the genus, for example, Leptasterias
polaris acervata.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 95

If these formae breed fairly true when isolated I presume it is simply a question
of terminology whether they be called formae or species. The fact remains, however,
that intermediate specimens occur at localities where only extreme examples of dispar
(but not nitida) have been found (Attu), and also where neither nitida nor extreme
dispar have been collected (Atka and Shumagin Islands).

In the following description the formae have been listed in the order of their
resemblance to L. camtschatica. It is a taxonomic episode that the race must have
a name which connotes a form morphologically so dissimilar to camtschatica.

Forma NITIDA, new forma
Plate 42, Figures 3, 3a-3c; Plate 49, Figure 2; Plate 50, Figure 1

This corresponds to the typical form of camtschatica. The abactinal spinelets
are much more numerous and do not form evident longiseries with the exception of
the carinals which are larger than the dorsolaterals, more crowded, and occur in
groups, close together, so that a carinal band of more crowded spinelets can be
differentiated. The marginal plates are rather high on side of ray. Just above them
is a typically narrow, regular channel—the supramarginal—containing a straight
series of papulae. Between this channel and the edge of the carinal band (4 or 5 mm.
in fully grown specimens) the spinelets are thickly scattered, with a tendency to form
short transverse lines, especially adjacent to the superomarginals. A few of these
lines extend entirely across the dorsolateral area and seem in some cases to be con-
tinued by the superomarginal combs.

Superomarginal spines three to five, usually arranged in an arcuate vertical
series of three or four proximally and three or two distally, with occasionally another
smaller spine on the adoral lobe of the plate. The spines are capitate slightly larger
or subequal to the dorsals, and form a regular series of combs rather high on the side
of the ray.

Intermarginal channel narrow. Inferomarginal spines two (usually one on first
four or five plates), much more robust and about twice as long as the superomarginals.
The end is usually swollen and turned upward slightly. Two series of actinal plates,
one reaching 0.8 the length of R, the inner about 0.5 R in fully grown specimens.
Spines, one to a plate, similar to the inferomarginal spines. The inner series of actinals
sometimes extends only a slight distance upon the ray.

Adambulacral spines similar to those of camtschatica. The first four to eight
plates have one spine; then a few may have two, followed by two and one in not
very regular alternation; then on outer part of ray the plates are monacanthid by
the suppression of the outer spine of diplacanthid plates. This results in alternate
plates having only a furrow spine, which is slender and more advanced into the
furrow than the clavate, subtruncate subambulacral spine of the intervening, original
monacanthid plates. Three pairs of adoral adambulacral plates, as in camtschatica,
are in contact interradially back of oral plates.

Mouth plates similar to those of camtschatica. In this forma the outer or smaller
of the two actinostomial or apical spines is missing.

Papular areas small, with few papulae. The abactinal do not form regular
longiseries except adjacent to superomarginal plates.

96 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Very small lanceolate pedicellariae are found on the abactinal surface. Ovate
to lanceolate acute, larger ones, as large as 0.8 mm. long, are found rather plentifully
in the marginal and actinal channels, on the proximal adambulacral spines, oral
spines, and along the furrow face of the adambulacrals.

Crossed pedicellariae, variable in numbers, are distributed as in camtschatica.
They are frequently abundant inferomarginally and actinally. The abactinal
measure 0.24 to 0.27 mm.

Madreporic plate inconspicuous, partly overhung by the numerous encircling
spinelets.

Anatomical notes.—There is nothing peculiar or unusual about the skeleton of
this form. The carinal plates are conspicuous when denuded and are imbricated
directly to form an irregular series. They appear to have been originally four
lobed; in adult specimens the contour is irregular. The marginal and actinal plates
have a warped 4-lobed contour, the upper or outer lobe being more advanced toward
end of ray (transverse axis of plate, oblique). The outer row or actinals nearly
reaches the end of ray while the inner attains about the middle, measured on side.
The dorsolaterals, smaller than either marginals or carinals, are irregular in contour
and without regularity in arrangement. The intervals of the skeleton are smaller
than plates on basal half of ray; slightly larger on distal half.

Variations —The dorsolateral spinelets become a little more widely spaced while
the carinal are aggregated into slight convex differentiated groups. These groups
are not as prominent as in typical dispar. LL. dispar may be achieved by the reduction
in the number of the dorsolateral and superomarginal spinelets. These incipiently
“acervate” examples are found at Kodiak Island (Karluk) (pl. 50, fig. 1), Unalaska,
Pribilof Islands (St. Paul and St. George). A number of specimens from St. George
(pl. 49, fig. 4) exhibit this variation in a marked degree and I have noted them under
dispar although they are really perhaps as close to nitida. ‘The spines of the convex
carinal groups are closely aggregated. At the base of the ray there are sometimes
one or two adradial clusters. Superomarginals, two or three.

As mentioned above this incipiently clustered condition of the carinal spinelets
is shown by a few specimens from Karluk, Kodiak, which resemble and may perhaps
be intermediate with L. hexactis f. siderea, which occurs with them, at low tide.

Type of forma nitida.—Cat. No. £1493, U.S.N.M.

Type locality—St. Paul, Pribilof Islands, Bering Sea, rocks at low tide. Col-
lected by William Palmer.

Distribution.—Pribilof Islands, Unalaska, and Kodiak (Karluk).

Forma NESIOTIS, new forma

Plate 42, Figures 2, 2a; Plate 49, Figure 3

This differs from forma nitida in having much more delicate spinelets. Typi-
cally the carinal spinelets are not any larger or only slightly larger than the other
abactinal spinelets and the mouth plates generally have two spines each at the acti-
nostomial end.

The abactinal skeleton is sometimes more open than in nitida, the papular areas
forming longiseries of which one above the superomarginal plates and one on either
side of the carinals is evident, while one between these two is likely to be broken up.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 97

Potentially there are six longiseries on the abactinal area. In some variations the
spinelets are so arranged that a reticulate or areolated pattern is formed. Such
specimens may be readily distinguished from forms of L. alaskensis by the presence
of a long series of actinal spines and by the small lanceolate straight pedicellariae of
the sides of ray.

The number of superomarginal spines is usually three or four in an arcuate ver-
tical series. Owing to their smaller size they appear to be more spaced than in
nitida. The inferomarginal and actinal spines are much slenderer than in nitida
but number the same.

The lanceolate straight pedicellariae are appreciably smaller than those of
nitida, measuring 0.3 to 0.45 mm. (PI. 49, fig. 2a.) The abactinal crossed pedi-
cellariae measure 0.16 to 0.21 mm.

This form predominates at St. Paul, Pribilof Islands, where it far outnumbers
nitida judging by the series collected by Dr. G. D. Hanna. The type was taken at
Village Point, July 25, 1920, among bowlders, between tides, along with L. alaskensis.
This forma has not been found elsewhere.

Color in life—Dr. G. Dallas Hanna collected at St. Paul, Pribilof Islands, 20
specimens of forma nesiotis and intergrades and made notes of the color variations.
The following notes refer to 11 specimens taken from an area 3 feet square, among
bowlders, near the village:

1. Dorsal surface olive spotted with dark brown; ventral surface light pink.

. Lighter olive, with more reddish spots.

. Spots very dark brown, almost black, the interspaces yellow.

. Dorsal spots black, interspaces pure yellow.

. Under side with yellowish tinge on the pink.

. Dorsal surface clearer green, the spots very dark brown; ventral surface,
darker pink.

The following specimens were taken July 25, 1920, at Village Point, between
tides:

1. Upper surface with large black blotch or blotches on disk of very irregular
form, but in every case filling center of disk and radiating upon each arm forming
thus a stellate area; remainder yellow, pink on either side of the groove.

a. Blotches very dark blood red, remainder a dirty gray; margins of furrow
light pink.

b. Blotches dark lavender, remainder gray; margins of furrow, light pink.
(Type of forma nesiotis.)

Type of forma nesiotis—Cat. No. E 1494, U.S.N.M.

Type locality.—St. Paul, Pribilof Islands, Bering Sea, August, 1920, collected
by G. Dallas Hanna.

was a

Forma DISPAR Verrill
Plate 42, Figure 4, 4a, 4b; Plate 49, Figures 4, 5, 6

Leptasterias ? dispar Vernrtui, 1914, pl. 16, fig. 7; pl. 106, fig. 3 (as Asterias acervata).

A 6-rayed form differing from forma nitida in having the carinal spines much
enlarged and usually grouped in nodular clusters; typically few dorsolateral spines
much small +r than the carinals; a conspicuous supramarginal channel, bounded above

98 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

in large specimens by a very regular series of small dorsolateral spines and below by
one or two series of robust superomarginal spines. R 66 mm., r 17 mm. R=3.88
r; breadth of ray at base 18 to 20 mm. (Attu Island).

Verrill’s type and only listed specimen of this variable form is a small example
from Unalaska. Under the caption of Asterias acervata he gives a good figure (pl.
106, fig. 3) of a variety from Nazan Bay, Atka. I have two specimens of the same
lot, collected by W. H. Dall, 1873.

The type is an extreme variant of the form and is farthest removed in appear-
ance from nitida. I have three small examples of this subforma from Unalaska
(type locality, pl. 49, fig. 6), two small and one large specimen from Attu (pl. 49,
fig. 5) at the end of the Aleutian chain. Of the intermediate stages between this
extreme and acervate examples of nitida there is a pretty complete series. Scarcely
two specimens are alike.

The large example from Attu (R 66 mm.) has the carinal line armed with very
prominent clusters of large capitate spines, three to five in a group, forming a zigzag
longiseries; 10 or 11 similar groups mark the primary plates of disk. These spines
are larger than any of the others, stand well.above the general level, and have
convex, finely striated tops, upward of 1.75 mm. in diameter. On the inner side of
the broad supramarginal channel which is abactinal in position is a line of relatively
small capitate spines, fairly well spaced and not reaching end of ray. Scattered
between these and the carinal groups are a relatively few similar spines which some-
times form a poorly defined series for a short distance. The spinulation between
the superomarginal and carinal series is characteristically meager, but this feature is
very variable in different specimens from the same locality. In the extreme as rep-
resented by the small type there may be only a single very interrupted series of small
dorsolateral spinelets, and a single irregular series of spaced subtruncate carinal
tubercles.

The superomarginal spines are either one or two to a plate, usually one in the
small type form, but two in large specimens, where they stand close together on the
plate and form a prominent duplicate series defining the dorsolateral margin of the
ray, which is more definite and prominent than in nitida. Spines cylindrical, slightly
capitate, striate and subtruncate, about 1.25 mm. long, and a little bigger than the
dorsolateral spines. The intermarginal channel, though variable, is broad proximally
in both large and small examples.

Inferomarginal spines usually one on the first three to six plates, then two, with
an occasional monacanthid plate; less often a plate carries three spines. A series of
similar actinal spines reaches about 0.8 R along ray, while a very short second series
is also present. These spines are cylindrical, round-tipped, and longer and thicker
than the superomarginals (about 2.5 mm.). In typical small specimens the spines
are more prominent and likely to be compressed; they form a bristling ventrolateral
border, and the actinal series is short.

The first four to six adambulacral plates are monacanthid; then follow about
20 diplacanthid plates; then a variable number alternate with one and two spines.
About the last fourth or fifth of the ray has monacanthid plates (a plate bearing a
furrow spine alternating with one carrying a subambulacral). The spines are simi-
lar in form to those of nitida. Toward the mouth plates they gradually lengthen

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 99

and become slenderer. In the median part of the ray the subambulacral spines are
cylindrical, or slightly clavate, round-tipped. The furrow spines are slenderer and
tapered. There are three pairs of contiguous plates back of the mouth plates.

Mouth plates very similar to those of camtschatica. There are two actinostomial
spines and one suboral. The smaller of the two apical spines is bent across the begin-
ning of the radial nerve. The other, about twice as long (2 mm.), is a trifle longer
than the median oral suture.

The series of actinal, intermarginal, and supramarginal papular areas are regular,
but the other dorsolateral areas are not at all regularly arranged. They vary greatly
in number according to the age of the specimen and appear not to be of diagnostic
value. There are only a few papulae to each group. In small examples they are
mostly single.

Straight pedicellariae are all small and are of the lanceolate and ovate com-
pressed type, quite inconsiderable in bulk as compared to the spines. The dorsal
are commonest interradially and along the supramarginal channel, and are lanceolate-
ovate, or ovate, blunt, or subacute; the intermarginal and actinal are similar but a
little larger. They occur in clusters on the oral spines and among the crossed pedi-
cellariae of the furrow spines.

Crossed pedicellariae, distributed as in nitida, are fairly numerous in large speci-
mens, but few, especially on the abactinal surface, in small ones. Their size varies
with that of the specimen. On large examples they measure 0.25 to 0.27 mm. (PI.
42, figs. 4a, 4b.)

Madreporic body circular, situated at about the middle of r, surrounded by a
complete or an incomplete ring of short capitate spines. Striae rather coarse, radiat-
ing from center.

In small specimens, as Verrill states, the ambulacral groove is broad. This
feature may be masked by the arching of the abactinal surface, so that the edges of
the furrow are drawn together.

Anatomical notes—In small specimens the carinal and superomarginal plates
are disproportionately large as compared to the irregular dorsolaterals. The triserial
arrangement of the dorsal plates, mentioned by Verrill, is lost in large examples by
the interpolation of plates between the carinals and the primary dorsolateral series
(that which borders the supramarginal channel). In typical large examples such as
that from Attu, the carinals are, nevertheless, still relatively large. (Pl. 49, fig. 5.)

The gonads open ventrally. The apertures are close to the interradial line, and
in line with the outer actinal row of papulae. Each is at the summit of a low promi-
nent papilla, and the specimen (R 57 mm.) taken at Attu, June 10-11, 1906, has
large but subspherical, nodular ovaries filled with ripe eggs. Another large specimen,
that described, taken May 29, 1892, at Attu, is a male and the gonads are of the
branched type common to the males of this genus. The gonads appear to be not
fully mature. A mature female, taken June 15, 1920, at Unalaska, has R only 26 mm.

Variations.—The small typical subforma as described by Verrill is unmistakable,
but this is an extreme having a minimum of abactinal spines and usually one series
of prominent superomarginal spines. The carinals are few and unusually large, as
if there were only a certain amount of lime to be expended—the larger the spines
the fewer their number. In larger specimens the number of carinal spines varies

100 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

from one to a plate to a crowded group of five or six, and the size, from slightly to
conspicuously larger than the dorsolaterals. The dorsolateral spinelets vary from
one incomplete longiseries to the equivalent of three, such specimens being in fact
nothing more than examples of nitida with grouped carinal and spaced dorsolateral
spinelets. 4

Shumagin Islands: Largest specimen, R 47 mm., has one or two superomarginal
spines, rather numerous dorsolaterals, and not very prominent acervate carinals.
The five specimens from the Shumagins suggest young acervata, but are not.

Chernofiski Bay, Unalaska: A specimen (47 R mm.) has rather narrow, arched
rays and proximally three or four superomarginal spines, in an arcuate vertical comb.

Unalaska, Alaska: In addition to three small typical examples there are nine
intermediates some of which may be called acervate examples of nitida. The spines
of the conical groups are not neat and compact as in the St. George Island variety.
Superomarginal spines usually two, sometimes one.

St. George Island, Pribilof Islands: These belong really to a subforma of nitida
rather than to forma dispar. Along the carinal ridge (the ray being well arched) is
a zigzag series of conspicuous, convex, compact groups of 8 to 12 unequal, but en-
larged, close-set spines; dorsolateral spines numerous, superomarginals two or three.
(See under f. nitida, pl. 49, fig. 4.)

Adakh Island, Andreanof Group, Aleutian Islands: A specimen resembles the
St. George examples but the groups are smaller; the superomarginals two; R 73 mm.
There are rather numerous coarse ovate intermarginal straight pedicellariae but they
are not of the form nor nearly as large as those of alaskensis. This large specimen may
be classified as an acervate specimen of forma nitida, or as dispar with numerous
dorsolateral spinelets. One specimen with much fewer dorsolateral spinelets.

Nazan Bay, Atka: The specimen figured by Verrill (1914, pl. 106, fig. 3) is much
smaller than the two recorded here (R 70 and R65 mm.). In these the carinal spines
are prominent and the dorsolaterals well spaced but more numerous than in the
example from Attu described above. Superomarginals two. Intermarginal straight
pedicellariae smaller than in the Adakh example.

Attu Island: A small example with two superomarginal spines connects the
extreme type form, from Chickagoff Harbor, Attu, with what might be called the
intermediate variety, of which there is one specimen (in addition to the large extreme
form described) collected at the same locality.

Remarks.—The following differences may be observed between approximately
equal-sized specimens of f. dispar and Leptasterias polaris acervata f. acervata; examples
of acervata with R 50 to 70 mm.; of dispar with R 66 mm. Acervata has a smaller
disk, and the rays are slenderer, especially at the base (R=4.5 to 5 r). The dorsolat-
eral spines are much more numerous, and a certain number of them are enlarged (as
are the carinals) and form spaced, scattered clumps which are strongly convex. The
supramarginal and intermarginal channels are narrow, the former being not at all
conspicuous, as in dispar. In typical dispar the superomarginal plates are more
prominent, the spines being a little longer. Small specimens of acervata (R 22 to 27
mm.) do not resemble equal-sized typical dispar but they do resemble some of the

intermediate stages of dispar from Unalaska. The latter lack the dorsolateral
convex groups of acervate spines.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 101

Type of forma dispar.—In Yale Museum.

Type locality —Dutch Harbor, Unalaska (forma dispar).

Distribution of forma dispar.—Attu Island to the Shumagin Islands and southern
Bering Sea (Pribilof Islands). The subspecies dispar extends from Kodiak Island
(forma nitida) to Attu (forma dispar).

Specimens examined.—Kighty.

Specimens of Leptasterias camtschatica dispar examined

| |
- f.
Locality | nitida | nesiotis| dispar | Remarks
—_ ou = — Sti ee ee ee _
Unalaska (Chernofski seven Siento 5 Salt eta Pid [exéhe-3 1) U.S.N.M.
Unalaska_.. a ee ee aia aCe «| eed 12 | 3 typical dispar; intermediates; U.S.N.M.; Stan-
| ford.
Round Island, Coal Harbor, Unga, Shumagin Is- |........|....-... 4 | U.S.N.M.; W. H. Dall; intermediates,
lands.
Popoff Strait, Shumagin Islands... ............-...|-.--..--|...----- 1] Do.
8t. Paul, Pribilof Islands.......................-.-- 8 2» 2)| U.S.N.M.,; Stanford; G. D. Hanna, H. Heath.
St. George, Pribilof Islands......_ Se oho tents) SN renee 4 Stanford; H. Heath,
Adakh Island, Aleutians. a ee See 2| U.S.N.M,; 1 Intermediate.
Atka Island, Aleutians (Nazan. Bay).. ebb. .S0Ssakel aces 2) U.S.N.M.
Attu Island (Chichagoff Harbor) _. Retain benirne iach & U.S.N.M,; W. H. Dall 1873; Albatross, Jane 10,
| 1906.

LEPTASTERIAS ALEUTICA, new species
Plate 43, Figures 1, la—lc; Plate 50, Figures 2, 3

Diagnosis.—Rays six. Differing from L. camtschatica dispar in having shorter,
thicker rays; more numerous, closer set abactinal spinelets, which have the appear-
ance of a rather even, close, granuliform covering; relatively larger superomarginal
plates, which with their groups and combs of four to six spines form a broader band
along the dorsolateral border of ray. Straight pedicellariae small, ovate or ovate-
lanceolate, acute or obtuse. Resembles a small L. aequalis without longiserial ar-
rangement of the abactinal plates. R 28 mm., r 10 mm., R=2.8 r; breadth of ray
at base 10 mm.

Description.—Rays short, robust. Abactinal surface arched, covered with many
small, close-set, uniform spinelets, which give the whole abactinal area the appear-
ance of being covered with granules. In small examples separate plates not easily
distinguishable; in largest specimens, such as type, the small papular areas serve to
mark off groups of spinelets. In the type, the largest plates (the carinals) carry 8
to 15 spinelets, the dorsolaterals 5 or 6. The spinelets are characteristically
thorny 0.3 to 0.4 mm. long on abactinal plates. The irregular carinals sometimes
have considerably more robust spinelets than dorsolaterals. Dorsolateral skeleton
an irregular reticulum with the eqiuvalent of about six longiseries of papular areas
for the whole abactinal area, only those adjacent to superomarginals at all regular.

Superomarginal plates with four to six spinelets (usually three or four in small
specimens) which distally form a vertical comb and proximally more often a group.
Exceptionally there are as many as eight toa group. The spinelets are a little longer
than the abactinals, rather stubby and subclavate, sometimes curved. They form a

63160—30——8

102 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

conspicuous band well up on the side of the ray, and are bounded below by a narrow
but fairly conspicuous intermarginal channel.

The inferomarginal spinelets stand two (sporadically three) to a plate. They
are subcylindrical and are both longer and stouter than the superomarginals. The
proximal spinelets may be nearly twice as long and 75 per cent thicker than supero-
marginals, and rather clavate in form. A series of actinal spinelets, similar to the
inferomarginals but slightly larger, extends for two-thirds the length of the ray. A
few spinelets of a second series may be present at the base of the ray. The infero-
marginal and actinal spinelets together form three ventrolateral series separated from
the broad band of superomarginal spinelets by the intermarginal channel.

Adambulacral plates diplacanthid for half or two-thirds the length of the fur-
row; the distals alternating diplacanthid and monacanthid; or the distal plates may
be irregularly diplacanthid. The furrow spinelet is, as usual, slenderer than the
adambulacral, which is subcylindrical and subclavate, about as long as the actinal
spinelets but considerably slenderer. Both become longer and decidedly slenderer
on the first dozen plates. The first five or six plates are monacanthid, and the first
three pairs are contiguous interradially.

The oral plates have the usual arrangement of spinelets: Either one or two at
the actinostomial end, and one suboral. The longer of the slender apical spinelets
is as long as or a little longer than the median suture of the plates, and longer than
in alaskensis.

Straight pedicellariae: There are a few small ovate or ovate-lanceolate, com-
pressed, bluntly pointed to obtuse pedicellariae in the supramarginal, intermarginal,
and actinal channels, and along the furrow face of the adambulacral plates, the
latter being quite small. They occur in clusters on the oral armature and mixed
with crossed pedicellariae on the furrow armature of the proximal half of the ray.
The lateral and actinal dermal pedicellariae are much slenderer than any of the
spines. The largest intermarginal measure 0.3 mm. (PI. 43, figs, 1b, Ic.)

Abactinal crossed pedicellariae few and scattered, in circles around the spines.
They form rings or part rmgs on the superomarginal spinelets, and not very large
clusters on the outer side of the inferomarginal actinal, and adambulacral spines.
Abactinal pedicellariae measure 0.18 to 0.2mm. (PI. 43, figs. la, 1a’.)

Madreporic body circular, surrounded by one or two: cycles of granuliform
spinelets and situated at about the middle of r, or a trifle nearer the center of disk.

Variations —This species is typically small with short, rather stout, rays and a
close, uniform covering of granuliform spinelets. It is sexually mature at R 14 mm.,
and the majority of the specimens have R less than 20 mm. The spine-counts in
these small examples are less than in the largest specimens, but the spines are more
uniform in size and distribution. Even in very small examples the actinal plates
extend for half the length of the ray.

The superomarginal spinelets in small specimens form an arcuate vertical series
of three or four with an extra spine on the adoral, convex side of the arc. This is
essentially the arrangement in large specimens where extra spines appear on the
adoral side of the are of usually five spinelets.

_ Specimens from Unalaska, and a part of those from Adakh and Kyska are not
typical and approach, sometimes closely, what appear to be young L. camtschatica

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 103

dispar {. nitida. The difference briefly consists of a slight enlargement of the carinal
spinelets and a disparity in the size of the dorsolateral spinelets, whereas in the
typical form the abactinal spinelets are all fairly uniform. The really problematical
specimens are small. It must be remembered that all young Leptasterias are pecu-
liarly difficult. Forms which are easily separable in the adult phase may converge
in the young stages so that exact differentiation is impossible.

Type—Cat. No. E 1495, U.S.N.M.

Type locality —Adakh Island, Aleutian Islands, July 2, 1893.

Specimens examined.—Forty-two from the following of the Aleutian Islands:
Agattu, 2 specimens; Amchitka, 2 specimens; Kyska (Kyska Harbor), 6 specimens
(W. H. Dall, 1873); Adakh, 8 specimens; Atka, 20 specimens; Unalaska, 4 specimens
(variety).

Remarks.—This species, which so far as known is confined to the Aleutian Islands,
reminds one of a small ZL. aequalis having irregular abactinal plates. It is probably
an offshoot of L. camtschatica. Some of the smallest specimens approach L. camt-
schatica dispar f{. nitida. L. aleutica has typically shorter, broader rays and more
numerous and close-set dorsolateral spinelets, which are arranged in groups without
serial alignment.

The resemblance to aequalis is perhaps due to the relationship which both
species bear to camtschatica. Some variants of L. heractes vancouveri closely resemble
L. camtschatica.

L. aleutica can be distinguished from the forms of L. alaskensis by the small,
compressed, ovate or ovate-lanceolate straight pedicellariae, numerous abactinal and
superomarginal spinelets, and long series of actinal spinelets.

LEPTASTERIAS ASTEIRA, new species
Plate 43, Figures 3, 3a-3e; Plate 50, Figures 4, 5

Diagnosis.—Rays six, rather short and robust; abactinal spinelets subcapitate,
generally one to a plate and pretty uniformly spaced, and not obviously areolated;
carinal series not distinguishable from the rest; abactinal papulae one to three, not at
all in series; superomarginal spines two; inferomarginal and actinal forming each a
single series, well separated; adambulacral spinelets, two, and one and two alternating;
ambulacral furrow narrow; straight pedicellariae small, ovate or elliptical, obtuse.
R 26 mm., r 8.5 mm., R=3 r; breadth of ray at base 9 mm.

Description —Abactinal spinelets short, robust, clavate to subcapitate, the
length being once or twice the breadth at tip, which is grooved and thorny. The
spinelets are not very dissimilar in size, are mostly one to a plate, and are spaced
one-half to two or three times their breadth but have the appearance of being pretty
evenly scattered, without the slightest differentiation of a carinal series, a reticulated
pattern of the abactinal spinelets only vaguely indicated in the larger specimens.

There is a fairly broad but variable and not very sharply differentiated supra
marginal channel. Superomarginal spinelets two, sporadically three, slightly larger
than the dorsals, fairly high on the side of ray. Intermarginal channel about as
broad as the supramarginal. Inferomarginal spinelets generally one, occasionally
two distally; actinals one, the series extending two-thirds or three-fourths the
length of the ray; both are about 50 per cent larger than the superomarginals, sub-

104 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

cylindrical to slightly clavate, and sometimes a trifle curved upward. The actinal
channel is a little narrower than the intermarginal.

The first five or six adambulacral plates are monacanthid; then about five or
six are diplacanthid; the remainder alternate more or less regularly with one and
two spinelets. The subambulacral spinelets are clavate, a little slenderer than the
actinals but often no shorter; the furrow spines are a little slenderer than the sub-
ambulacrals but instead of tapering, as is usual with furrow spinelets, are more often
a trifle clavate. One or two pairs of adambulacral plates meet interradially behind
the mouth plates.

Oral spinelets two; a tapering spine at the inner apex of each plate, shorter
than the interradial suture; one suboral spinelet similar to the first few subambu-
lacrals which are slenderer than further along the ray.

Abactinal papulae mostly single and inconspicuous; intermarginal and actinal
(sometimes two series proximally) in twos and threes.

Straight pedicellariae (pl. 43, figs. 3a-3c) small, in side view ovate, or broadly
elliptical, very obtuse, distributed typically for the genus, the largest being the
actinal interradial dermal. Front profile of jaw, usually ovate or broadly lanceolate.
The dorsal are very small, as are those along the furrow margin. An intermarginal
measures 0.27 to 0.33 mm.; an actinal interradial about 0.3 mm.

The sheaths surrounding the abactinal spinelets are rather pulpy and prominent,
often contiguous, but the crossed pedicellariae (0.2 mm.) are few, usually only one
to three to each spine. The marginal are more numerous but not abundant; the
inferomarginal, actinal, and the adambulacral form very small clusters. (Pl. 43,
fig. 3d.)

Madreporic body small, situated one-half r from center.

Anatomical notes —The hepatic coeca are large and reach nearly to the end of
the ray. The intestinal coecum is a rather slender, 2-lobed sac. The gonads open
ventrally by a genital papilla close to the interradial line. The ovaries are sub-
globular, the eggs large and relatively few (specimen with R 21 mm.). ;

The abactinal skeleton consists of an irregular small-meshed reticulum of very
numerous small plates without a differentiated series of carinals. At the nodes are
mostly three or four lobed primary plates (usually monacanthid) connected by one
intermediate, elliptical, often monacanthid, ossicle. The spaces are larger and the
plates slenderer toward the end of the ray. The superomarginal plates are four
lobed, but at the base of the ray the transverse lobes only may be present on many
inferomarginal plates, since the intermarginal and actinal papular areas are very
frequently confluent. A second series of small actinal plates (more numerous than

the outer series) is irregularly developed at the base of ray, but generally is not
spiniferous.

Type.—Cat. No. E 1496, U.S.N.M. ‘

Type locality —Adakh Island, Andreanof Group, Aleutian Islands.

Distribution.—Pribilof Islands, Shumagin Islands, Andreanof Islands.

Specimens examined.—Pribilof Islands (St. George and St. Paul), shore, 4 speci-
mens. Adakh Island, Aleutian Islands, shore, 32 specimens. Kyska Island, Rat
Islands, Aleutians, 9-14 fathoms, 4 small specimens. Sanborn Harbor, Nagai,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 105

Shumagin Islands, 1 specimen. Round Island, Coal Harbor, Unga, Shumagin
Islands, 1 specimen.

Remarks.—This species may be distinguished from alaskensis by the form of the
small straight pedicellariae, by the absence of prominent groups of carinal and dorso-
lateral spinelets, by the generally more uniformly distributed dorsal spinelets which
are of a pretty uniform size, by the narrower abactinal area and higher position of
superomarginals on the side of the ray, and by the well developed series of actinal
plates. It differs from camtschatica in lacking any trace of serial arrangement of the
abactinal spinelets and in the absence of any crowding or differentiation of the carinal
spinelets, the same distinction separating it from dispar. It further differs in having
shorter rays, the intermarginal and actinal papular areas confluent, and very obtuse-
ovate or elliptic, small straight pedicellariae.

Asteira is probably one of the camtschatica group of species, as evidenced by the
well developed actinal plates. It appears to be distinct from all the forms of
camtschatica and heractis. The pedicellariae are rather distinctive.

LEPTASTERIAS LEPTODOMA, new species
Plate 43, Figures 2, 2a, 2b; Plate 50, Figure 6

Diagnosis.—Rays six. Skeleton weak; abactinal spinelets small, rather widely
scattered; carinals not clearly differentiated; superomarginal, inferomarginal, and
actinal spines typically one to a plate, forming three definite lateral series well spaced
from one another; adambulacral plates monacanthid; apical oral spines rather long;
lanceolate dermal straight pedicellariae; integument rather thick pulpy, and thrown
into irregular transverse folds; abactinal integument rather flaccid. Rays slender,
well arched. R 29 mm.,r7mm., R=4-+r; breadth of ray at base, 7 mm.

Description. —Abactinal skeleton weak, entirely obscured in alcoholic specimens
by pulpy integument which is raised in low, irregular, transverse folds and circular
disks surrounding the widely spaced small spinelets. These cushions seldom carry
pedicellariae abactinally. Abactinal skeleton very irregular, the plates thin, small,
irregularly elliptical, oblong, three or four lobed; skeletal intervals small to large;
usually a series of large intervals directly above superomarginal plates. Lobed
plates usually carry a single short, tapered, subtruncate spinelet (two or three on a
few plates of disk). These are spaced 3 to 10 spine lengths apart and sometimes
form three very irregular longiseries, or else are without serial order. They vary
greatly in number but for the genus are few and widely spaced.

Superomarginal spinelets, a little longer than abactinals, one to a plate, sur-
rounded by a pulpy disk bearing one to several crossed pedicellariae, They form »
very regular series rather high up on the side of the ray.

Similarly there is a single series of inferomarginal, and one of actinal spinelets
(extending two-thirds to three-fourths the length of the ray). In the largest speci-
mens there is a second short series of actinals. The spinelets are slender, terete,
blunt, and increase in length toward the furrow, the actinals being longest (upward
of 1.5 mm.) and sometimes curved. The intermarginal and actinal channels (both
lateral in position) are broad, so that the three lateral series of spines are well separated
and the actinals are also fairly well spaced from the adambulacrals. The inter-

106 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

marginal and actinal papular areas are frequently confluent, forming very large sub-
quadrangular spaces reaching from the superomarginal to the actinal plates. It
may happen that a whole series of inferomarginal plates is thus disconnected.

On either side of ray there are three fairly regular series of papulae—an actinal,
an intermarginal, and a supramarginal, with one to four in each cluster.’ The abacti-
nal, one to three in each area, are very irregularly distributed and the areas are vari-
able in number. Usually the papulae are few.

Adambulacral plates monacanthid. The spinelets are very slender, terete,
blunt, a little shorter and considerably slenderer than the actinals. The spinelet of
alternate plates is situated on the furrow margin while the other is spaced back from
margin. In the type there is very little difference in size between the two; but in the
largest specimen from Unalaska, the outer is longer, heavier, and slightly compressed
at the truncate tip. The inner tapers a little, as is usual with furrow spines. First
pair of adambulacrals contiguous interradially, conspicuously longer than second

air.
: In the type, 5 of the 10 oral plates carry 2 suboral spines, one carries 3, the
rest 1. Other specimens generally carry one. There are two apical spines, both
tapered, slender, the inner a little longer than the plate, the outer (adjacent to furrow)
nearly as long. Both are longer than in alaskensis.

Straight pedicellariae (pl. 43, fig. 2), slender to broadly lanceolate, are scattered
over lateral abactinal regions; others occur in the intermarginal and actinal channels,
in clusters on the oral spines, and sparingly on the furrow spines. Very small ones
are scattered along the furrow face of the adambulacral plates. Intermarginal
pedicellariae measure about 0.5 mm. long.

Crossed pedicellariae, very scarce abactinally. A few form rings around the
superomarginal spines (sometimes absent), and small clusters on the inferomarginal,
actinal, and adambulacral spines. The abactinal and superomarginal (pl. 43, fig. 2a)
measure 0.2 to 0.25 mm. long.

Madreporice body variable in size, medium, situated a little nearer to margin
than middle of r.

Young.—The smallest specimen has R 8 mm., and like larger examples has a
thick pulpy skin, thrown into folds or welts by irregular deep creases. There are a
few carinal spines and one series of superomarginals, one of inferomarginals, and one
of adambulacrals, the latter well spaced from the inferomarginals. The spinelets
are all slender, and the carinals are nearly obscured by the integument.

Anatomical notes.—The spinelets are fewer and the plates thinner than is usual
in 6-rayed species. The primary spiniferous plates are irregularly roundish or
short-lobed and are connected by from 1 to 3 overlapping, flat, spineless ossicles.
(Pl. 43, fig. 2b.) These form the connectives of a very irregular mesh, which abactin-
ally may be open, or fairly close. The spaces are larger toward the end of the ray
than on the disk and base of arm. The superomarginal plates are larger than the
inferomarginal and either three or four lobed. The latter frequently have only the
dorsolateral lobes developed. They are then not connected in series, but only with
the superomarginals and actinals. When the longitudinal connective is present it
is quite slender.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——-FISHER 107

The gonads open ventrally. The ovaries are nearly globular and are mature
in an example with R 25 mm., and in another with R only 15 to 17 mm. (Agattu,
June 8, 1906).

Type—Cat. No. E 1497, U.S.N.M.

Type locality —Humboldt Bay, Shumagin Islands.

Distribution —From the Shumagin Islands to Attu and Agattu (at end of Aleu-
tian chain).

Specimens exramined.—Twenty-four.

Specimens of Leptasterias leptodoma examined

j Collection
mens
ORR eel ete este chee erm te tale 1) U.S.N.M., W. H. Dall.
etek: opt oatc <b hace as 4| U.8.N.M.
tiesectandiec alepehisabei seis a ipa sitesindale shsbte stat we oi A eee 5 | Stanford, H. Heath, G. D. Hanna; U.S.N.M.
Nazan Bay, ‘Atka Island, Aleutian Islands _. Se EAA a 2| U.S.N.M., W. H. Dall, 1873.
Atka Island, Aleutians. ............ ..do. 1} U.S.N.M.
Adakh Island, Aleutians. . tain wi «do. 1 Do.
Attu Island, Aleutians........__- a pee AI 1 Albatross, June 10-11, 1906.
Agattu Island, Aleutians... ..... Sikes Sew Shore rocks - 9 Albatross, June 8, 1906.

Remarks.—L. leptodoma is notable for the pulpy, welted integument, very irregu-
lar, almost papery, abactinal skeleton, and few abactinal spinelets and pedicellariae.
It may be distinguished from other small 6-rayed species by the three regular lateral
longiseries of marginal and actinal spinelets and by the very few small abactinal
spinelets.

The well-developed actinal plates differentiate the species from any of the forms
of alaskensis; while the absence of any carinal differentiation prevents confusion with
the various forms of heractis, notably forma aspera which has rather heavy pulpy skin.

LEPTASTERIAS HEXACTIS (Stimpson)
Plate 44; Plate 45, Figure 2; Plate 50, Figure 7; Plate 51; Plate 52; Plate 54, Figure 6.

Asterias hexactis Stimpson, Proc. Boston Soc. Nat. Hist., vol 8, 1862, p. 272. —Verritt,
Trans. Conn. Acad. Sci., vol. 1, 1867, p. 326.—Be.t, Proc. Zool. Soc. London, 1881,
p. 495.

Leptasterias hezactis Vernivi, part, Shallow-water Starfishes, 1914, p. 126, pl. 25, figs. 7, 8
(type).

Leptasterias epichlora alaskensis var. siderea Vernritt, Shallow-water Starfishes, 1914, p.
137, pl. 16, figs. 3, 4 (Yakutat Bay).

Leptasterias epichlora miliaris var. regularis Verritt, Shallow-water Starfishes, 1914, p.
139 (Cape Fox, Alaska).

Leptasterias epichlora plena Verniii, Shallow-water Starfishes, 1914, p. 140, pl. 58, fig. 1
(Comox or Sooke, British Columbia).

Leptasterias epichlora var. plena Verriiy, Shallow-water Starfishes, 1914, plates, p. 118.

Leptasterias epichlora pugetana Vennii1, Shallow-water Starfishes, 1914, p. 142 (Puget Sound).

Diagnosis.—A small 6-rayed, polymorphic species, allied to camtschatica, having
slender to robust rays, a well developed actinal series of plates, irregular dorsal spinu-
lation, one to five (according to forma) superomarginal spinelets, and usually well

108 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

differentiated carinals. Differing from alaskensis in having a long series of actinals,
a narrow dorsolateral area, conspicuous carinal series, superomarginals higher on side
of ray, and in lacking conspicuous bivalved pedicellariae.

Heretofore the name hexactis has been applied to almost any small 6-rayed
Leptasterias with slender rays and relatively few spinelets. Verrill reproduced Stimp-
son’s original description but his own published observations were confined to speci-
mens of L. pusilla from Monterey. What his “numerous specimens from Puget
Sound and the Queen Charlotte Islands” may have been I have no way of knowing,
since they can not be located. Certainly duplicates of Stimpson’s puzzling types
are not common in that region.

It is necessary to repeat that among the numerous small 6-rayed sea stars of the
northwest coast of America specific lines are exceedingly difficult to draw; but nowhere
is there such a confusion and intermingling of forms as in the sounds of Washington
and British Columbia. Specific lines which are fairly distinct elsewhere here break
down and one of the reasons is probably hybridization, as suggested by the presence
of a minority of intermediate and freakishly aberrant specimens. Hybridization
would be likely and simple enough if the breeding seasons coincide; but it is difficult
to prove that it takes place.

I have used hexactis in a different and much broader sense than has Verrill, to
include the common 6-rayed Leptasterias of the Puget Sound region having irregular
abactinal spinulation, a fairly regular and narrow carinal band of spinelets, and usually
monacanthid or diplacanthid superomarginals (forma regularis), This ranges from
Cape Flattery to Kodiak Island, while an intergrading paucispinous variation (aspera)
can be traced as far as the Shumagin Islands. A multispinous form (plena) appears
to intergrade with both forma regularis and heractis in the Puget Sound region;
while its northern counterpart, forma siderea, intergrades with regularis and probably
also with L. camtschatica dispar (through forma nitida at Kodiak).

I have studied Stimpson’s type and type series. Probably if an attempt had
been made to find the most puzzling specimens in the Puget Sound region no better
selection could have been made. The type series, No. 1368, contains three examples
of forma hexactis with R 32 to 40 mm. (pl. 51, figs. 2, 3), a small relatively stouter-
rayed specimen with R 20 mm., and monacanthid superomarginals, referable to
forma regularis (pl. 51, fig. 1) and 14 equally small, stout-rayed examples of a sub-
form of plena (figs. 5-9). Whether these were collected together is not known, but it
is very probable.

In the original description, reproduced by Verrill, Stimpson has incorporated
some of the characters of these small plena, as, for instance: ‘‘In some specimens the
lateral spines are distinct from the ventrals, being separated from them by a channel,
and forming a crowded row of confluent clusters like the dorsals. Dorsal spines small
and numerous, in little heaps, which being confluent in a longitudinal direction form
three or five (according to the distance from the disk) rows, separated from each other
by corresponding rows of papuliferous depressions.” This refers to small plena
having R about 20 mm. such as Plate 51, Figures 6, 8; Figure 5 is somewhat inter- -
mediate with regularis. In the following sentence he says: ‘These spines in some

specimens, however, are fewer and do not form heaps.’’ This refers to the type and
two cotypes; Figures 2, 3.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 109

The relationships of heractis, in the narrow sense, are very obscure. I doubt if
there is a homogeneous forma heractis. These gaunt, rather paucispinous specimens
with one or two superomarginal and rather few irregular, dorsolateral spinelets
behave like fortuitous variations, probably of forma regularis. The small specimen
(pl. 51, fig. +) in the type series suggests this and I have what are apparently a few
variants of regularis (San Juan Island) which resemble the cotypes of forma heractis.
They appear to me to be something distinct from the paucispinous forms of ran-
couveri, for instance.

After camtschatica, the name heractis has priority for any 6-rayed Leptasterias.
It is unfortunate, therefore, that the types are what they are. I believe, however,
that the relationship of this series of specimens is with forma regularis and not at all
with aequalis, despite the peculiar behavior of vancouveri. Both regularis and hez-
actis, among other items, lack the two or three enlarged lateral teeth of the terminal
lip of the crossed pedicellariae, such as are characteristic of aequalis, This is an
insignificant character but in the almost hopeless mess of 6-rayed forms in the Puget
Sound region, everything helps.

No adult specimens with slender rays (forma heractis) have been taken north of
Boundary Bay (between Washington and British Columbia) which is another curious
fact. As forma regularis is the prevalent variety of heractis—the species—in the
Puget Sound region, so also forma regularis and its less spiny counterpart, forma
aspera, are the predominant types over the extensive coast between Boundary Bay
and the Shumagin Islands. Forma siderea and plena are not so common, unless
possibly on more exposed outer shores.

The picture of the species heractis should be based upon regularis and aspera
(which are really subformae of a variable forma) rather than upon forma heractis
which is a rare, problematical variation. The fact that the type of the species is
what it is constitutes an unfortunate episode in the taxonomic history of Leptasterias.

Forma HEXACTIS (Stimpson)

Plate 44, Figures 1, la-lf, 5; Plate 50, Figure 7; Plate 51, Figures 2, 3

Superomarginal spinlets two or one, sporadically a group of three; inferomargin-
als usually one. Rays slender, fairly long and attenuate distally; abactinal spinelets
cylindrical, tapered or slightly clavate, well spaced; a fairly regular carinal series of
single spinelets or of groups of two or three (occasionally four or five); dorsolateral
spinelets usually one, sometimes two to a plate proximally forming two irregular
longiseries and distally one, or in extreme cases one only throughout. In this case
there are a few scattered spines at base of ray. One series of actinal spinelets extend-
ing one-half to three-fourths length of ray. Crossed pedicellariae with the last
external tooth of terminal lip sometimes slightly enlarged; straight pedicellariae
triangular ovate to slender ovate, or triangular lanceolate. R 41 mm., r 8 mm.,
r= 6.9,

Specimens examined.—Stimpson’s types, three specimens, No. 1368, U.S.N.M.;
Puget Sound, two specimens, Plate 50 Figure 7, Stanford collection; San Juan
Islands, Wash., Stanford collection, one specimen; Boundary Bay, British Columbia,
Stanford collection, six specimens.

110 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Forma ASPERA, new forma

Plate 44, Figures 4, 4a—4c; Plate 45, Figures 2, 2a; Plate 51, Figure 4; Plate 52, Figure 6

Diagnosis.—Differing from forma regularis with which it freely intergrades in
having relatively few dorsolateral spinelets; usually a definite adradial and supra-
marginal band free of spinelets; typically a single series of superomarginal, infero-
marginal and actinal spinelets; adambulacrals monacanthid; rays six, robust, typi-
cally with a thick skin; papulae large. R 43 mm., r 13 mm., br. 13 mm., R=3.3 r
(St. Paul, Kodiak).

In this paucispinous, but robust phase of hexactis there are extreme specimens
with a’single prominent carinal series of usually contiguous subcylindrical truncate or
subcapitate spinelets; a single zigzag series of widely spaced dorsolaterals with a few
scattered spinelets at base of ray; superomarginals in one regular series, inferomar-
ginals in one or two; actinals in one series; meshes of abactinal skeleton rather open.
Then there are others in which accessory smaller carinal spinelets are added on some
of the lateral lobes of the plates, usually on the proximal part of ray only. (This
phase of spininess corresponds to that of Pisaster brevispinus forma paucispinus.)

These specimens when dried have a rather rough, meager appearance in spite of
the stoutness of the rays. Owing to the thornlets of the spines the surface is raspy.

Typical specimens are found at Nagai, Shumagin Islands; Kukak Bay, Shelikof
Strait; St. Paul, Kodiak; Litnik Bay, Afognak Island; Prince William Sound (Cor-
dova); Port Etches, Alaska; Funter Bay, Lynn Canal; Wrangell; WhiteWater Bay,
Alaska; and Alert Bay, Vancouver Island.

Well developed types from St. Paul, Kodiak Island, have a regular prominent
series of close-set, subcapitate, striate, unequal, carinal tubercles. The dorsolateral
area is very irregularly beset with relatively few, widely spaced, unequal tubercles
becoming sometimes transverse groups of three or four. These increase in number
by interpolations until a broken areolated pattern of regularis is achieved. Some of
the dorsolateral tubercles are conspicuously heavier than the carinals or they are
subequal. There is usually a well-defined bare zone above the regular single series
of subcylindrical or tapered superomarginals. The latter define the dorsolateral
margin of ray. Frequently there is a similar adradial channel. Intermarginal
channel well marked. Inferomarginal spinelets usually one, sometimes two, tapered,
blunt, a little longer and heavier than upper series; actinals similar to above extending
in a single series far along ray. Adambulacral spinelets subclavate, about as long as
actinals, but much slenderer, gradually lengthening on first 6 to 10 plates; beyond
base of ray the alternates are slenderer and more advanced into furrow; mouth plates
with fairly well developed actinostomial spine, about as long as plate; occasionally
a second small companion; suboral longer than first adambulacral and actinostomial.

The specimens from Kukak Bay, Shelikof Strait, Alaska, are notable for having
unusually slender ossicles and large papular areas. The spinelets are weaker than
normal and straight pedicellariae small.

Specimens from the vicinity of Cordova, Prince William Sound, exhibit an
uninterrupted series from typical aspera to paucispinous examples of regularis.
Some specimens of both forms have well developed ovoid straight pedicellariae on
the abactinal, lateral, and actinal surfaces. The skeletal meshes are large.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 111

Alcoholic examples usually show a pulpy skin, prominent sheaths around the
spinelets, and large papulae. This is particularly true of southern Alaskan and
Alert Bay examples, but is not of geographic significance since the Nagai and Afognak
specimens are similar.

Crossed pedicellariae not very numerous, 0.19 to 0.25 mm. long. (Pl. 44,
fig. 4a.) In northern examples the lateral teeth of the terminal lip are characteristi-
cally slender. Straight pedicellariae ovate, slender ovate, tapered, the tip of jaw
either narrowed or slightly spatulate; largest in interradial channel, sometimes
(Cordova) fairly conspicuous. (PI. 44, figs. 4b, 4c.)

Specimens collected at St. Paul, Kodiak, April 12, 1926, by Mr. S. P. Smith,
were carrying large masses of eggs on the oral side of disk.

Color in life—Specimens from Kodiak (St. Paul) only a month in formalin,
were colored dull maroon, sometimes very deep, mottled with cinnamon or pinkish
cinnamon.

Type of forma aspera—Cat. No. E 1498, U. S. N. M. (from Hopkins Marine
Station).

Type locality of forma aspera.—St. Paul (Kodiak), Kodiak Island.

Distribution —Nagai, Shumagin Islands to Alert Bay, Queen Charlotte Sound,

British Columbia.
Forma REGULARIS Verrill

Plate 44, Figure 6; Plate 51, Figure 1; Plate 52, Figures 3-5
Leptasterias epichlora miliaris var. regularis VERRILL.
Leptasterias epichlora miliaris var. subregularis VERRILL.
Leptasterias epichlora pugetana VERRILL.

Diagnosis.—Differing in appearance from L. alaskensis in having a long series
of monacanthid actinal plates; a well marked carinal series of small groups of spine-
lets; superomarginals well up on side of ray; small to medium sized ovate or ovate-
lanceolate straight pedicellariae; mostly monacanthid adambulacral plates.
Superomarginal spines one or two (sporadically three); inferomarginals two; a well
developed inter and supra marginal channel. Rays six, rather short and robust;
skeleton stout, irregular dorsolaterally; dorsolateral spinelets rather numerous, and
coarse, scattered or forming a poorly defined reticulum.

This is one of the few forms of Leptasterias described by Verrill of which authentic
specimens can be found. I have carefully examined one from Cape Fox, the only
listed locally. This is on the mainland opposite the northern end of the Queen
Charlotte Islands.

The Cape Fox example has rather fewer abactinal spines than the majority of
the specimens from the Puget Sound region and certain similar ones from Sitka,
Loring, and Homer, Alaska, to which Verrill’s name pugetana would be applicable.
This subspecies was based upon an example from Puget Sound having the unusually
large ovoid pedicellariae which are found also on some specimens of undoubted
regularis (Cordova, Alaska; Victoria, British Columbia). Pugetana simply repre-
sents a more multispinous phase of regularis and occurs among specimens from
Cooks Inlet, Alaska, to Neah Bay, Washington.

Verrill’s specimen of regularis is therefore a rather paucispinous example of
forma regularis. It is necessary to correct a serious error in Verrill’s short diagnosis

112 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

of variety regularis (1914, p. 139). He says: ‘‘The second short interactinal row of
ossicles bearing spines is not present, as it is usually in other varieties when of the
same size, and this is probably the cause of the slenderness of the ray.’’ A second
series of actinal ossicles occurs in well-developed specimens of camtschatica and
aequalis but never in the alaskensis series, and never, so far as I have observed, in
hexactis. Verrill more than once mistook young Hvasterivas for 5-rayed examples of
some of his varieties (as, var. subnodulosa 1914, p. 139). These have regularly two
series of actinals. Verrill’s Leptasterias macouni is an Hvasterias.

The largest specimens which I have seen were collected by Mrs. Ida S. Oldroyd
at Friday Harbor, San Juan Islands, Wash. (pl. 52, fig. 3). Rays medium slender
to medium robust; R 39 mm., r 12mm.; ray width 9 toll mm.; rays rather blunt
and stubby, sometimes inflated. Abactinal spinelets, stout, subcylindrical to sub-
capitate truncate, arranged in irregular broken reticulum, the carinals forming an
irregular multiple series—three to six spinelets per plate; dorsolaterals two or three
to six to a plate. Papular areas in the equivalent of six abactinal longiseries, only
those above superomarginals being at all regular; three to six abactinal papulae
to an area.

Superomarginal spinelets two, sometimes three to a plate proximally, sporadically
one. Inferomarginals proximally one, then two; actinals one, these forming a trans-
verse comb of two or three with the inferomarginals. They are subcylindrical or
tapered, blunt, sometimes upturned at end. Adambulacrals with one spinelet except
near base of ray where for a short distance there are one and two. Oral plates with
one actinostomial spinelet a little shorter to a little longer than plate and one suboral
a trifle longer than proximal adambulacral spinelets.

Straight pedicellariae ovate or triangular ovate, compressed, the end of jaw
with sometimes two or three denticles. Good sized ones measure 0.4 to 0.5 mm.
Crossed pedicellariae not very abundant, the abactinal 0.2 to 0.22 mm. long. They
are not particularly diagnostic. (PI. 44, fig. 6.)

Three specimens from the vicinity of Victoria have mostly monacanthid supero-
marginals, while the others have two or three spinelets. The straight pedicellariae
are sporadically larger, or more numerous, in certain specimens. Such examples
formed the basis for Verrill’s Leptasterias epichlora pugetana.

Type of forma regularis—In Yale Museum.

Type locality of forma regularis —Cape Fox, Alaska (opposite Dixon Entrance).

Distribution —Puget Sound and Vancouver Island to Cooks Inlet. The northern
examples are less spinous than the southern and less easy to differentiate from aspera
which predominates in the north.

Forma SIDEREA Verrill
Plate 44, Figures 2, 2a; Plate 51, Figure 1; Plate 52, Figures 1, 2; Plate 54, Figure 6

Leptasterias epichlora alaskensis var. siderea VERRILL.

Diagnosis.— Differing from forma regularis in having still more numerous abac-
tinal spinelets which are usually crowded into an areolated pattern; superomarginal
spinelets two or three as against usually one of northern examples of regularis;
straight pedicellariae small lanceolate or ovate lanceolate.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 113

This is still spinier than f. regularis and seems to be a northern counterpart of
f. plena. Siderea occurs at Karluk, Kodiak, with a variety of L. camtschatica f. nitida
which it resembles, and with which it possibly intergrades. The two forms are
evidently very closely related. Siderea differs in having the dorsal spinulation more
areolated, without longiserial arrangement. A comparison of Plate 52, Figures 1, 2,
with Plate 50, Figure 1, will answer better than detailed description. This variety
of nitida may constitute an intergrading subforma since no large typical nitida has
been found at Kodiak.

Verrill’s type of siderea is lost but his figure is duplicated by a specimen from
Karluk, Kodiak Island. (PI. 52, fig. 2.) The abactinal spinelets are medium robust,
capitate, rather truncate, 0.4 to 0.6 mm. long, and the vertical ridges are armed with
flattened often leaflike thornlets similar to those of L. aequalis. Spinelets not so
prominent along the carinal line as in the Middleton Island specimen. The adam-
bulacral spinelets are robust, clavate, irregularly one and two, or only one to a plate;
actinal spinelets still stouter and a little longer; inferomarginal spinelets subequal to
actinals; superomarginals intermediate between inferomarginals and abactinals.
The apical mouth spines are stout, about as long as plates. Straight pedicellariae
rather inconspicuous, elliptical ovate to lanceolate ovate, compressed. The crossed
pedicellariae (0.22 mm. long) do not have comspicuous teeth on border of terminal lip.

Two specimens from Anchor Point, 20 miles west of Homer, Cooks Inlet, are
remarkable for their large size. The larger has R 80 mm.,r18 mm., ray width 20 mm.
These specimens do not have the maximum number of dorsolateral spinelets, which
are coarse and sometimes tubercular; and the proximal superomarginals have some-
times only one spine, while the distals have two. Straight pedicellariae rather
plump, lanceolate to ovoid. These specimens may be classified either as multispinous
regularis or as less spiny siderea.

Type of forma siderea.—Lost.

Type locality of forma siderea.—Not stated; Yakutat, one of the localities
mentioned by Verrill, may be designated.

Distribution.—Kodiak Island and Middleton Island to Yakutat Bay.

Forma PLENA Verrill
Plate 44, Figure 3; Plate 51, Figures 5-9; Plate 52, Figure 7

Leptasterias epichlora plena VERRILL.
Leptasterias epichlora var. plena VERRILL.

Diagnosis.—A southern more multispinous counterpart of siderea which differs
in having the abactinal spinulation somewhat more clumped, the groups of spinelets
sometimes subdivided into five very irregular multiple longiseries of spinelets by six
irregular longiseries of papular depressions; superomarginals three to five usually
unequal, in a vertical series or a group; occasionally six or seven on a few proximal
plates and two on distal; inferomarginals two or three; actinals one; adambulacrals
one and two in alternation over most of ray; a few proximal plates sometimes consecu-
tively diplacanthid; furrow spinelet of diplacanthid plates slenderer than the others
which are subcylindrical to clavate. R 40 mm., r 12 mm., br. 12 mm., R=3.3 r

Forrester s and).

114 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

This group ranges from Puget Sound to southern Alaska and probably inter-
grades with siderea which it much resembles. It also suggests a coarse spined form
of L. aequalis with indefinite abactinal longiseries of spinelet groups. The spinelets
are like those of siderea in size and structure.

Straight pedicellariae longer than broad, triangular lanceolate or lanceolate
ovate, similar to those of siderea. Crossed pedicellariae (pl. 44, fig. 3), with the
lateralmost tooth of terminal lip slightly enlarged.

I have examined Verrill’s type which is figured (1914, pl. 58, fig. 1). The
specimens from Forrester Island resemble the type, which is said to be from either
Comox or Sooke, Vancouver Island. Practically all the other specimens I have seen
from the Puget Sound region, including those in the type series of hexactis, are small
(R 10 to 26 mm.), and easily confused with small L. aequalis. (Pl. 51, figs. 5 to 9.)
It is a hopeless task to differentiate such confused forms by words. However, these
small plena differ from aequalis in having fewer, coarse spinelets, the coordinated
groups being more irregular; or else the abactinal spinelets are arranged in a close
reticulum. Both phases differ from regularis in the more numerous abactinal spine-
lets and the presence of three to five superomarginal spinelets in an angular series
oragroup. (Puget Sound; Alert Bay, Vancouver Island.) Two or three specimens
of Stimpson’s type series of hexactis (1368 a, 6) stand between these depauperate
plena and the actual type specimens, while one (1368 d), is intermediate with regularis.

Type of forma plena.—Victoria Memorial Museum, Ottawa.

Type locality of forma plena.—‘Sooke or Comox, B. C.”’ (label of type specimen).

Distribution —Puget Sound region, Vancouver Island, and southern Alaska.

Type of Leptasteriae hexactis —No. 1368, U.S.N.M., Dr. C. B. Kennerly.

Type locality —Puget Sound, Wash.

Distribution.—Shumagin Islands to Cape Flattery, Wash., chiefly intertidal: to
25 fathoms in the Puget Sound region.

Specimens ecamined.—Three hundred and twenty-two.

Specimens of Leptasterias hexactis examined

Number of specimens of forma—

Locality

} Collection
her- | regu-
laris

aspera siderea | plena

actis

Nagai, Shumagin Islands........-.--------|-------- We a U.S.N.M.

Kukak Bay, Shelikof Strait peno | oooaaaa| Stanford, H. C. McMillin.
Weoulak island) s= 2. 2. . onscreen cco | Secneene 1 Coll. Wosnessensky, 1858, from Acad. Sel.,
U.R.S.S.

St. Paul, Kodiak Island.._......-....-_._- |p==--=== BSilGsececes 1S eee eee Stanford, S. P. Smith; U.S.N.M., W. J.
| Fisher.

Karluk, Kodiak Island-._.._..-.......-__- (eee eeepc uaee se eee Miho a eee Stanford, C. Rutter.

Arognak laland =<. 2.) ee aes 1, |Ee2" ea | ee hue saa U.S.N.M., Albatross.

Middleton Island, Gulf of Alaska._______- peaee SA Pie 1| dean U.S.N.M., W. H. Dall.

Port Graham, Cooks Inlet_.._........_._- [oaweeeee potaaosts V2 lo escculeves co U.S.N.M.

Seldovia, Cooks Inlet.........-..-....-... |---~--=-|-------- 12 enol neers Do.

Homer, Cooks Inlet._.__.---....._- Banner eee eens) torneo eal Medarene PO eeece an Do.

Cordova, Prince William Sound_____._...|..._-.-- 117 LO 9 eee Stanford, A. W. Greeley, H. C. McMillin.

Observation Island, Cordova..............|...----- 12 Die cae [teensy U.S.N.M.

' Some specimens more or Jess intermediate between aspera and regularis; not extremes of either forma.
7 See note under siderea; somewhat intermediate with regularis.

ASTEROIDEBA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 115

Specimens of Leptasterias hexactis examined—Continued

Number of specimens of forma—

Collection

pe. ee U.8.N.M., W. H. Dall.

Punter Bay, Lynn Canal. ................ U.S.N.M., Albatross.

Sitike, AGUA... asipdncasignannsewenncaeyvese Stanford; U.8.N.M.
WENNER ioe naprnansccecnncepee noee----|-e------| U.B.N.M., W. H. Dall, W. H. Jones.
pe a ee eS eee ee U.S.N.M.

Security Bay, Alaska... U.8.N.M., W. H. Jones

White Water Bay, Alaska.. ad Do.
Cape Fox, Southeast Alaska __ Yale, W. R. Coe.
Forrester Island, Southeast A 5S | ie Stanford, H. Heath.
Fort Rupert, Vancouver Island. se 4 U.8.N.M., Harlan Smith.
Alert Bay, Vancouver Island Stanford; U.8S.N.M.
Vancouver Island (Comox or Sooke) al Victoria Memorial Museum.
Boundary Bay, British Columbia. | n ...| Stanford, F. W. Weymouth.
Victoria, British Columbia. U.8.N.M., J. E. Benedict, Wm. Palmer,
j Albatross.
Orcas, San Juan Islands, Wash............ | Stanford, E. C. Starks.
San Juan Islands, Wash.................-- Cal. Acad. Sci., shore to 25 fathoms, Ida
Oldroyd.
PRINT tina ceca thane gseatniptmresenredy U.S.N.M., Dr. C. B. Kennerly.
Ne Si Eo J ho le gad ane U.8.N.M., D. 8. Jordan; Stanford, E. C.
} Starks.
HASOIR 08 WM fo soncccacecsnneiseGdudacns Stanford, Ira Cornwall; U.S.N.M., D. 8.
Jordan.
Neah Bay (Cape Flattery), Wash. U.S.N.M., J. G. Swan.

} Some specimens more or less intermediate between aspera and regularis; not extremes of either forma.

1 See under rancoureri. These are part of the series of rancoureri and are sufficiently large (R 30 mm. or more) to preclude the
possibility of their being the young of multispinous eancoureri. 3 specimens are pretty clearly intermediate with regularis.

«No. 1368, U.S.N.M., type and 2 paratypes.

# No. 1368, U.S.N.M., part of type series of Asterias hezactis.

* William Head west of Victoria. This is similar to the most regular of the San Juan Islands version of hezactis.

LEPTASTERIAS HEXACTIS VANCOUVERI (Perrier)
Plate 45, Figures 1, la-le; Plate 46, Figure 1; Plate 53

Asterias vancouveri Perrier, Ré¥. des Stell., 1875, p. 64; Ann. and Mag. Nat. Hist., ser. 4,
vol. 17, 1876, p. 35 (Vancouver Island).
Leptasterias vancouveri VerRILL, Shallow-water Starfishes, 1914, p. 125.

Diagnosis —Rays six. A problematical and very variable form, in its extreme
development resembling a coarse-spined L. aequalis but practically intergrading with
L. hexacitis. Rays moderately stout, tapered; spinelets typically arranged, by groups
or singly in longiseries; carinals three to five, dorsolaterals one to three, supero-
marginals two to five, inferomarginals two to one, actinals one per plate (two series
proximally); dorsolateral area typically narrow; superomarginal plates broad, with
typically three, four, or five spinelets in a vertical comb; usually six longiseries of
dorsal papular areas.

I examined Perrier’s type in the British Museum. It is one of the extreme, more
multispinous variations of the race having prominent regular carinal plates, fairly
regular dorsolaterals and four or five superomarginal spinelets.

116 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

I have given this extraordinary form subspecific rank in order to emphasize its
pecularities—structural, developmental, and geographic—and to free hexractis and
aequalis from a further encumbrance of formae.

Its range so far as known is restricted to the general region of the confluence of
the Strait of Georgia, Strait of Fuca, and Puget Sound. The large series which I
have examined was collected among the San Juan Islands, chiefly by Mrs. Ida Oldroyd.

Its younger stages are different from those of aequalis in having fewer, more
widely spaced dorsal spinelets (arranged in pretty regular longiseries). Its structural
pecularities concern the wide range of its variations. These variations include a
few specimens which appear to be intermediate with L. hexactis f. regularis, which
occurs with vancouveri at the San Juan Islands. Other extermes are closely similar
to L. hexactis f. heractis as noted below.

In order to arrive at some semblance of order the specimens will be divided into
groups on the basis of the number of superomarginal spinelets.

I. Superomarginal spinelets, four or five, rarely six, in a vertical series; a broad
conspicuous band of these combs extending along the dorsolateral curvature of the
ray (pl. 53, fig. 3); superomarginal plates broad and regular; inferomarginal spinelets
longer than superomarginal, slightly curved, two proximally and one or two distally;
a series of slightly longer actinal spines reaches far along ray while a second series
is usually present on the proximal third or half of ray. There is a definite channel
above the superomarginal spinelets and a narrow intermarginal one; in addition to
the dorsolateral papulae (see below) an intermarginal and two incomplete actinal
series (large examples). Adambulacral spines subcylindrical to subclavate, slender,
usually two at base of ray; then one and two, or one only.

a. Abactinal spinelets have the appearance of forming fairly regular longiseries (proximally two,
distally one) on either side of the regular carinal series. Spinelets short, typically robust,
cylindrical to subeapitate, round-tipped to subtruncate, coarser, more widely spaced and
fewer than in aequalis of the same size; dorsolateral spinelets one to three per plate, usually
two or three; carinal spinelets three to five; usually six discernible longiseries of abactinal
papulae. No small specimens.

b'. Carinals conspicuously broadened, nearly as wide as the dorsolateral area. Dorsolateral
plates very compact (one specimen only, R 60 rhim., r 10 mm., br. 10.5 mm.) (Pl. 53,
fig. 1.)

b Carinala narrower than either dorsolateral area (pl. 53, figs. 2, 3); groups of dorsolateral
spinelets (not individual spinelets) in more or less distinct longiseries separated by longi-
series of papulae (six for abactinal area); rays moderately robust; 11 specimens from R
32 mm. to R 54 mm. (latter with R=5.4r). Type of vancouveri belongs here.

a*, Abactinal spinelets similar in numbers and form to a! but not arranged in longiseries of groups
or only very indefinitely; even the carinal series of groups may lose its distinctness; irregu-

larity largely due to papulae falling out of longiserial alignment; R 55 mm.; eight specimens,
three more or less intermediate with a!. (Pl. 53, fig. 4.)

II. Superomarginal spinelets averaging three; in some specimens being proxi-
mally four, distally three; or rather irregularly three or four; or three, with two
spines on a minority of the distal plates; or three distally and two irregularly on some
on the proximals; dorsolateral plates with one to three spinelets but more often one
or two; frequently only one inferomarginal spine.

a‘. Abactinal spinelets arranged in fairly regular longiseries; six longiseries of papular areas in well
grown examples; carinal plates all rather narrow with three to five spinelets. Differing

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——-FISHER 117

from Ia! in that the dorsolateral plates carry one or two rather than two or three spinelets.
Whatever factors reduce the number of superomarginal spinelets act also upon the abactinal.
Fourteen specimens, R 26 to R 45 mm., including two or three intergrades with a’. Shorter
rayed, more robust specimens tend to have less regular abactinal spinelets. Twenty-seven
specimens R 15 mm. to R 25 mm. and from quite selnder (R 24 mm. br. 5.5 mm.) to decidedly
plump (R 23 mm.; br. 7.5 mm.) rays; dorsolateral spinelets usually solitary. Some of
these are probably young Ia', without the adult, full complement of spinelets.

a, Similar to Ia’, spines varying in coarseness; rays medium robust, none very slender; seven
specimens, R 22 mm. to 42 mm. ‘Two other specimens intermediate between a! and a
and with subcapitate, spaced spinelets coarser than in a’ resemble L. camtschatica. (PI.
53, fig. 5.)

III. Superomarginal spinelets two, sporadically three; dorsolateral plates with
usually solitary spinelets, as a consequence of which the spinelets appear well spaced ;
carinals with usually two to three spinelets; inferomarginals usually one, sporad-
ically two.

a', In the most regular examples, there are two longiseries of solitary, spaced, dorsolateral spinelets
on either side of the carinal series of plates, which carry three (two to four) spinelets; six
abactinal longiseries of small papular areas. Nine specimens R 28 mm. to R 42 mm.,
slender to medium robust rays; 42 specimens, R 7 mm. to R 25 mm., slender to robust rays.
A considerable number of these are evidently young of Ila', or even Ia'. (PI. 53, fig. 6.)

a?, Abactinal spinelets not in evident longiseries. Three variants are shown. (PI. 53, figs. 7, 8, 9.)
Figure 8 has the long rays of vancouveri but the abactinal spinelets are arranged much as
in L. hexactis forma regularis, and many of the superomarginals are monacanthid. This
and two smaller similar ones are intermediate in appearance between vancouvert and
regularis, but whether biologically so I am unable to say. Five specimens, R 30 mm. to R
54mm. The largest has somewhat swollen rays and three or four spinelets on a few supero-
marginals. There is a slight indication of serial arrangement of abactinal spinelets as
in Figure 7 (lateral parts of abactinal area). Straight pedicellariae figured. (PI. 44, fig. 5.)

In the foregoing enumeration of variations there is a pretty complete series of
gradations from forms (IIIa') which are difficult to separate from L. hexactis £,
hexactis (and others, IIa’, which appear to be intermediate with f. regularis) to one
which can easily be differentiated on the basis of three or four superomarginals (11).
This in turn passes without hiatus into typical vancouveri (Ia') which resembles
large, coarse spined aequalis and is therefore quite distinct from Class III.

In Class II there is a deviation toward the small variety of L. heractis forma
plena with which the type of heractis shows close relationship.

If large specimens of vancouveri are placed side by side with aequalis they may
be told at a glance: rancouveri has coarser, less numerous abactinal spinelets (espe-
cially on the carinals). If the small phase, or young, of Class II is also the young
of Ia, then the difference is even more marked, since the young of Ia, from having
conspicuously fewer spinelets, are quite unlike the young of aequalis. In this case,
the similarity between the adults of aequalis and vancouveri would result from
convergence.

The mouth plates do not offer any help. The actinostomial spine is about as
long as plate and usually has a small lateral companion.

Madreporic body conspicuous, about mid r, Crossed pedicellariae fairly
abundant, in circlets around the abactinal and superomarginal spinelets, They are
not particularly diagnostic, the enlarged tooth on outer side of lip being sometimes

118 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

slightly developed, sometimes suppressed. Straight pedicellariae ovate or triangular
ovate similar to those of aequalis. (PI. 45, figs. 1¢, 1d, le.)

Type.—In the British Museum.

Type locality—‘ Vancouver Island.” The specimen is so like the prevalent
form from the San Juan Islands that I think it probably came from that region.

Distribution—Puget Sound and adjacent regions of the Strait of Fuca and
Strait of Georgia; shore to 60 fathoms.

Specimens ecamined.—One hundred and twenty-six.

San Juan Islands, Wash. (Friday Harbor and Orcas Island) intertidal; a few
from 20 to 25 fathoms.

Seventy specimens of Classes I and II; 56 specimens of Class III, some of which
are young of vancouveri, others either forma hevactis, or facsimiles.

Remarks.—Some of the specimens of Class III, enumerated above, are L.
hexactis forma hexactis for ordinary purposes, and no diagnosis can be given which
will separate them effectively from the types of L. heractis. Yet some of Class III
are undoubtedly the young of Class II, probably also of Class I (typical vancowvert).
In all probability the type specimens of heractis have nothing to do with vancouwveri,
but are variants of regularis. The logic therefore is that some of the specimens of
Class IIT are genetically quite different from L. heractis, although so similar in

appearance.
LEPTASTERIAS PUSILLA, new species

Plate 45, Figures 3, 3a-3d; Plate 46, Figures 2, 2a—2d; Plate 54, Figure 5; Plate 55, Figures 1, 2, 7.
Leptasterias hexactis VERRILL, part, Shallow-water Starfishes, 1914, p. 127 (Monterey).

Diagnosis.—Differing from the slender phase of L. hexactis f. hexactis in its con-
stantly small size, in the different dentition of the crossed pedicellariae and different
shape of the straight pedicellariae which are constantly narrow-ovate or elliptical in
contour. Rays six, slender, subterete, tapering from a narrow base; disk small.
R 22 mm.,r5mm., R=4.4r (varying to 5r); breadth of ray at base5 mm. Spinelets
small, slender, spaced, the carinal plates with proximally one to three and the dor-
solaterals with usually one spinelet, all without serial order or else in five poorly defined
longiseries; superomarginals, two, sometimes three, inferomarginals two; actinals one.

Description.—Abactinal spinelets (pl. 46, figs. 2, 2a, 26) slender, longer than
thick, terete to subclavate, thorny, one to three to a plate on disk; on rays one to
three to a carinal plate; ordinarily one, occasionally two to a dorsolateral. Spinelets
0.3 to 0.4 mm. long, spaced by about their own length. They are arranged in about
five not very regular longiseries of which the median or carinal, composed of groups
of two or three spinelets, is not conspicuous as in aequalis. The dorsolateral plates
occasionally carry two spinelets. In some specimens the spinelets are not arranged
in definite longiseries. Size of papular areas variable. In one variety they are longer
than plates of ray and the skeleton appears open, while in the other the areas are
subequal to the plates or smaller. Papulae one to an area.

Superomarginal plates not conspicuous, four lobed with two slender spinelets
similar to but a trifle longer than the dorsals, one on the dorsal, one on the ventral,
and occasionally a third on the adoral lobe.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 119

Intermarginal channel well marked with a series of single papulae. Infero-
marginal spinelets two, the upper slender, subequal to or a little longer than the supero-
marginal, the lower 50 to 100 per cent longer, and much stouter, blunt. First three
to five, with one spinelet. A single series of similar actinal spines extends one-half
to three-fifths R. There are five fairly regular longiseries of lateral and actinal spines;
two superomarginal, two inferomarginal, one actinal.

Adambulacral armature: One spinelet on first three or four plates, then two
spinelets to a plate for a varying number; then alternately one and two, fairly regu-
larly. Some specimens have diplacanthid plates extending to middle of ray or even
farther. The spinelets are slender, terete, slightly clavate, round-tipped, a little
smaller than the actinals. The furrow spinelet is usually slightly slenderer than the
outer. First pair of adambulacrals in contact interradially and sometimes inner
corner of second pair in contact.

Mouth plates (pl. 45, fig. 3c) with the usual three spinelets. The inner of the
two actinostomial spinelets usually equals the plate in length, while the suboral is
appreciably longer.

Crossed pedicellariae (pl. 46, fig. 2d), distributed as in L. aequalis, are not espe-
cially abundant, there being one to six around each abactinal spine but attached to
the plate. In alcoholic specimens in which the skin is somewhat swollen they appear
nearly as thick as the spinelets, but shorter. There is usually one (on some examples
one to three) on each adambulacral spinelet; lacking on some spinelets.

The extreme lateral teeth of lip are not enlarged on outer side of jaw; length,
0.18 to 0.19 mm. Compare with L. aequalis (pl. 46, fig. 3) and L. heractis f. heractis
(pl. 44, fig. 1a).

Straight pedicellariae more elliptical, less triangular in contour and relatively
larger than in L. aequalis and L. heractis. (PI. 45, figs. 3, 3a, 35.) They are about
0.4 mm. long and are found very sparingly in the intermarginal channel, in the actinal
interradial channel, on proximal adambulacral spines and oral spines, and occasionally
on proximal inferomarginal and actinal plates, while smaller ones are found along
the furrow face of the adambulacral plates.

Madreporic body small, situated at middle of r and surrounded by a circle of
spinelets. Striae few, relatively coarse.

Color in life: Mottled olive green and pale straw color; carmine and pale straw
color; blotched with purplish red on a lighter ground.

Anatomical notes.—The skeleton (pl. 45, fig. 3d) in addition to one series of rather
small 4-lobed carinal and on either side two series of larger 4-lobed marginals, con-
sists in the adult of twice two series of dorsolaterals and on proximal part of ray of
twice two series of actinals. The marginals are the largest plates, and the supero-
marginals have a prominent descending lobe. The inner row of actinals are always
very inconspicuous since they do not carry spines.

Variations —This species is less variable than L. aequalis. The rays are never
short and stout as in some (but not all) examples of L. aequalis f. nana. They vary
in the degree of slenderness. As arule in large specimens R is about 5r. The skeletal
meshes vary from being smaller than the plates to appreciably arger. In the first
case the skeleton appears to be close-knit; in the latter decidedly open. The length
of the abactinal spinelets varies. Short-spined specimens are often difficult to separate

120 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

from L. aequalis f. nana of equal size. The latter may have slender rays, but have
more numerous closely-placed abactinal spinelets. The form of their straight
pedicellariae is a fairly reliable recognition mark, but these are sometimes absent in
very small specimens, and are occasionally narrower than normal in aequalis.

Type.—No. E 1499, U.S.N.M. (from Hopkins Marine Station).

Type locality Monterey Bay (Cypress Point), Calif.

Distribution Monterey Bay and Pillar Point (south of San Francisco Bay).

Specimens ecamined.—Numerous examples from Monterey Bay (Pacific Grove)
and the region between Monterey Bay and Carmel Bay, Calif.

One specimen from Pillar Point, San Mateo County.

Remarks.—During January this species carries eggs and embryos clustered in the
‘oral cavity of the actinal surface, and the tiny young leave the mother in February
either before or while aequalis is laying. The young of the latter are ready to fend
for themselves in late April or early May. There is thus a fundamental biological
difference between aequalis and pusilla.

Pusilla is not unlikely an extension of hexactis stock southward to Monterey.
But unlike hexactis of the north, it is comparatively homogeneous. Inasmuch as
pusilla shows certain constant differences, it seems best to give it specific rank since
hexactis is such a polymorphic species. Pusilla is a counterpart of forma heractis.

Verrill used this species as a basis for his comparison of hexactis and aequalis and
was hence led to erroneous conclusions as to the straight pedicellariae of hevxactis.

LEPTASTERIAS AEQUALIS (Stimpson)

Plate 46, Figures 3, 3a—3e; Plate 47; Plate 54, Figures 1-4; Plate 55, Figures 3-6; Plate 71, Figures
2, 2a—2c

Asterias aequalis Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 273.—VERRILL,
Trans. Conn. Acad. Sci., vol. 1, 1867, p. 327.

Leptasterias aequalis Verriti, Shallow-water Starfishes, 1914, p. 128, pl. 16, fig. 8; pl. 18,
figs. 1, 2; pl. 25, figs. 5, 6; pl. 56, fig. 5—Fisupr, Ann. Mag. Nat. Hist., ser. 9, vol. 12,
1923, p. 599.

Leptasterias aequalis var. compacta VERRILL, Shallow-water Starfishes,1914, p. 130, pl. 56,
fig. 5.

Leptasterias aequalis var. nana VERRILL, Shallow-water Starfishes, 1914, p. 132.

Leptasterias aequalis var. concinna VERRILL, Shallow-water Starfishes, 1914, p. 132

Diagnosis.—Size small to medium; rays six, slender to moderately stout, arched,
slightly swollen at base; abactinal and lateral surfaces compactly covered with gran-
uliform spinelets which form groups, aligned in more or less definite longiseries by
small papular areas; carinal series conspicuous, with numerous spinelets; supero-
marginal series broad with an angular or vertical series of four to eight spinelets;
inferomarginals with two longer spinelets; one long series of similar actinal spinelets;
crossed pedicellariae with enlarged, external lateral tooth on terminal lip; straight
pedicellariae few, small, triangular-ovate. Typical large specimen, R 52 mm., r 13
mm. R=4 r; breadth of ray at base, 12 to 14 mm. (PI. 54, fig. 4.)

Description.—The abactinal and lateral surfaces of the body have the appearance
of being covered with a uniform close granulation. In alcoholic specimens this
appears more compact by reason of the papulae and swollen sheaths of pedicellariae.
The spinelets increase gradually in length from the carinal toward the superomarginal
but are not precisely uniform on any one plate. They vary in length from about 0.35

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS--FISHER 121

to 0.7 mm. and are broader at top than at base. The finer structure is characteristic.
The vertical ridges carry expanded bladelike, hyaline, sharp thornlets, more de-
veloped than those of heractis.

The abactinal skeleton is compact and the groups of spinelets are usually arranged
in fairly definite longiseries, frequently interrupted and often not at all regular on the
dorsolateral plates. In typical specimens there is a broad carinal band of spinelets,
upward of 10 to a plate, and on either side the equivalent of two dorsolateral series,
that next to the superomarginals the more regular. There are three dorsolateral
longiseries of papular areas, smaller than the plates. Dorsolaterals with from one to
six spinelets. Plates not to be differentiated without preparation.

Superomarginals broad, the series bending upward at base of ray; plates armed
with a vertical (transverse) angular series, or shallow chevron, of five to eight spinelets
proximally (four or five in smallish specimens) and about five distally (two or three
in small examples). Spinelets similar to but slightly larger than the dorsolaterals and
usually slightly bent as in Plate 47, Figure la. The superomarginal band of spinelets
is broader but less compact than the carinal.

Intermarginal channel very narrow. Inferomarginal plates with two (occasion-
ally three) spinelets, the upper subequal to largest superomarginal spines, the lower
spine usually much larger, flattened and subtruncate at tip; sometimes both are
enlarged, especially distally. End of spinelets slightly upcurved. In compressed
spinelets the ventral border is curved as in a broad-tipped knife, while the upper
border is straight, or only slightly upcurved. Sometimes the spinelets are sub-
spatulate. Proximal plates may carry only one spine.

One series of actinal plates extends three-fourths to four-fifths R, and each plate
carries a single spine very similar to the larger inferomarginal. A second series of
smaller plates, armed each with a small spine (about size of smaller inferomarginal),
extends in large specimens about two-fifths to over half R, and is absent in most
small ones.

Adambulacral plates variable. The plates of proximal third or half of ray have
one and two spines in irregular alternation, sometimes continuing to near end of ray;
or the plates of outer part have only one spine; the proximal plates, or a majority,
may carry two spines while the distal are alternately monacanthid and diplacanthid,
or else only monacanthid; or else all the plates are monacanthid with the exception of a
few scattered diplacanthid plates proximally. In a series of monacanthid plates
alternate ones have a slenderer, more tapered spine somewhat advanced into the
furrow. These plates lack the outer spine of the couple. The other plates, lacking
the furrow spine, carry a somewhat heavier subclavate or subspatulate spine, usually
turned away from the furrow. The first two or three pairs of adambulacral plates are
in contact interradially. First plate somewhat longer than second.

Mouth plates of the usual small size. For form and armature see Plate 47,
Figures 2, 2a. While the longer of the two actinostomial spines is usually as long or
a little longer than the plate, there is considerable individual variation, and specimens
from Monterey Bay may have this spine considerably shorter than the plate. The
suboral spines are usually a little longer than the adoral adambulacral spines.

Papulae in small areas, few to an area, but rather conspicuous in living specimens.
There are about three dorsolateral longiseries, one intermarginal and two actinal.

122 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Crossed pedicellariae (pl. 46, figs. 3, 3a-3d) with the usual distribution. They
are attached to plates at base of dorsal spinelets but are not sufficiently numerous
to form wreaths as in L. hexactis. Interrupted wreaths are present on superomarginal
spinelets and small tufts on outer side of inferomarginals, actinals, and adambulacrals.
The abactinal pedicellariae measure 0.19 to 0.22 mm. in length and are characterized
by the increased size of the external lateral teeth of the terminal lip of the jaw as
shown in Plate 46, Figure 3. Sometimes only one tooth is enlarged, sometimes
several. The adambulacral pedicellariae have a somewhat longer lip and are slightly
larger than the abactinal (0.24 mm.).

Small straight pedicellariae (pl. 47, figs. 3, 3a-3d) are found chiefly on mouth
and proximal adambulacral spines, very sparsely in interradial, intermarginal, and
actinal channels and just above the superomarginal plates; small pedunculate ones
(0.2 to 0.4 mm.) are scattered along furrow edge of adambulacrals. They are fairly
broadly triangular in contour, compressed, and the tips of the jaws may be somewhat
unguiculate. They are usually very scarce outside of the ambulacral and oral plates
and are never conspicuous. The largest (interradial) are about 0.5 mm. im full-
grown males.

Madreporic body conspicuous, surrounded by a close palisade of spinelets;
external margin at middle of r; striae fine, branching, or interrupted.

Color in life-—Large specimens are usually olive green, either uniformly dark, or
dark mottled with paler green; actinal surface whitish, ashy, or pale pinkish gray;
tube feet very pale translucent gray green. Small specimens may be dull rose pink
or mallow purple blotched with pomegranate purple, whitish on under side; others
are gray green mottled with darker olive. A medium-sized specimen of forma
aequalis: General color of abactinal surface rich greenish brown rather faintly marbled
with dark raw Siena and pale Hooker’s green. The ends of fluted spinelets pale
greenish yellow; pedicellariae same; papulae pale greenish to bottle green, golden at
ends; madreporic body cream color; marginal spines Naples yellow becoming more
and more greenish toward end of ray; tube feet Naples yellow.

Specimens collected under rocks at Port Renfrew, Vancouver Island, by Dr.
John C. Brown: ‘Color variable, often gray browns; some are orange with gray-
brown specks and blotches all over; some are light coral pink (like coralline algae) and
still others are bright coral red.”’

Anatomical notes—Skeleton (pl. 46, fig. 3e): In most specimens the skeleton of
the ray consists of regular longiseries of conventionally 4-lobed or somewhat lozenge-
shaped plates with the exception of the dorsolateral area where the regularity is
broken. There are essentially two dorsolateral series; that adjacent to the carinals
rather zigzag and separated from the second or outer by a fairly regular longiseries
of papular pores. The dorsolaterals are quite regular in contour. In a few specimens
examined the serial arrangement of abactinal plates was entirely broken up on the
proximal half of ray. The superomarginal plates are much the broadest of any.
The interbrachial septum consists of strong imbricating plates.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——FISHER 123
Forma AEQUALIS (Stimpson)
Plate 54, Figures 1, 2, 3, 4; Plate 55, Figure 6
Leptasterias aequalis var. compacta Verriiy, 1914, p, 130, pl. 56, fig 5.
Leplasterias aequalis var. concinna Verrity, 1914, p. 132.

The typical and prevalent California form is that with fairly compact granuliform
abactinal spinulation. There is a fairly regular carinal band of more closely set,
numerous granuliform spinelets. A semblance of regularity is given to the dorso-
lateral groups on account of the frequently regular longiseries of papular pores.
Both very large (R 55mm., pl. 54, fig. 4) and small examples of this phase are common.
A preponderance of the largest are males. The average breadth of ray is that of the
large specimen. Slightly stouter and slenderer rayed examples occur. Smaller
specimens (pl. 54, fig. 2) are likely to be stouter rayed, but not invariably so. Nor-
mally, abactinal crossed pedicellariae are not abundant. A slender-rayed variant has
a complete circle of crossed pedicellariae around each abactinal spinelet. Straight
pedicellariae of the slenderer and broader forms, as figured, occur in this typical phase.
A variant has the carinal band narrower, somewhat less regular, and the dorsolaterals
less regular. It is but a short transition to specimens which, while having granuliform
abactinal spinelets, show no evident serial arrangement of the groups of spinelets.

The type specimen (No. 1372, U.S. National Museum, Monterey) has R 20 mm.,
r 6.5 mm., and is a small example of what I have called the typical phase. It has a
subregular arrangement of the abactinal granules and three or four, rarely five,
superomarginal spinelets.

Verrill’s var. compacta and var. concinna seem to me to be only well-developed
specimens of the type form. His figure of the type of var. compacta indicates a small
specimen although it is described as “the most perfectly developed variety of
aequalis.” It is not clear wherein it differs from var. concinna since both are large,
have numerous spinelets, crowded on the plates, and the clusters are arranged in
three to five abactinal longiseries. I have examined the type of concinna. ‘‘Mon-
terey to Straits of Fuca.”

Forma NANA Verrill
Plate 55, Figures 3, 4, 5
Leptasterias aequalis var. nana, part, Verrivy, 1914, p. 132.

For ecological work it is convenient to recognize a small form which is charac-
terized by slender spinelets typically fairly evenly spaced without arrangement in
longiseries, or with this arrangement rather vaguely indicated. There are usually
only two or three slender (not short, obovate) carinal spinelets to a plate, and some
few specimens (including the largest, R 31 mm., r 8 mm.) have rather numerous
abactinal pedicellariae. In the largest specimen they form complete cirelets around
each abactinal and superomarginal spine. This specimen has four or five supero-
marginal spinelets, two inferomarginal, and one long and one very short series of
actinals. In the more characteristic small specimens, with R upwards of 17 mm.,
there are three to five slender superomarginals, two very slender inferomarginals and
one short series of slender actinals.

There are slender and stout rayed examples, the former sometimes difficult to
separate from small specimens of pusilla. Certain specimens on account of their
intermediate characters can not be placed definitely either in forma aequalis or

124 BULLETIN 76, UNITED STATE NATIONAL MUSEUM

forma nana. It seems probable that Verrill included young f. aequalis in his variety
nana.

Both formae carry eggs while still small. I have a specimen of nana with R
10 mm. which carries young nearly ready to leave (March 30, 1921). Monterey.

Type of Leptasterias aequalis.—Cat. No. 1372, U.S.N.M.

Type locality —Monterey, Calif. (A. S. Taylor).

Distribution.—San Juan Islands, Wash., and Strait of Fuca to Santa Catalina
Island, Calif.

Specimens of Leptasterias aequalis examined

| Number

Locality of speci- Collection
mens

Neah Bay, mouth of Strait of Fuca......-.-------------------------- | 5 | U.S.N.M.
Port Renfrew, British Columbia (Strait of Fuca) -----.-.------------ 7 | Stanford, J. C. Brown.
Friday Harbor, San Juan Islands, Wash -----.---.--------------- see 9 | Stanford, F. W. Weymouth.
OrescentnG ley GAN one se neater ae een eee ee | 22 | Stanford, C. L. Hubbs.
Fort Bragg, Mendocino County, Calif........--..--.---------------- | 2 | Stanford, R. P. Hays.
Beara ROreATICISCOO SES, CNL el ee ee | 1 | U.S.N.M., John Hornung.
Mionterey BBY, @HUl 92 - = oo semen ee | Many.| Stanford, W.K. Fisher.

Oayueas, Callfitce > tet -SEeE 2 oo Ee aes eae a nate man 8 | Stanford, Myrtle Johnson.
Seariger orties Va barighs QU tet ln re ee 1| U.S.N.M. H. W. Henshaw.
Santa Catalina Island, Calif__-_- eee | 2) U.S.N.M.

Remarks.—It is questionable if typical large aequalis reaches the Puget Sound
region. The Neah Bay specimens (R 21 to 25 mm.) appear to be a variety of forma
aequalis. The specimens from Friday Harbor have the smaller and more granuli-
form spinelets of aequalis as well as the peculiarities of the crossed pedicellariae but
fewer abactinal spinelets than typical Monterey specimens of the same size (R 21
to 35 mm.) This lot of specimens is easily separable from vancouveri, yet any
specimen from the San Jaun Islands must be viewed with suspicion and allocated
with extreme caution. Something connected with environment plays havoc with
specific boundaries.

It is possible that ZL. aequalis intergrades with L. hexactis forma plena, and °
thence through siderea with L. camtschatica dispar forma nitida.

LEPTASTERIAS ALASKENSIS (Verrill) emended
Plate 48, Figures 2, 3, 5; Plates 56, 57, Figures 1, 2

Asterias epichlora var. alaskensis VERRILL, Amer. Nat., vol. 43, 1909, p. 549.

Leplasterias epichlora VeRRILL (not Brandt), part, Shallow-water Starfishes, 1914, p. 132.

Leptasterias epichlora alaskensis VerRRiuL, Shallow-water Starfishes, 1914, p. 136, pl. 28.
fig. 1, 2; pl. 85, fig. 1-1d.

Diagnosis.—Rays six. Differiag from L. camtschatica in the irregular arrange-
ment of the abactinal spines, usually in a reticular pattern, without indication of
longiseries; in the position of the superomarginals low on side of ray; in the feeble
development of actinal plates which form only one series extending to middle of
ray, or less; in the presence of large bivalve ‘‘stone-hammer” pedicellariae on the
lateral and actinal portions of ray. R 55 mm., r 15 mm., R= 3.7 1; breadth of
ray at base, 16mm. (Unalaska).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 125

Verrill believed, erroneously, that this species commonly includes 5-rayed
examples. He apparently employed alaskensis in a curious way to cover the 6-rayed
specimens while epichlora was used in a rather vague, comprehensive sense, for all
the varieties combined. The type of alaskensis is from Unalaska (Dutch Harbor)
but he cites localities from Puget Sound to the Aleutian Islands. I have never
seen a 5-rayed specimen and I believe such are extremely rare in the adult state,
since I have examined very many more specimens than were available to Verrill.
Some of his supposed 5-rayed specimens are forms of young Evasterias troschelii
(for example, variety subnodulosa).

I have restricted the name alaskensis to that portion of the species (of which
Verrill’s type is an example) which ranges from Attu Island to Cooks Inlet and
probably a little further south. The range of this race corresponds closely to that
of L. camtschatica dispar.

Verrill writes that the species has a complete series of peractinal ossicles each
bearing one large spine and that in large specimens a short second row is found at
the base of the rays. This is not a fact. One means of distinguishing alaskensis,
its races and formae, from L. camtschatica and allies is by the feeble development
of its actinal plates and spines, which never form a complete series and only in large
. specimens reach the middle of the ray. A second row is never present. The absence
of a real actinal area allows the marginal plates to drop low on the side of the ray,
while in camtschatica, dispar, and allies they are rather high on the lateral face of ray.

Forma ALASKENSIS (Verrill) emended
Plate 48, Figures 2, 2a-2f; Plate 56

The following description is of the type forma and its major variations.

Description of forma alaskensis.—Spinelets numerous, irregularly arranged in
single lines surrounding the papular areas, which are unequal in size and without
longiserial alignment except immediately above the superomarginal plates. Spine-
lets unequal, small but robust, usually quite definitely subcapitate, or capitate with
a rounded or truncate, striated extremity. Those along the carinal line are likely to
be more closely placed, and either subequal to the others or a trifle larger. Dorso-
lateral region, between the carinal and superomarginal plates, well filled in with
spines which are rather evenly graded, as to length, with the superomarginals.
There is a distinct tendency in specimens from Unalaska and westward for the carinal
and a variable number of dorsolateral spines to form groups of two, three, four, and
sometimes five, which, by reason of their slightly more prominent plates and superior
size, give an uneven surface to the abactinal area (somewhat as occurs, in a more
exaggerated way, in L. polaris acervata).

Lower on side of ray than in L. camtschatica is a well marked supramarginal
channel and a regular double series of cylindrical, often subcapitate, round-tipped or
subtrunecate, but occasionally slightly tapered, superomarginal spines generally a
little longer and thicker than the abactinals, or even very decidedly so. Sometimes
there are three spines, but never regularly.

Below these is the regular intermarginal channel, followed by a double row of
inferomarginal spines longer and heavier than the superomarginals, and in large

63160—30——9

126 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

specimens there is a single actinal series extending less than half the length of the ray.
These actinolateral spines vary from 50 to 100 per cent longer and thicker than the
superomarginals, are quite variable in form, but are generally swollen and tapered
to a blunt tip; or subcylindrical; or a trifle curved; or many of the spines are com-
pressed, of even width; or else expanded toward the tip and subtruncate (Adakh).

Adambulacral spines alternate typically one and two, fairly regularly. They
are much slenderer and slightly shorter than the inferomarginals, and the furrow
spine of diplacanthid plates is decidedly slenderer and more tapered than the others,
which vary considerably, form a cylindrical to a terete or compressed-clavate round-
tipped form. Adoral carina composed of two or three pairs of contiguous plates.

Actinostome small; mouth plates small, with three spines each—a fairly long,
tapered suboral, and two apicals, of which the inner is a little longer than the median
oral suture; and the outer spinelet one-half or two-thirds as long; occasionally sub-
equal. Actinostomial margin of combined mouth plates slightly broader than the
opposite margin adjacent to first adambulacrals.

Papulae in alcoholic specimens are conspicuous and appear everywhere, irregu-
larly, among the spines and pedicellariae. The supramarginal, intermarginal, and
proximally one actinal series of areas are regular. The dorsal areas are so variable
in size and the number of papulae per area increases so with age that it isnot possible |
to state any precise number of papulae. Anywhere from 3 to 12 occur in the dorsal
areas of good-sized specimens, 3 to 6 in the intermarginal, and usually only 1 to 3
in the actinal.

The straight pedicellariae (pl. 48, figs. 2, 2a) afford one of the readiest means of
identifying specimens. Characteristic bivalved pedicellariae, having subrectangular
jaws a little longer than broad or slightly broader than long, of variable but large
size, with the edge of the jaw denticulat e, occur, sometimes abundantly, on the dorso-
lateral region of the ray, intermarginally, and on the actinal surface, especially in
the interradial region. Sometimes the jaws are shaped like a miniature pecten shell.
These pedicellariae, when well developed, have jaws 1 mm. broad, which exceeds the
thickness of the abactinal spines, and even that of the inferomarginal spines, unless
these are exceptionally heavy. The bivalved pedicellariae are by no means uniform
in size. The largest are found at the base of the ray in the intermarginal and supra-
marginal channels, and actinally.

Smaller ovate and lanceolate straight pedicellariae of divers sizes occur along the
furrow margin, on the oral spines, and in the clusters of crossed pedicellariae on the
furrow spines of diplacanthid adambulacral plates.

Crossed pedicellariae (pl. 48, figs. 2c, 2d) are usually only moderately abundant
abactinally. They are spaced, sometimes widely, around the base of the abactinal
and superomarginal spines and form clusters on the outer side of the inferomarginal
and adambulacral spines. The abactinal measure 0.225 to 0.24 mm. long.

Skeleton: The structural features differ from the arrangement in L. camtschatica
as follows: Abactinal area much broader, the superomarginals being actinolateral in
position; carinal series extremely irregular, sometimes not marked off from other
plates; dorsolateral skeleton more open, very irregular, and plates very irregular in
contour; marginal plates rather smaller with usually more attenuate lobes; actinal

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 127

plates small, reaching to middle of ray; a few rudimentary spinelses plates may ex-
tend a short distance beyond middle. (PI. 48, fig. 2/.)

Madreporic body situated at the middle of r, conspicuous, slightly convex, with
rather fine branched radiating striae. It is encireled by a variable number of
spinelets.

Variations.—Forma alaskensis: The typical form which has just been described
is rather variable, especially in the western part of its range. A comparison of
specimens from a series of localities, extending from Kodiak westward to Attu,
reveals a gradual change in addition to local peculiarities.

Western examples gradually become larger and there is a tendency for the
major abactinal spines to form aggregates of two to five, to become incipiently
“acervate,”’ so that the abactinal surface is uneven, not rather even as is the con-
dition in Kodiak and most Unalaskan examples. The bivalved pedicellariae, which
at Kodiak are fairly well developed, become larger and heavier, and reach their best
development from Unalaska to Adakh. To the westward of Adakh they decrease
in numbers and in breadth. Attu and Agattu examples in this character are inter-
mediate with asiatica.

Kodiak Island: The specimens in respect to straight pedicellariae are inter-
mediate between L. alaskensis multispina and L. alaskensis alaskensis. The lateral
bivalved pedicellariae are distinctly smaller than in examples from the Aleutian
Islands. Karluk examples, on the whole, have more spinelets, the superomarginals
carrying two or three spinelets. (PI. 56, fig. 2.) These specimens favor multispina.
Some of them resemble pribilofensis but have narrower straight pedicellariae. St.
Paul examples have fewer, more widely spaced spinelets, in common with others
from Seldovia, Port Graham, Kukak Bay, and the Shumagin Islands. All these
have weaker spinelets than typical Unalaskan examples, and usually only one supero-
marginal spinelet, but here and there two. The straight pedicellariae are smaller than
normal. They intergrade freely with typical alaskensis. (See forma shumaginensis.)

Unalaska: Type locality. At Unalaska the specimens frequently have the
larger abactinal spines in groups of two to four or even five, especially in the carinal
series. The pedicellariae are well developed, with wide jaws and have a rounded
often irregularly denticulate distal margin.

Atka: The specimens are characterized by the prominence of small groups of
scattered abactinal spinelets and by rather greater disparity between the largest and
smallest dorsal spinelets. There is considerable variation in the number of bivalved
pedicellariae. The proximal inferomarginal plates usually carry only one spinelet,
the distal two. The actinal series of spinelets is absent or quite short. In com-
parison with an average example from Kodiak, the spinulation is sparser, much
coarser, and much less uniform.

Adakh: The specimens from Adakh, another island of the Andreanof group,
resemble those from Atka, but there are also seven specimens taken July 5, 1893,
which have the abactinal spinelets almost exactly typical. The largest has R 82 mm.
Like the others, it has a typical number of marginal spinelets and the actinal series
extends half the length of the ray. All the spinelets are quite stout, but the marginal
and ventral especially so, these being heavy, clavate, or even clavate-spatulate.
The bivalved pedicellariae are numerous and large, the jaws being very often decidedly

128 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

broader than high, especially in those of the supramarginal and intermarginal chan-
nels. The distal margin is finely denticulate and is often slightly produced in the
middle. This is a prolongation of a low median ridge or carina which is sometimes
found on the outer face of the valve.

Amchitka Island: This is one of the Rat Islands between the Andreanof Group
and the Near Islands (Attu and Agattu). The smaller of the five specimens resemble
forma shumaginensis. The largest (R 38 mm.) has a more even surface than the
Atka variety and also resembles the sparser spined intermediate specimens from Attu
and Agattu. The bivalved pedicellariae are mostly smaller than typical with jaws
usually higher than wide on outer half of ray.

Attu and Agattu: The specimens from Agatiu (pl. 56, fig. 3), the largest of
which has R 65 mm., r 18.5 mm., are rather more uniform in appearance than those
from Attu (fig. 4). They represent a recognizable subrace of alaskensis notable for
the robust abactinal spinelets which are capitate, subtruncate, and do not define the
papular areas in reticulate pattern, but tend to form groups which along the carinal
line are crowded and sometimes a trifle elevated (especially in young specimens).
These groups occur also on the dorsolateral region but with a few exceptions are not
so well defined. The straight pedicellariae are about intermediate between those of
alaskensis and those of asiatica.

Numerous young specimens from Agattu (R 6 mm. to 25 mm.) have the carinal
groups of two or three spinelets relatively more prominent than in large examples,
while on the dorsolateral area the spinelets, of two or three sizes, are rather widely
scattered. The lateral pedicellariae are thick-ovate with straight-sided or slightly
tapered jaws, not greatly unlike the larger sort found in asiatica.

The specimens from Attu (pl. 56, fig. 4) agree with those of Atka in having
considerable disparity in size between major and minor abactinal spinelets, and in
having the former in prominent, though small, groups which give a very uneven
appearance to the abactinal surface. This is simply an exaggeration of the condition
in Agattu examples. Four specimens are practically the same as those from Atka.
The others have rather coarser spinelets. The bivalved lateral pedicellariae are rather
larger than in the Agattu examples, but the jaws are not so wide as in Adakh speci-
mens, that is, are not wider than high.

There seems to be in this region (the Near Islands) a fairly definite subrace
intermediate between alaskensis and asiatica, as exemplified by the majority of
specimens from Agattu and some of those from Attu. Then there are (especially
at Attu) a number of variants which may be the result of crossing of alaskensis with
dispar. One evidence of this is the instability in the size of major pedicellariae;
another the formation of prominent groups of carina! spinelets and the dwindling in
size and number of the dorsolateral spinelets.

Forma SHUMAGINENSIS, new forma
Plate 48, Figure 3, 3a; Plate 57, Figure 1

These specimens differ from large, well-developed Unalaskan examples in having
slenderer, cylindrical or slightly tapered spinelets which are only occasionally sub-
capitate. They stand not very close together on the skeletal ridges and do not form

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—-FISHER 129

a clearly evident reticulate design. Rather they appear to stand irregularly spaced;
carinal series not at all or only feebly differentiated; no acervate groups. Bivalved
pedicellariae relatively narrow.

Superomarginal spinelets usually one to a plate, slightly tapered, blunt, subequal
to the dorsolaterals to 50 per cent larger; spinelets well spaced, low on side of ray, much
slenderer than in typical alaskensis. Inferomarginals proximally one or two, distally
irregularly one or two or regularly two, conspicuously larger than superomarginals,
typically tapered, blunt (not clavate). Actinal spines either absent or represented
by a very short series, similar to inferomarginals. The small plates extend scarcely
a third length of ray. Intermarginal channel proximally fairly broad.

Bivalved pedicellariae rather variable but with the jaws considerably narrower
than high, and smaller than in typical alaskensis. In a relatively few pedicellariae
the jaws are nearly as broad as high and untapered when viewed from the outer
face but in the majority the jaws are tapered. (PI. 48, figs. 3, 3a.)

Adambulacral spines alternately one and two; or at base of ray irregularly one
and two, distally one; or less often one throughout ray (Sanborn Harbor, Nagai).
Two pairs of postoral adambulacrals in contact. Oral spines three, as in alaskensis,
but not quite so long nor so broad.

Some of the specimens from Humboldt Bay have a much weaker than normal
abactinal skeleton, with relatively slender ossicles and large meshes four or five be-
tween opposite superomarginal plates.

The rays vary in caliber from slender to medium stout. Largest example R
40 mm., r 11 mm., R=3.6r.

The variety from Seldovia, Cook’ Inlet, was carrying eggs April 9, 1892.

Type of forma shumaginensis.—Cat. No. E 1500, U.S.N.M.

Type locality Humboldt Bay, Shumagin Islands.

Distribution —Cooks Inlet to Shumagin Islands.

Remarks.—This is a small race or form which is found in the region between
Unimak Island (where nearly typical alaskensis occurs) and Cooks Inlet. It inter-
grades with a variety of forma alaskensis on Kodiak, and probably elsewhere. It is
none too homogeneous. The characteristic smaller spinelets, mostly monacanthid
superomarginals, and smaller straight pedicellariae are best exhibited by the Shu-
magin Island specimens.

Forma PRIBILOFENSIS, new forma

Plate 48, Figures 5, 5a; Plate 57, Figure 2

This is a medium-sized rather slender-rayed variety with R averaging about 40
mm. and very numerous, uniform abactinal spinelets compactly placed and defining
small, numerous papular areas. Abactinal surface evenly arched, with a regular, not
lumpy surface. The spinelets are generally much more delicate than in f. alaskensis
and very definitely form single or double palisades on the ridges of the skeleton,
defining a very evident reticulate pattern on the abactinal surface. Although the
spinelets are often crowded along the carinal line they do not form convex groups.
This form resembles the common variety of alaskensis from Kodiak but the rays are
slenderer and longer and the bivalved pedicellariae larger.

130 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Among the large series there are however some short, stout-rayed examples, not
differing from the above in spine characters. These have broader pedicellariae than
both shumaginensis and the small-spined intergrades with multispina which are found
at Karluk, Kodiak Island.

The abactinal spinelets are short, broadening toward the striate usually sub-
truncate tip, which is usually subcapitate and provided with numerous minute
points. The superomarginal spines, two or three in a vertical series to each plate,
are only a little larger than the adjacent dorsolaterals, but the inferomarginals, two |
to a plate are decidedly longer and heavier, and are generally cylindrical or clavate,
subtruncate. There is a single short series of similar actinal spines, these with the
inferomarginals forming transverse series of three at base of ray. They are from 1.3
to 2 mm. long, while the superomarginals are 0.6 to 0.8 mm., and the abactinal about
0.5 mm. long. The adambulacral spines alternate one and two.

The lateral, dorsolateral, and actinal ‘‘dermal”’ straight pedicellariae are of the
characteristic broad jawed, bivalve, goniasterid sort (pl. 48, fig. 5a) generally thicker
than the adjacent spines. The largest are in the supramarginal and intermarginal
channels; but smaller ones occur among the inferomarginal spines. Some of these
may have narrower jaws.

Color in life-—Dr. G. Dallas Hanna, who collected a large series at Village Cliffs,
Village Point, and Zapadni Reef Rookery, St. Paul Island, made careful color notes
and recorded 25 variations, some of them minor. The following are the principal
color phases:

1. Upper surface very dark green, borders of papular areas dark brown; actinal
surface and tube feet light transparent green, the margins flesh color (15 specimens).

a. Margins pink instead of flesh color (one specimen).

b. Margin of ventral surface lighter pink, and tube feet flesh color.(two speci-
mens).

c. With small bright purple spots uniformly distributed on upper surface; flesh
color below, margins translucent green (one specimen).

2. Similar to 1 but upper and lower surfaces slightly lighter (10 specimens).

a. Gray muddy green above, flesh color below (2 specimens; see 12).

3. Upper surface very dark brown, papular areas black; tube feet light dirty
green; furrow bordered by flesh color (14 specimens).

a. Lighter brown above, papular areas almost black (12 specimens).

b. Upper surface still lighter brown, papular areas black, tube feet dark green,
lighter green, or flesh color (four specimens).

c. Dark brown all over (one specimen).

4. Upper surface dark buff; papular areas small, jet black; tube feet dirty green
bordered by flesh color (one specimen).

a. Lighter buff above; actinal surface all flesh color (two specimens).

5. Upper surface purple, sides brown; actinal surface flesh color (one specimen).

6. Upper surface brick red all over; tube feet light dirty green bordered by pink-
ish flesh color (two specimens).

7. Upper surface light red with a few irregular blotches of blood red (one
specimen).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 131

8. Upper surface lavender (one specimen).

a. With a few little spots of dark red (one specimen).

9. Upper surface maroon (two specimens).

10. Upper surface very dark purple; under surface flesh color (one specimen).

11. Upper surface very dark blood red with dirty blotches (one specimen).

12. Light gray above, with dark spots which become merged into a large area
in the center of the disk; flesh color below (four specimens).

Type of forma pribilofensis—Cat. No. E 1501, U.S.N.M.

Type locality of forma pribilofensis—St. Paul, Pribilof Islands, rocks, low tide.

Type of Leptasterias alaskensis—Not present in Yale Museum collection, 1927.

Type locality of Leptasterias alaskensis—Dutch Harbor, Unalaska (Harriman
expedition). The type locality is not designated in the text, but on page 172,
opposite Plate 85, is said to be Dutch Harbor.

Distribution —Attu and Agattu (more or less intermediate with asiatica) to
Kodiak Island and Cooks Inlet, Alaska, intertidal to six fathoms.

Specimens examined.—Four hundred and twenty.

Specimens of Leptasterias alaskensis examined

Locality Collection
MARION, RUM 52 hidden ti cadad Soke bienacndcawess Acad, Sci. U. R. S. 8., Wosnessensky, 1858.
PEIN OD ic sd steal atl yt he wieistee tol Stanford, C, Rutter.
Ree IE! RON etalk remtniedpiacvelenduncanedin|eu bligae ....--.-| Stanford, C. Rutter; 8. P. Smith.
Seldovia, Cooks Inlet........... ..| U. 8. N. M., Albatross, 1992.
Port Graham, Cooks Inlet......... .| U. 8. N. M., 1872.
Chignik Lagoon, Alaskan Peninsula. | U.S N. M.
Kukak Bay, Shelikof Strait.............. .| Stanford.
Unga, Shumagin Isiands.........................--|..--.--- | U.S. N. M., W. H. Dall.
Humboldt Bay, Shumagin Islands. ................ U. 8. N. M., Albatross.
Nagal, Shumagin Islands........................... 0.
Unimak Island, Alaska } , C. H.
St. Paul, Pribilof Islands. {

LEPTASTERIAS ALASKENSIS ASIATICA, now subspecies
Plate 48, Figures 1, la-ld; Plate 57, Figures 3, 4, 5

Diagnosis.—Similar to L. alaskensis but the broad bivalved pedicellariae of that
form represented by narrower-jawed ovoid ones. Abactinal spines numerous, of
rather uniform length, robust, subcapitate, arranged in a recticular pattern. R 55
mm.,rl5imm. R=3.7 r; breadth of ray at base, 15 mm.

Description.—The spines are robust, the superomarginal and abactinal being
capitate and subtruncate, with striate tips. The abactinal spines are generally of
uniform, or nearly uniform length, though of different sizes and rather compactly
placed, outlining the small papular areas and forming an irregular reticulate pattern.

132 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The carinal spines are not typically larger than the dorsolaterals although more
numerous to a plate, and in some but not all specimens they form an irregular band |
due to greater crowding. The spines do not form conspicuous, often elevated groups
as in Atka and Adakh, and most of Attu and Agattu examples of alaskensis. There
are typically two superomarginal spines, two inferomarginals (on the first three or
four plates, one) and a short series of actinals (scarcely reaching the middle of R).
The first three or four superomarginals may have three spines, or only one; the distals
sometimes have three. The inferomarginals are heavier than the superomarginals,
sometimes terete, sometimes slightly clavate. On the outer part of the ray the
lower spine is usually heavier than ths upper, or the upper two, for three occasionally
occur.

Adambulacral spines one and two, in fairly regular alternation, and very similar
to those of alaskensis. Three adoral pairs of adambulacral plates are in contact
interradially. The oral plates are also similar to those of alaskensis and the outer
of the two apical marginal spines is frequently absent.

The straight pedicellariae (pl. 48, figs. 1, la-1d) are of the common compressed
ovate form, and are smaller than the broadly wedge-shaped ones of alaskensis. In
some pedicellariae the jaws, as seen from the back or outer side, taper narrowly to a
blunt or acute point (fig. 1@) while in others (usually near the base of ray or in axillary
channel) the jaws are untapered, sometimes slightly constricted at the middle, and
end in two to four teeth (fig. 1b). These are more common in Medni examples, and
are intermediate with the pedicellariae of alaskensis. Small ovate and lanceolate
pedicellariae occur in clusters on the oral spines, among the crossed pedicellariae of
the furrow spines and on the furrow face of the adambulacrals. The latter are the
smallest of all.

Crossed pedicellariae distributed as in alaskensis. They vary considerably
in number.

Madreporic body as in alaskensis, surrounded by a complete circle of spinelets.

Variations.—It is to be noted that the straight pedicellariae of Medni examples
are more variable than are those of Bering Island specimens. Abactinal straight
pedicellariae are very scarce outside of the supramarginal channel, and the number
of marginal and actinal ones is quite variable. In the arrangement of the abactinal
spines there is much less variation than in specimens of alaskensis from Attu and
Agattu. The abactinal surface of asiatica resembles that of some stout-spined
examples of alaskensis from Kodiak in which the pedicellariae are also narrower
than in typical alaskensis from Unalaska westward. The largest specimen, from
Medni, has R 62 mm. There are five specimens from Simushir, Kuril Islands, which
constitute a distinguishable form (pl. 57, fig. 5), differing from typical asiatica in
smaller size, fewer and relatively larger abactinal spines, which do not clearly define
papular areas but appear scattered; and in the lateral spatulate pedicellariae of
conspicuous size which occur along with ovate conical ones. This may be similar
to the ordinary Kamchatkan form, of which there are no specimens.

Type.—Cat. No. E 1502, U.S.N.M.

Type-locality—Nikolski, Bering Island, shore rocks, June 15, 1906.

Distribution—Known from Simushir (Kuril Islands), Bering Island, and Medni
Island (Commander Islands).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 133

Specimens exramined.—Sixty-six.

Bering Island (Nikolski), shore rocks, 29 specimens, June 15, 1906, Albatross.

Bering Island (Nikolski), 5 to 10 fathoms, stones, algae, 1 specimen, August L5
1879, Vega expedition.

Medni Island, shore rocks, 31 specimens, June 13, 1906, Albatross.

Simushir (Milne Bay), Kuril Islands, 5 specimens, June 23, 1906, Albatross.

LEPTASTERIAS ALASKENSIS MULTISPIN A, new subspecies
Plate 48, Figure 4; Plate 57, Figures 6, 7

Leptasterias epichlora alaskensis Vernit1, part, 1914, p. 136.

Diagposis.—Differing from L. alaskensis f. alaskensis of the Aleutian region in
having slenderer rays, slenderer spinelets, and notably smaller bivalved straight
pedicellariae. Abactinal spinelets unequal; quite slender ones associated with stouter,
subclavate, scarcely subcapitate spinelets arranged in a close irregular reticular
pattern or without definite arrangement; carinal spinelets not differentiated. Super-
omarginal spines, usually two, or one at base of ray, typically slender; inferomar-
ginals usually two (some times one); actinals small, sometimes wanting and never
extending beyond middle of ray. R 45 mm., r 10 mm.; breadth of ray at base,
10 or 11 mm.

This race is a southern extension of alaskensis, which reaches its eastern limit
at Kodiak Island and Cooks Inlet. Multispina shows its relationship by the numbers
and arrangement of the abactinal spinelets (which however are weaker) and by what
might be called the relics of the characteristic bivalved pedicellariae. The position
of the superomarginal plates rather low on the side of ray and the reduction of the
actinals to a short series of rather inconstant development indicate a close relation-
ship with alaskensis.

Description —Abactinal surface well arched so that ray appears cylindrical with
a flattened, narrow, actinal surface. Abactinal spinelets, without any regular
arrangement, are very numerous, fairly slender, rather uniform in length but unequal
in caliber, cylindrical, tapered, or slightly clavate but not normally capitate. Since
they stand on a close irregular reticulate skeleton they show a similar pattern unless
interrupted by the absence of a spine here and there. In some specimens the spine-
lets are shorter and of more uniform thickness.

Superomarginals generally two, or irregularly one proximally; occasionally three,
slightly tapered, blunt, only a little longer than adjacent abactinals. Supramarginal
channel fairly broad proximally. Inferomarginals conspicuously longer, especially
the lower of the usually two spinelets; sometimes three spinelets are present forming
a triangular group.

The adambulacral spinelets normally alternate fairly regularly one and two,
as is so commonly the case in this genus. The spinelets are nearly as long as the
inferomarginals but much slenderer, terete, slightly tapered, blunt, while the furrow
spine of the diplacanthid plates is shorter, slenderer, and usually a little more tapered
then the outer. The posture of the spines frequently is such that there appears to
be three series along the furrow margin. The first three pairs of adoral adambulacral
plates are in contact interradially while those of the fourth pair are very close together.

63160—30——10

134 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The first six plates usually have but one spine. The first plate is longer than the
succeeding plates.

The actinostome is small and sunken as is usually the case when the adoral
carina is narrow. The mouth plates, at the sutural margin adjacent to the first
adambulacral plate, are produced into a sort of flange so that the combined pair
is here conspicuously broader than the first pair of adambulacral plates. This char-
acter is less pronounced in some varieties. Each mouth plate has three spines, a
very short one close to the actinostome, and usually bent over the beginning of the
radial nerve; a second, about 50 per cent longer, stands above it (as viewed from
below); and a suboral, similar to the first adambulacral spine, near the outer margin
of the plate.

The abactinal papular areas, except for a fairly regular series adjacemt to the
superomarginals, are quite irregular in form and arrangement, and variable in num-
ber and size. Probably three or four papulae to an area would be an average. In
the intermarginal areas there are three or four proximally, and in the actinal, one
to three. The numbers vary so with age that they afford no taxonomic ammunition.

Crossed pedicellariae occur rather sparsely on the sheaths of the abactinal spines
and in dried specimens at the base of the spines; in rather scanty wreaths around
the base of the superomarginal spines, and as clusters or half wreaths on the outer
side of the inferomarginal and adambulacral spines.

Straight pedicellariae: The large lateral and actinal straight pedicellariae afford
the easiest means of distinguishing multispina from alaskensis. Good-sized ovate
pedicellariae are present in the supramarginal, intermarginal, and actinal inter-
brachial channels; and also sometimes in the actinal channel, and less often on a few
of the proximal inferomarginal and actinal spines. Smaller ones of the same sort
are sometimes sparsely scattered on the dorsolateral plates. All are broadly ovate
as seen from the side, but the back of each jaw is narrower than the height, and tapers
slightly. The breadth increases gradually in specimens from more northern localities,
until in typical alaskensis from Unalaska and the Aleutian Islands, the jaw is broader
than high in well developed specimens, and the whole pedicellaria is relatively larger
and heavier than in southern examples. (PI. 48, fig. 4.) The pedicellariae have a
few denticulations at the jaw tip. Other smaller, broadly lanceolate, ovate, or
subtriangular pedicellariae are present on the furrow margin, furrow spines (along
with crossed pedicellariae), and oral spines.

Type—Cat. No. E 1503, U.S.N.M.

Type locality —Wrangell, Alaska.

Distribution —Southeast Alaska to northern Vancouver Island.

Specimens eramined.—Seventeen.

Specimens of Leptasterias alaskensis multispina examined

‘ Number
Locality of speci- Collection
mens
Southern AIAG e ees sno tan cccut rete eC eeE eo Ree ee 6 | U.S.N.M., Dr. W. H. Jones, U. S. N.
WIAnOl A LOBNA ee seen ora took: Secon eee wee es et 8 | U.S.N.M., Dr. W. H. Jones, U. 8. N.
Chasina Bay, Prince of Wales Island, Alaska........_...____________ 1) U.S.N.M., T. H. Streets.
Alert Bay, Vancouver Island, British Columbia......-____.......___ 2\ U.S.N.M., Dr. W. H. Jones, U. S. N., February,

1882.
—— te ee I Le ld tee tt el

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS--FISHER 135

Remarks. — Multispina intergrades with alaskensis in the region of Kodiak and
the Shumagin Islands. It resembles forma shumaginensis, which, in a sense, is an
intermediate stage between alaskensis and multispina, perhaps if anything a little
nearer the latter. Multispina has more numerous abactinal spinelets and rather
smaller lateral straight pedicellariae. (Compare pl. 57, figs. 1 and 6.)

Subgenus NESASTERIAS, new

Diagnosis. —Differing from Leptasterias s. s. in the form and small size of the
crossed pedicellariae which have a very broad terminal lip; in the thick, distally
broadly expanded, circumspinal sheaths; in the reduction of the adoral carina to one
pair of adambulacral plates in partial contact interradially; in the incipient degenera-
tion of the dorsal skeleton (dorsolateral skeleton a single series of ossicles usually
not in connection with carinals). Gonads opening ventrally; actinal plates in 0 to 2
incomplete series.

Type—Leptasterias stolacantha Fisher.

Remarks.—The curious small sea stars upon which this subgenus is based are
readily recognizable by the very large obconic sheaths which surround the spines and
which carry on the distal convex end numerous diminutive crossed pedicellariae.
The latter are of a form which differs from that running pretty constantly through
the genus Leptasterias, as exemplified by groenlandica, hyperborea, arctica, miilleri,
littoralis. The sheaths remind one of those found in Coronaster, Rathbunaster,
Pycnopodia, and Lysastrosoma, and are relatively much fleshier than the sheaths of
L. hyperborea or of L. tenera forma compta.

The dorsal skeleton is beginning to disintegrate, the connectives between the
single dorsolateral series and the carinals, as well as between the consecutive carinals,
having largely disappeared. The abactinal integument is therefore very soft and
flexible.

The adoral carina is as poorly developed as in some species of the Pedicellasterinae.
Only the adcentral, or inner, two-thirds or three-fourths of the interradial margin of
the first adambulacral plates are in contact. The species is therefore intermediate
so far as this character is concerned, between the Asteriinae and Pedicellasterinae.
It occupies somewhat the same position in the Asteriinae as Tarsaster and Ampheraster
in the Pedicellasterinae. Its position in the Asteriinae is indicated by the skeletal
structure and the presence of pedicellariae on the adambulacral spines, while the
structure of the gonads suggests an alliance with Leptasterias. The distinct form
of the crossed pedicellariae, however, argues for only a distant connection with
typical species of Leptasterias. The group is not so close to typical Leptasterias as
is the camtschatica-polaris section.

LEPTASTERIAS STOLACANTHA, new species
Platel3; Plate 18, Figures 4, 5

Diagnosis.—Rays five, R 32 mm., r 5.6 mm., R=5.8 r; breadth of ray just
beyond base, 9 mm.; rays sharply constricted at base, arched, gradually tapered
to a blunt point; disk small, slightly convex; ray uniformly beset with delicate,
slender, tapered spinelets surrounded by a relatively very broad ruff of small crossed

136 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

pedicellariae carried on the expanded summit of a retractile sheath which covers
all but the tip of spinelet; spinelets and sheaths on disk smaller than those of rays;
serial arrangement not very evident except on sides of ray; actinal plates in 0 to 2
incomplete series; adambulacral spines two, the inner with a group of pedicellariae,
the outer only sporadically; only one pair of postoral adambulacrals in contact;
crossed pedicellariae very small, with an unusually broad denticulate terminal lip;
straight pedicellariae small, ovate, or lanceolate, compressed.

Description—Rays abactinally well rounded, and disk sometimes slightly
higher than base of rays. Spines very delicate, slender, tapered, terete, bluntly
pointed, rather widely spaced, each invested by a thick, distally widely expanded,
sheath the convex distal surface of which bears very many crowded, small, crossed
pedicellariae of characteristic form. These sheaths usually cover all but the tip
of the spine, and sometimes extend beyond the tip. In alcoholic specimens the
expanded terminal portion is subcircular or irregular in outline as viewed from the
end of the spine, 1 to 1.5 mm. broad, and resembles a furry ruff or stole. The spines
are all very similar, the abactinal being a little shorter than the marginal and in
less evident longiseries. Regular serial arrangement is theoretically present owing
to the arrangement of the plates, but some of the carinals, dorsolaterals, and supero-
marginals carry two spines, or even three, in the transverse series on the plate.
As shown in Plate 18, Figure 5, this breaks up the regularity of the longiseries of
spines. The space between the spines varies greatly. Sometimes the stoles nearly
touch; sometimes they are widely spaced. In gravid specimens there is usually
a broad naked area on either side of the carinal series (which, in such cases, is likely
to be fairly regular). The abactinal spines (which are roughened on the distal
portion) are 1.5 to 1.8 mm. long.

The skeleton (pl. 13, fig. 1) is characterized by a partial degeneration of some of
the abactinal connectives, by which the carinal plates are more or less (and irregu-
larly) isolated. There is a carinal, a dorsolateral, 2 marginal, and 0 to 2 incomplete
series of actinal plates as shown in the drawing, where also the position of the isolated
rather inconspicuous papulae are indicated. There is evidently started in this
species a process similar to that which took place in the ancestors of such diverse
genera as Lysasterias, Adelasterias, Pycnopodia, Rathbunaster. Degeneration of the
abactinal skeleton is characteristic of very cold water forms, and particularly those
of the antarctic.

The inferomarginal plates carry a single spine with a complete and large ruff
of pedicellariae. In some specimens a majority of the superomarginal plates also
carry but one spine; proximally, however, there are usually two or three. The
actinal spines are rather small and do not occur in any except the largest specimens
(R 30 mm.).

Adambulacral plates regulatly with two slender subequal spines about 1.5 mm.
long. Most of the outer spines of the proximal half of ray (and sporadically those
of distal half) bear at mid height a thick cluster or pad (sometimes a complete
wreath) of very small crossed pedicellariae similar to those of the other spines,
while a few of the inner spines carry sporadically a smaller cluster. The first plat2
is longer than those following and is in contact with its fellow along two-thirds or
three-fourths of the interradial margin, the outer portion of this margin being gen-

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-——-FISHER 137

erally free. This pair of plates is monacanthid and constitutes all there is of an
adoral carina, the second pair being widely separated interradially. A young speci-
men from Kyska with R 6 mm. has the first pair of adambulacrals separated
interradially.

Mouth plates rather narrow, each with two spines, an inner slender, blunt
actinostomial as long as, or a little shorter than the median suture and a longer
suboral similar to the first adambulacrals. This spine rarely carries a group of small
lanceolate straight pedicellariae.

Crossed pedicellariae small (0.125 to 0.135 mm. long), with very broad terminal
denticulated lip. (Pl. 13, figs. 5, 5a—5e.) They form a thick mat on the expanded
convex terminal portions of the circumspinal sheaths, upward of 8 to 10 occurring
along the radius, from spine to periphery; and they form smaller clusters on the
adambulacral spines, as detailed above.

Straight pedicellariae small, broadly lanceolate to ovate truncate, with narrow
jaws. The abactinal and lateral, scattered on the integument measure 0.25 to 0.37
mm. long (pl. 13, figs. 6, 6a. 66). Actinal pedicellariae are lanceolate with the tips
of jaws curved, sharp, and crossed; similar smaller ones occur in groups on the
furrow face of the adambulacrals and at base of inner oral spine (fig. 6c).

Madreporic body small, with few coarse striae, situated near margin of
disk.

Color in fresh alcohol whitish, or yellowish white.

Tube feet in four series, rather crowded; the pores are broadly lanceolate,
not very crowded.

Gonads very large in male; numerous lobulated divisions arise pinnately from
the axis which extends sometimes nearly to the end of ray, the longer divisions
being on the mesial side. The gonads are attached to the body-wall just above
and between first and second inferomarginals (pl. 13, fig. 1, g), the duct passing
downward to open on a papilla just exterior to the fourth adambulacral plate. The
two papillae stand close together at the actinal end of the axillary channel. (PI.
13, fig. 3, g.)

Type.—Cat. No. E. 1504, U.S.N.M.

Type locality —Station 3322, north coast of Unalaska, Aleutian Islands, 53° 28’
45’’ N., 167° 23’ 50’’ W., 35 fathoms, black sand, bottom temperature 42.4° F.,
three specimens.

Distribution.—Bering Sea, in the vicinity of the Aleutian Islands; from Unalaska
to Kyska, 43 to 59 fathoms (depth at Kyska not recorded).

Specimens examined.—In addition to the types, 12 specimens:

Station 3319, north coast of Unalaska, 53° 40’ 30’’ N., 167° 30’ W., 59 fathoms,
black sand, bottom temperature 40.8° F., six specimens.

Station 3321, vicinity type locality, 54 fathoms, dark mud, bottom temperature
41.5° F., three specimens.

Station 4777, Petrel Bank, Bering Sea, off Semisopochnoi Island, 52 to 43 fathoms,
fine gravel, one specimen.

Kyska Island (Kyska Harbor), Aleutian Islands (177° 30’ E.), W. H. Dall
(189-1015; 188-1014; 173), two specimens,

138 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Genus STENASTERIAS Verrill
Stenasterias Verritt, Shallow-water Starfishes, 1914, p. 145.—Type, Asterias macropora
Verrill.—Fisner, Ann. Mag. Nat. Hist., ser. 9, vol. 12, 1928, p. 598.

Diagnosis.—Rays six, slender. Differing from Leptasterias in having unusually
large triangular ambulacral pores separated by thin plates. Skeleton very compact,
the plates closely imbricated, in longiseries, with very small intervals; marginal
plates wide; actinals small, a short second series present; plates closely covered with
clusters of minute spinules; adambulacrals diplacanthid; reproduction unknown.

Remarks.—I have not seen a specimen of this small sea star. Verrill’s types
have disappeared. It is probable that his specimens were all young. The following
account is rearranged from the original description.

STENASTERIAS MACROPORA (Verrill)
Asterias (Leptasterias) macropora VerritL, Amer. Journ. Sci., vol. 28, 1909, p. 65, fig. 10,
p. 64. (Rays said to be five!)
Stenasterias macropora VERRILL, Shallow-water Starfishes, 1914, p. 145, pl. 50, fig. 7; pl. 74
fig. 4; pl. 84, figs. 5-5z.

Description —Disk small; rays six, slender constricted at base, convex, with
median row of more prominent ossicles. R15 mm.,r3mm.,R5r. Ambulacral
furrow unusually wide and open; tube feet and pores unusually large, the pores tri-
angular, overlapping by their acute angles and separated only by thin plates.

Abactinal plates rather closely imbricated, the spaces between being very few
and small, with only one or two papulae; carinal plates thicker and larger, subtri-
angular, with acute cusps and concave edges. These overlap the plates proximal to
them. Dorsolaterals in three longiseries, proximally, then two, subquadrate and
subtriangular in outline. (Verrill, 1914, pl. 84, fig. 54.) Dorsal spinelets numerous,
small, short, stumpy, usually truncate or slightly clavate, but not much enlarged at
tip and not much longer than thick; in small groups on the larger plates, isolated
on the smaller ones.

Inferomarginal plates > rhombic, imbricated, overlapping the outer actinals,
bearing usually two or three small tapered spines, like the actinals; two series of
very small actinal papular spaces.

A single row of somewhat quadrate, overlapping, actinal plates which bear one
or two small, rather stout, tapered spines; a series of small oblong connective ossicles
between these and the adambulacral plates, on the basal part of ray, each of which
may also bear a spine.

Adambulacral plates strong, unusually thick radially, mostly diplacanthid;
spinelets long, slender, tapered, acute or subacute, carry a few small compressed
lanceolate straight pedicellariae.

Straight pedicellariae compressed, acute, lanceolate, of rather large size but few
in number are present in actinal interradial region, and similar smaller ones along
the edges of the ambulacral grooves. Crossed pedicellariae “few and minute.”

“The specimens of this species are small and poorly preserved, having lost many
of their spines. It appears to be allied to ZL. aequalis more nearly than to other

4‘ The description is confused. The author says that the marginal plates next to the “‘ peractinals’’ (hence the inferomarginal
series) runs along the under side of the rays, but “curves upward to the dorsal side at the disk.” This is most unusual behavior for

inferomarginal plates and if true is peculiar to Stenasterias. I suspect however that superomarginals were confused with infero-
marginals at base of ray.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 139

species, but it is much more slender than that and has much larger ambulacral pores,
while the dorsal plates are closely imbricated.
“Sitka (Harriman expedition); Queen Charlotte Islands (G. M. Dawson).”’
Remarks.—Unfortunately the specimens of this species have disappeared, so that
nothing can be added to the original description. In the first notice of the species
the rays are described as five, which is probably a typographical error, as nowhere
in the definitive account is any such variation indicated.

Genus EVASTERIAS Verrill

Evasterias VeRRILt, Shallow-water Starfishes, etc., 1914, pp. 51,151. Type, Asterias troschelit
Stimpson.—Fisuer, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, p. 599.

Diagnosis.—Asteriinae differing from Asterias in having numerous actinal plates
(each bearing one or two spines) arranged in from three to six longiseries which
alternate with longiseries of large actinal papulae; inferomarginal plates lateral
rather than actinal in position. Abactinal skeleton irregularly reticulate, usually
robust, but composed of small irregular plates (the primaries three or four lobed, the
secondaries variously irregularly elliptical or elongate); spines short, stout and slender
mixed, or slender only, or robust only, terete, tapered, conical, or subglobose; adam-
bulacral plates small, diplacanthid, or irregularly diplacanthid and triplacanthid,
the spines carrying prominent clusters of crossed and straight pedicellariae; adoral
carina composed of three to five pairs of adambulacrals; gonads open dorsally, on
the disk, close to the interradial lines.

Remarks.—This genus, so far as known, is confined to the intertidal zone and
shallow water of the north Pacific from the Okhotsk Sea to central California. It
is most abundant from southern Alaska to Puget Sound, where in great inlets it vies
for dominance with Pisaster ochraceus.

In fact it bears considerable superficial resemblance to Pisaster, especially to
the numerous varieties of P. ochraceus. I have seen short-rayed examples of Evas-
terias troschelii forma alveolata which might easily be mistaken for slender-rayed
Pisaster ochraceus forma ochraceus. The resemblance concerns the dorsal spinulation
and the multiple longiseries of actinal plates. Pisaster, however, almost always has
rays which are broader at the base and evenly tapered; monacanthid adambulacral
plates; no pedicellariae on the adambulacral spines; much more sunken actinostome;
very characteristic fureate straight pedicellariae.

The modifications and distribution of the abactinal spines exhibit curious
analogies in the two genera. Corresponding forms are indicated in the following list.

Pisaster giganteus. Evasterias echinosoma.

P. ochraceus f. ochraceus. Ev. troschelii f. alveolata.

P. ochraceus f. confertus. Ev. troschelii f. acanthostoma.
P. ochraceus f. nodiferus. Ev. troschelii f. troschelii.

EVASTERIAS TROSCHELII (Stimpson)

? Asterias epichlora Branpt, Prodromus, 1835, p. 270.

Asterias epichlora Stimpson (part), Bost. Journ. Nat. Hist., vol. 6, 1857, p. 528.

Asterias troschelii Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 267—Pernrnter,
Révision des Stellérides, 1875, p. 71—Bett, Proc. Zool. Soc. London, 1881, pp. 495,
505.—Warrteaves, Trans. Roy. Soc. Canada, vol. 4, 1887, p. 116.

140 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Asterias brachiata Pprrier, not Linnarus, Révision des Stellérides, 1875, p. 65; Gulf of
Georgia.

Asterias (Diplasterias) epichlora pz Lorioz, Mém. Soc. Phys. et d’Hist. Nat. Genéve, vol.
32, 1897, p. 19, pl. 3, figs. 2, 2d.

Asterias saanichensis pr Lortot, Mém. Soc. Phys. et d’Hist. Nat. Genéve, vol. 321, p. 897,
p. 23, pl. 2, figs. 3, 83a-3d, 4, 5. Saanich Inlet, Vancouver Island.

Asterias victoriana Vurritt, Amer. Journ. Sci., vol. 28, 1909, p. 68; Victoria, British Colum-
bia. Shallow-water Starfishes, 1914, p. 124, pl. 58, fig. 1; pl. 54, figs. 1, 2; pl. 69, fig. 4;
pl. 82, figs. 1-le.

A. troschelii (St.) var. rudis Verrity, Amer. Nat., vol. 43, 1909, p. 542; Victoria, British
Columbia.

Asterias acanthostoma VERRILL, Amer. Nat., vol. 48, 1909, p. 543; no locality.

Leptasterias macowni VurRiLL, Shallow-water Starfishes, 1914, p. 124; Departure Bay,
British Columbia.

Leptasterias (?) inequalis VerRitu, Shallow-water Starfishes, 1914, p. 117, pl. 78, fig. 2; text,
figs. 4, 5 (probably very young specimen, Orca, Prince William Sound).

Leptasterias epichlora miliaris var. subnodulosa VeRRiLL, Shallow-water Starfishes, 1914,
p. 139; Wrangell, Alaska.

Evasterias troschelii Verritu, Shallow-water Starfishes, 1914, p. 151, pl. 22, figs. 1, 2; pl.
25, figs. 1, 2 (type); pl. 26, figs. 1, 2; pl. 62, fig. 1; pl. 106, figs. 1, 2.

Evasterias troschelii var. rudis VERRILL, Shallow-water Starfishes, 1914, p. 158.

Evasterias troschelii var. densa VpRRiLL, Shallow-water Starfishes, 1914, p. 161; Victoria,
British Columbia.

Evasterias troschelii var. alvcolata VERRILL, Shallow-water Starfishes, 1914, p. 162, pl. 62,
fig. 1; Departure Bay, British Columbia.

Evasterias troschelii var. subnodosa VERRILL, Shallow-water-Starfishes, 1914, p. 163; Wran-
gell, Alaska.

Evasterias troschelii var. parvispina VERRILL, Shallow-water Starfishes, 1914, p. 163, pl. 106,
figs. 1, 2; Sitka, Alaska.

Evasterias acanthostoma VERRILL, Shallow-water Starfishes, 1914, p. 165, pl. 20, figs. 1, 2;
pl. 24, fig. 3 (type); Popof Island, Alaska.

Diagnosis.—Rays five, rarely six; actinal plates in three, sometimes four series
at base of ray; superomarginal plates lateral in position, defining ambitus; adam-
bulacral armature diplacanthid, or diplacanthid and monacanthid. Abactinal spines
exceedingly variable: (1) Large and much smaller ones intermingled without the
formation of a reticulate pattern (forma troschelii, pl. 60, fig. 1); (2) large and much
smaller spines, grading imperceptibly into uniformly coarse spines, but arranged in
reticulate pattern (forma alveolata, pl. 62); (3) spines uniformly very small and
arranged usually in a reticulate pattern (forma acanthostoma, pl. 64, figs. 2, 3; pl. 65).
Rays generally long and fairly slender, but very variable. Straight pedicellariae,
small lanceolate, without terminal teeth.

Evasterias troschelii is one of the most variable among many variable species of
the northwest coast. Its forms very closely parallel those of Pisaster ochraceus and
many specimens when preserved resemble slender rayed examples of that species.
Forma acanthostoma parallels forma confertus of Pisaster ochraceus; forma alveolata
resembles forma ochraceus; while forma troschelii and its acervate intergrades with
alveolata correspond to forma nodiferus (of P. ochraceus).

It is not possible to identify with certainty Brandt’s Asterias epichlora, from
Sitka, although in all probability the green phase of forma alveolata was referred to.
Brandt’s description is as follows: Diameter of the moderate sized subdepressed
disk scarcely exceeding a thumb-breadth. Rays five, conical, unequal, subdepressed,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—FISHER 141

about three thumb-breadths in length (i. e., R=6 r). Dorsal surface dull green,
covered with crowded white spines, of which the summit is capitate, arranged in a
reticulate pattern. Lower surface pale flesh color.

Professor Verrill applied epichlora to the commoner 6-rayed Leptasterias of the
intertidal zone of the southern Alaskan coast (two species). I have seen but one
5-rayed example of these species among several hundred examined; and if we adopt
an inch as a thumb breadth, the size given by Brandt does not conform to the size
or proportions of the Leptasterias, The proportions, inequality of rays, their slightly
depressed form, and color of Brandt’s species agree fairly well with small examples
of Evasterias which | collected at Departure Bay, British Columbia. However, in
order to avoid a great deal of confusion which would follow a shift of the name to
Evasterias, epichlora had best be consigned to the limbo of disused names. It is
certainly misapplied in connection with any 6-rayed Leptasterias.

The variations are so numerous and confusing in Evasterias troschelii that no
attempt will be made to describe the species. Rather the chief formae will be
analyzed as well as the material permits,

Forma TROSCHELII (Stimpson)

Plate 58, Figures 1, la, 4, 6, 6a; Plate 59, Figures 3, 3a; Plate 60; Plate 61, Figure 1; Plate 64
Figure 1

Asterias troschelii Stimpson, 1862, p. 267.

Asterias victoriana VERRILL, 1909, p. 68; 1914, p. 102.
Leptasterias macount Verriiy, 1914, p. 124.

Evasterias troschelii var. rudis Vern, 1914, p. 158.
Evasterias troschelii var, subnodosa VERRILL, 1914, p. 163.

This is the forma to which the type of the species belongs. Plate 60, Figure 1,
is an extreme having widely spaced and rather slender major spines, and unusually
few minor spinelets. Figure 2 is a stouter-rayed specimen having robust subtruncate
tubercular major spines. The minor spines are in lines here and there outlining
poorly defined areas. Plate 64, Figure 1, is a slenderer-rayed specimen in which
the minor spines have increased in number and it is intermediate with forma alveolata
having both coarse and slender spines, as Plate 62. When full grown, fat-rayed
specimens of troschelii have the spines rather widely spaced, as in Plate 61, Figure 1.
This is Verrill’s variety rudis—simply a growth stage.

The type forma is extremely variable and it is practically impossible to find two
specimens alike. There are slender and stout rayed examples. The major spines
are well spaced, or they may be clustered in twos and threes, or may form sub-
nodular prominences. The major spines are relatively slender or are very thick,
truncate, and many times the diameter of the minor spinules. The latter are few
and irregularly scattered between the major spines; or they may suggest a very
poorly defined reticulum.

The salient feature of this forma is the wide disparity in size between the major
and minor dorsal spines which do not form a meshwork design. Stimpson says:

The large ones are few in number, shorter but much thicker than the ventrals, capitate, with
flattened heads, and are arranged in a pretty regular though somewhat zigzag median row of about

142 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

25 spines, crowded near the disk, but further apart near the extremity of the ray. Between this
row and the marginal row there are scattered a few more of the larger kind, sometimes in clusters
or short rows of three or four. On the disk they form a more or less distinct pentagon, within
which there is another circle and a spine of large size in the center. The spines of the smaller
kind, minute, slender, and truncated, are scattered between the large ones. (Stimpson; see pl.
60, fig. 4, type.)

The large spines are striated at the summit, and in large specimens the smaller
sort are also pretty definitely grooved. The smaller spines sometimes form trans-
verse rows, but they do not define the papular areas, or complete the reticulation as
in alveolata, and intermediates. Four small specimens from near Tacoma have the
larger spines abnormally magnified and the smaller all but absent, while one from
Orea, Prince William Sound, Alaska, with R 65 mm., has the larger spines distinctly
acervate.

The superomarginal spines are slenderer than the larger abactinal, usually
tapered and blunt, and form a very regular series which curves up at the interradius
to the level of the madreporite. While the spines are generally single, sometimes a
small spinelet accompanies the larger. In giant specimens the principal spine is
often subcapitate, truncate, and striate.

The inferomarginal spines (usually two te a plate proximally) are a little longer
and slenderer, pointed, blunt, or subtruncate, varying individually. In giant speci-
mens there are usually two spines to a plate. The intermarginal channel is definite
and well defined.

Actinal plates in four series in very large specimens; in three series, in medium-
sized; and in two series in small specimens (R 20 mm.). In giant specimens they
generally carry two spines in an oblique row, except for the plates of the short inner
series which are monacanthid. In medium-sized examples there is usually one spine
to a plate; or three rows of actinal spines at the base of ray, then two, and finally one
on the outer half or third of the ray. But in giant specimens, where the plates are
diplacanthid there may be seven rows of actinal spines at the very base of the ray,
although the duplicate series of each range of plates is evident enough by rather
wide channels between the longitudinal rows of plates. The submarginal channel
is nearly as broad as the intermarginal channel. The spines are similar to the infero-
marginals and are variously cylindrical, tapered, or swollen, with blunt-terete or
blunt-compressed, or obliquely dressed extremities; or the whole spine is a trifle
curved upward. In very large examples the spines are shallowly grooved at the
tips. The spines are slenderer and longer than the larger abactinal spines, but in
medium-sized examples may be subequal in caliber.

The adambulacral armature is alternately one and two spines to a plate, with
one on the first five or six (adoral) plates, and two on a variable number (usually
relatively few) of succeeding proximal plates. The spines are slender, tapered, blunt,
the furrow spine, as is usually the case, being the slenderer. Near the mouth the
spines become longer and slenderer forming a dense chevaux-de-frise over the acti-
nostome. In a giant example, R 270 mm., the adoral adambulacral spines are 8 mm.
long (or subtend the are of the first eight adambulacral plates). At one-fourth R,
measured from the mouth, the spines are 5 or 5.5 mm. long. Medium-sized speci-
mens have two, giant specimens three (or four) adoral pairs of plates in contact
interradially.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 143

Mouth plates relatively small, with two unequal, apical and one longer, slender,
tapered suboral spine. As the animal grows larger, the actinostome becomes more
sunken and the mouth plates relatively inconspicuous. The tube feet are crowded
about the mouth, and the entrance to the furrow is only about as wide as the breadth
of the combined mouth plates. It widens rapidly, reaching the maximum width
at the sixth or seventh adambulacral plate.

The papular areas vary enormously with size of specimen. In giant examples
the whole abactinal surface as well as the lateral and actinal channels are a mass of
small papulae closely packed in larger areas (with narrow skeletal trabeculae sepa-
rating them). In medium-sized examples the papulae are also numerous, but the
areas are relatively smaller, though closely placed.

Straight pedicellariae (pl. 58, fig. 4) inconspicuous, compressed, broadly lanceo-
late, largest in lateral and actinal channels; absent or few on dorsal surface. They
occur on the oral and adambulacral spines, and not very numerously on the furrow
face of the adambulacral plates; length 0.5 to 0.8 mm.

Crossed pedicellariae (pl. 58, figs. 1, 1a) very numerous around the spines, among
the papulae, and on the surface of the plates between the areas, where they have well-
developed peduncles. Spine wreaths on big specimens very thick. They are com-
plete on the abactinal and both series of marginal spines; on actinal and adambu-
lacral spines the pedicellariae are in clusters. Abactinal pedicellariae 0.27 to 0.28
mm. long. Larger actinal and adambulacral pedicellariae, 0.31 to 0.36 mm. long.

Color in life, variable. A small specimen from Departure Bay, British Columbia,
gray green. A medium-sized specimen, general tint plumbeous, the papular areas
reddish brown; actinal surface flesh color.

Madreporic body prominent, slightly convex, with fine, intricate striae situated
about mid r.

Young.—Very small specimens resemble coarse-spined Leptasterias. The small-
est example (Orcas, San Juan Islands, Wash.) has R only 9 mm., but there is already
a series of actinal plates and spines. The young are rather easily recognized as
they have the peculiarities of the adult, even greater discrepancy in size between
the two sorts of spines. The larger spines are capitate, striate, subtruncate, and
form a very definite carinal series, with a few scattered dorsolaterals. There is a
regular series of superomarginals, of inferomarginals, and of actinals, the latter not
extending the entire length of the ray.

Type and type locality —That of species.

Remarks.—Although only the distal part of a ray of Stimpson’s type remains
(No. 1306, U.S.N.M., pl. 60 fig. 4), it is unmistakable. It closely resembles Plate 60,
Figure 1, which may be considered, therefore, as representing typical troschelii.
Verrill’s interpretation of troschelii, as represented by his Plate 26, is consequently
not correct.

Verrill’s Asterias victoriana (1914, p. 102, pl. 53, fig. 1) is a somewhat distorted
specimen of forma troschelii, having few minor spines. It resembles Plate 60,
Figure 2 (San Juan Islands, Wash.) but has slightly slenderer primary dorsal spines.
No. 603, Stanford collection, Insect Island, British Columbia, is nearly identical
with the type of victoriana.

144 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Verrill’s variety rudis, the type of which I have seen, is a giant form of troschelir
resembling Plate 61, Figure 1. His variety subnodosa (type from Wrangell, Alaska)
is a variant of forma troschelii, with large primary spines somewhat acervate in
places. It is one of the narrow rayed varieties between forma troschelii and the
variety of alveolata in which the alveolations are breaking up and primary spines are
differentiating as in Plate 62, Figure 1. I have examined the types of these two
varieties in the Yale Museum.

I have examined the type of Leptasterias macouni Verrill which was taken at
Departure Bay, British Columbia, and is now in the Ottawa Museum. It is, I
think, a 6-rayed, young, Evasterias troschelii probably forma troschelii. The radii are
41 mm. and 5 mm. The outstanding feature of this specimen is the presence of
four series of actinal plates (unheard of in a small Leptasterias) and the behavior of
the superomarginals, the series of which curves up strongly at the interbrachium.
The intermarginal channel is narrow and does not widen at the base. The first
series of actinals extends nearly to the tip of ray (within a few inferomarginals of
the tip); second row about three-fourths the length; third row, which is quite com-
pressed about half the length; there are four or five compressed plates of a fourth
row at base of ray (one or two of them have a tiny spinelet). Superomarginals
mostly with one spine, occasionally with a shorter fellow; proximal inferomarginals
with two, distal with one; first series of actinals proximally with one or two, distally
with one; other actinals with one spine. Adambulacral spines, slenderer than the
actinals, alternate two and one; on the diplacanthid plates the inner spine is advanced
somewhat into the furrow. The primary abactinal spines are mostly short, thick,
and stumpy, somewhat variable in size, while the relatively few minor spines are
shorter and much slenderer. Most of the carinal plates have one or two stubby
major spines and about two minors while the dorsolaterals have usually one major
and about two, quite slender, minors.

Six-rayed examples of Hvasterias troschelii are rare. I have one with R 72 mm.
from Departure Bay, British Columbia. It is close to the particular variation
shown by Plate 60, Figure 1, but the rays are relatively short (R=4.5 r).

The largest specimen which I have examined (station 4222) has R 355 mm.,
r 46 mm., br. 58 mm.

Forma ALVEOLATA Verrill, emended
Plate 58, Figures 2, 2a—2c, 5, 7, 7a, 7b; Plate 59, Figures 1, la; Plate 61, Figures 2, 3; Plates 62, 63;
Plate 66, Figure 1

? Asterias epichlora Branpt, 1835.

Asterias epichlora Strmpson, 1857, p. 528.—De Loriol, 1897, p. 19.

Asterias brachiata Perrier, 1875, p. 65.

Evasterias troschelii ‘‘typical form’’ VERRILL, 1914, p. 153, pl. 26, figs. 1 and 2.
Evasterias troschelii var. alveolata VERRILL, 1914, p. 162, pl. 62, fig. 1.
Evasterias troschelii var. parvispina VERRILL, 1914, p. 163, pl. 106, figs. 1, 2.

Forma alveolata is far from being homogeneous. It is a convenient pigeon hole
for a large number of small varieties which differ from typical troschelii in having
the abactinal spines, which are coarser than in acanthostoma, arranged in a reticulate
pattern. The spines are typically more numerous than in troschelii; and when the
spines are unequal in size, the smaller are much more numerous than in troschelit.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 145

Size medium to large, but I have seen no giants. Largest specimen R 205 mm.,
r 27 mm. br. 34 mm.

It is not possible to describe this forma with any degree of precision. The
principal variants will be indicated by means of figures.

(I) In its most generalized phase this forma may be described as a corase-spined
acanthostoma, the spines being short, subequal, and defining a reticulum. (PI. 63,
fig. 1.) This variation intergrades with acanthostoma; there are slender and thick-
rayed specimens and considerable range in the number and caliber of the spinelets.
Clayoquat Sound, Departure Bay, Gabriola Inlet, British Columbia; Straits of Fuca,
San Juan Islands, Wash.; Monterey Bay, Calif.

(II) The spines increase greatly in number and become fairly packed on the
carinal and dorsolateral plates, the papular areas becoming smaller. (Pl. 63, figs.
2,3.) These multispined forms intergrade with I; and a similar variation in acan-
thostoma differs only in having smaller spines and usually more numerous supero-
marginal spines. Fort Rupert and Departure Bay, British Columbia; Tongass,
Alaska; Monterey Bay, Calif.

(111) Disparity in the size of the spines gradually increases from a slight difference
as in plate 62, figure 4, through a series: Plate 61, figure 2; plate 62, figure 2, 3; plate
61, figure 3 and plate 60, figure 3, which have the spines as unequal as forma troschelii.
Plate 60, figure 3, may be classed as an aberrant troschelii, or as an intergrade.
Unalaska to Puget Sound.

(IV) Acervation of the major dorsal spines is developed in various degrees—
slight to very prominent (pl. 62, figs. 1, 3; pl. 63, fig. 3). Departure Bay and Bayne
Sound, British Columbia; Puget Sound.

Each of the above varies as to length and breadth of ray; that is, each variety
has longer and shorter as well as slenderer and thicker rayed representatives. I
intergrades with II, III, and IV; II and IV intergrade; III and IV intergrade.

In forma alveolata the abactinal spines are individually very similar to those of
forma troschelii. They vary from cylindrical truncate, through clavate forms, to
capitate, a little higher than broad to broader than high; and from capitate to a
curious flaring obconic form with a convex summit broader than the height. (Puget
Sound, Departure Bay, pl. 62, fig. 2.) The summit of the spines is finely striate.
The smaller spines, also striate, are cylindrical to clavate, obtuse to truncate, with
a certain number slightly tapered and bluntly pointed. When there is disparity
in the size of the dorsal spines the smaller are usually more numerous, stand in straight
or zigzag single file on the trabeculae and outline the skeletal intervals. The heavier
spines form a pretty definite carinal series either singly or in groups, and dorsolater-
ally occupy (singly or in groups of three to five) the principal nodal points of the
skeleton. From the formation of groups (pl. 62, fig. 4) it is only a step to an acervate
condition (IV, above).

Specimens from Unalaska have characteristically few major spines but numerous
much smaller minor spines. (Pl. 61, fig. 2; pl. 62, fig. 3.) These resemble forma
acanthostoma with a few heavy spines sprinkled over the surface, but the supero-
marginals are mostly monacanthid. They differ from troschelii in their more numerous
secondary spines which have a reticulate arrangement.

146 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The superomarginal spines are similar to those of forma troschelii, but often
not so regular. They partake of the character of the abactinal spines, varying in
robustness, but falling behind the abactinal spines in thickness. They vary from
cylindrical to capitate, truncate, the tip being striated, and sometimes asymmetrical
and oblique. Spines are usually single, but often accompanied by one or two, much
smaller, slender, slightly tapered. Very much less commonly there are two or three
subequal spines, but the specimens are not typical in other ways.

The inferomarginal spines are slenderer and longer than the superomarginals,
more or less clavate and bent, one or less often two to a plate. Proximally there are
three series of actinal plates, with three series of spines. In some specimens many
of the actinals are diplacanthid, so that the armature is crowded. I have not been
able to find any distinctive difference in the inferomarginal or actinal spines, since
the individual and age variations are very confusing. The third or inner actinal row
of plates is short as a rule.

Adambulacral spines very similar to those of troschelii, generally alternating
one and two to a plate, with two on a few plates following the adoral carina. They
are typically slender, subterete, blunt (or slightly tapered in the case of the alternate
furrow spine). In some heavy-spined examples they are narrowly spatulate, with
a shallow groove at end. Adoral carina composed of three to six contiguous pairs
of adambulacral plates in specimens with R 120 to 150 mm.; in specimens having
R 60 to 70 mm. there are usually three pairs in contact. The broader rayed examples
appear to have the fewer number in contact.

Actinostome very small, mouth plates closely similar to those of troschelii, having
two unequal apical spines and a longer, slender suboral.

Papulae very abundant, completely filling the spaces between spines and masking
the small ones in alcoholic specimens.

Crossed pedicellariae (pl. 58, figs. 2, 2a-2c) vary somewhat in minor details
and increase slightly in size in specimens from Unalaska. Monterey specimen;
length, 0.225 to 0.24 mm.; Puget Sound, 0.23 to 0.24 mm.; Unalaska, 0.31 to 0.34
mm.—all abactinal. The actinal and adambulacral are somewhat larger.

Straight pedicellariae (pl. 58, fig. 5) of small size and lanceolate in form, similar
to those of typical troschelii, are fairly abundant in most specimens in the inter-
marginal, interbrachial, and actinal channels, along the furrow face of the adam-
bulacral plates, and on the adambulacral spines; absent or very scarce on abactinal
surface.

Color in life very variable. Notes made at Departure Bay, British Columbia.
Red phase: (a) Clear flame scarlet, the spines and a very narrow surrounding zone,
pale cream color. (6) Vermilion, spines Saturn red to orange; or orange vermilion
above, larger spines and collars of pedicellariae, white. (c) Chinese orange, spines
ee white. (d) Brick red (Indian red), spines whitish. (e) Dragon’s blood
red.

Brownish phase: (a) Yellowish dark russet, spines paler. (6) Russet to burnt
umber; disk mottled with wood brown; spines whitish; skin flecked with pale Nile
blue. (c) Papulae deep ferruginous; spines whitish with basal zone of bluish gray.
(d) Mottled rich brown and gray; spines pale blue gray. These are pretty generally
pale clay color or faded yellow below.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 147

Green phase: (2) Arms banded dark sage green and bright olive; spines cream
color. (b) Arms olive green, groups of spines cream color, yellowish about their
bases. (c) Dark sage green without bands, spines whitish (also young with spines
pale sage green). (d) Major spines acervate—mottled olive green and cream color
(specimen similar to pl. 63, fig. 3). (e) Cream color, papular areas apple-leaf green
(specimen like pl. 63, fig. 2).

A specimen of the green phase may have been the type of Brandt's Asterias
epichlora.

Anatomical notes —Abactinal plates: There is considerable individual and age
variation. The abactinal reticulum is, of course, always irregular. The carinal
plates are four lobed and overlap over a third of their length, forming a zigzag or
irregular series. In alveolata there is one large spine and one to four small ones
on each carinal, but in troschelii the large spine is often without an accessory. Dor-
solateral plates of rather unequal size, irregular in shape, sometimes subtriangular
and connected by secondary ossicles also irregular as to size and shape. Marginal
and actinal plates closely imbricated, decreasing in size from the superomarginal
to the inner actinals. The plates are arcuately lozenge shaped to four lobed, the
dorsal lobe the longest. In the case of the actinals and inferomarginals it strongly
underlaps the descending lobe of the plate, just above. There are basically three
series of actinals, with a short fourth series in very large examples, in contrast to
the six (or seven) of E. echinosoma.

The actinostomial ring (pl. 59, fig. 1) is small, the actinostome being about as
broad as the length of the first 10 ambulacral ossicles (or the first pair plus nine).
The median suture between the two halves of the first enlarged pair is serrate and
the length of the dorsal crest is that of the next six plates—a very common propor-
tion in the Asteriidae. The odontophore has a characteristic form reminding one
of the valve of a small chiton shell. The interbrachial septum is small owing to
the small size of the disk. There is a pillar of plates extending upward from the
odontophore, and between this and where the true arm plates end at the inter-
brachial angle there is an uncalcified membranous septum in medium-sized speci-
mens. In the giant examples the whole septum appears to be filled with plates,
and in these specimens the rather triangluar entrance from the disk coelom to that
of the arm is surprisingly small and filled by a much folded lobe of the ventral
stomach.

The gonads open dorsally, close to the interradial line, about one-third r from
the margin; and in general form are elongate fusiform with numerous fusiform
lobes on the mesial side, these being subdivided into short globular lobules. The
intestinal coecum is tripartite; the stomach is rather crowded in the small disk.
and the hepatic coeca are large and extend nearly to the end of the long rays. No
Polian Vesicles.

Type locality of forma alveolata.—Departure Bay, British Columbia.

Remarks.—This forma corresponds to Dr. William Stimpson’s Asterias epichlora,
as several specimens labeled by him are in the collection of the United States National
Museum. He considered Asterias troschelii to be a distinct species.

I think Doctor Stimpson’s identification of Brandt's epichlora is correct; green
specimens are not uncommon. It would lead to endless confusion to adopt epi-

148 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

chlora, however, since Professor Verrill has used the name for the common 6-rayed
Leptasterias of the Alaskan coast, in spite of the fact that Brandt writes that his
epichlora had five rays.

The name alveolata was given by Verrill to a specimen taken at Departure
Bay, British Columbia, and now in the Ottawa Museum. It is one of the swollen-
rayed variants, such as Plate 62, Figure 2, but with the spines asin Plate 63, Figure 1.
The papular areas are sunken, accentuating the skeletal mesh, since the specimen
was dried before being properly preserved.

The specimen figured by Verrill (1914, pl. 26, figs. 1 and 2) as typical troschelit
belongs to forma alveolata. It is very similar to Plate 62, Figure 4, and has not
the peculiarities of the type of troschelii. Verrill’s Plate 22 represents a very imma-
ture specimen from Juneau, Alaska. It is a young alveolata with considerable dis-
parity in size between major and minor abactinal spines. The areolations are not
so distinct as in the foregoing specimen. ‘This is characteristic, however, of many
young specimens, which pass through a stage in which they resemble forma troschelit.
The young of Pisaster ochraceus usually have acervate dorsal spines. Much fewer
adults are so characterized, so that the young are in many cases transiently forma
nodiferus.

The type of Verrill’s variety parvispina is a very immature alveolata which was
taken at Sitka. Its peculiarities are partly due to immaturity. Among examples
of troschelii and of alveolata having two sizes of dorsal spines, certain ones not infre-
quently have unusually slender minor spines, while the major spines are somewhat
less often quite slender.

Forma alveolata is therefore much more inclusive than Verrill’s variety alveolata.
This name has been adopted since Asterias brachiata Perrier, 1875, is invalidated
by A. brachiata Linnaeus.

Most of the specimens which I collected or examined at Departure Bay, num-
bering several hundred, were accompanied by a commensal polynoid annelid, Halo-
sydna fragilis, which lives among the tube feet, usually at base of ray, or in the

actinostome.
Forma ACANTHOSTOMA (Verrill)

Plate 58, Figures 3, 3a, 3b; 8, 8a—8c; Plate 59, Figure 3b; Plate 64, Figures 2, 3; Plate 65; Plate 66,
Figure 2
Asterias acanthostoma VERRILL, 1909, p. 543, footnote.
Evasterias acanthostoma VERRILL, 1914, p. 165, pl. 20, figs. 1 and 2; pl. 24, fig. 3.

Plate 64, Figure 3, shows a typical specimen from Unalaska and Plate 66,
Figure 2, the actinal surface of the same; Plate 64, Figure 2, is one of the most
typical of the Tongass, Alaska, specimens. Plate 65, Figure 1, is a large slender-
rayed variant from Orcas Island, San Juan Islands, Wash., and Figure 2 a thick-
rayed specimen from Victoria. Plate 65, Figure 3, is an enlargement of a ray of
the Unalaska specimen. Plate 63, Figure 2, is a problematical variant from Tongass,
Alaska. Although taken with acanthostoma, it and other specimens may perhaps be
more appropriately classed as very slender spined alveolata.

Forma acanthostoma intergrades with alveolata. It is not possible to place
certain specimens definitely in either forma since they stand midway between.
Typical specimens differ from forma alveolata in having uniformly small abactinal

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 149

spinelets which stand in single file on the irregular reticulum of the skeleton and
divide the abactinal area into areolae, or are more grouped and scattered, so that
the reticulation is not so evident; superomarginal spines in combs or groups of three
to five; adoral adambulacral spines slightly longer. In mature specimens R=5 to
7.6r and 4 to 6.5 br. Largest specimen, R 310 to 330 mm., r 46 mm., R= about 7 r.

The abactinal spines are all small and of nearly uniform size, except near the
tip of the ray, where in some but not all specimens they are unequal, the larger
doubtless being homologous with the major abactinal spines of alveolata. The
spinelets are numerous, nearly equal, either tapered or clavate, with rough sulcated
tips, and they either stand in single file on the skeletal meshes, very definitely sub-
dividing the dorsal area into rather irregular, numerous areas (the papular areas),
or, less often, they do not form any very evident reticulations. In the type speci-
men: ‘‘Most of those on the basal part of the rays stand in short transverse or oblique
rows or combs, varying from 2 or 3 to 10 or more, but many stand singly or in small
clusters. On the distal third of the rays they become much more crowded and
stand in small groups or singly; on the disk they form irregular short rows or imper-
fect reticulations.”’ (Verrill.) Several specimens from Unalaska and Victoria exhibit
this imperfect reticulation but it is only an individual variation. (PI. 65, figs, 1, 2.)
More often the spinelets are arranged in a definite reticulate pattern, in which case
the spinelets are more numerous. (PI. 64, figs, 2, 3.) On the proximal half of ray
these reticulations are more or less lengthened transversely, so that the spinelets
appear to form transverse combs of variable length, because the transverse trabeculae
are much longer than the connecting longitudinal ones. The carina! series may not
be distinct or it may be well marked, as in most of the Tongass examples, in which
the spinelets stand in several ranks along the radial line.

The superomarginal spines are similar to the dorsals but a little longer and
stouter. There are generally three, four, or even five to a plate, in an arcuate vertical
comb; or a comb of three, with an extra spine on its adoral side. Distally the combs
are reduced to two spines, and near the tip there is usually only one spine. The
spines are sometimes subequal; or the fourth and fifth spines, when present, may be
smaller; or the median spine of the series may be somewhat enlarged over the others.
Intermarginal channel broad in large specimens; in medium sized ones narrower
than the superomarginal.combs.

Inferomarginal and actinal spines similar in form to those of alveolata, rather
crowded, and increasing in size from the inferomarginals (which are a little longer
than the superomarginals) toward the furrow. Inferomarginal spines are usually
two to a plate (sometimes three, and at base of ray one). The spines are clavate,
or tapered, obtuse, or bluntly pointed, finely striated at the ends. There are usually
three rows of actinal plates at the base of the ray; rather exceptionally four (Verrill
mentions five, but I have never found so many in my specimens). Plates of the
outer row bear two or three spines in an oblique series; the others generally two;
whence the crowded condition of the actinal armature. The spines vary greatly
in caliber from a slightly swollen form with a tapered blunt terminal half, to broadly
clavate, subtruncate. In the very large specimens the interactinal channels are well
marked but tend in others to be obliterated by a crowded armature.

150 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The adambulacral spines stand alternately one and two to a plate, with a
variable number of diplacanthid plates following the adoral carina. The spines are
similar in form to those of alveolata, but a little longer toward the actinostome. This
character is not of much practical use, owing to variability and the difficulty of
finding a method of comparison. In acanthostoma the longest adoral subambulacral
spine (of the second, to fourth plate) is about as long as the first eight adambulacral
plates measured on the furrow face; and in forma alveolata it is about seven plates in
length—perhaps 10 to 14 per cent less. In forma troschelii the spines are usually
still shorter. There are generally five pairs of adoral adambulacral plates joined to
form a narrow adoral carina and the actinostome is very contracted, as in alveolata.

The oral plates are not appreciably different from those of alveolata, there being
two (sometimes three) apical, and one suboral spines.

The madreporic body is large, with fine, very irregular, striae; its outer edge is
about mid r.

The papulae are very numerous and distributed as in alveolata; the abactinal
papular areas are often smaller and more numerous, but there is no constant dif-
ference.

The straight pedicellariae are similar to those of troschelii—lanceolate, blunt or
pointed, small—and are similarly distributed.

The crossed pedicellariae are very numerous among the papulae as well as
surrounding the spines. They have the distribution characteristic of the genus.
The abactinal pedicellariae measure about 0.30 to 0.31 (Unalaska); 0.2 to 0.26
(Tongass); 0.23 to 0.24 (Victoria). In profile the upper part of the outer surface of
the jaw is flattened or even slightly depressed. Northern specimens have larger
pedicellariae.

Variations.—The principal variations of the type form have already been noted.
The abactinal spines vary in robustness as well as in the precise shape. In the largest
specimen from Victoria, British Columbia, they are cylindrical or slightly tapered,
two or three times as high as thick, blunt or truncate, with a suleated tip. This is
the usual form but a number of spines especially near the end of the ray are thicker
and clavate. There is less difference in spine form than in alveolata. The differences
in the arrangement of the abactinal spines—by which the papular areas are more or
less completely fenced in—constitute one of the major variations. There is also a
conspicuous difference in the robustness of the rays. The largest specimen from
Victoria, British Columbia, with R 310 to 330 mm., and considerably shrunken in
drying has the ray 60 to 65 mm. broad at base, or about one-fifth R. The stoutest
specimen has the arm breadth about one-fourth R (Unalaska), while another from
the same locality has the arm breadth one-fifth to one-sixth R.

Tongass, southeast Alaska, Plate 64, Figure 2. The specimens from this locality
have been referred to acanthostoma, although they present certain points of difference
which at first sight give them a very distinctive appearance.

The rays are very slender and long, R equalling about 6% br measured at widest
part. The rays taper very gradually from this narrow base. The abactinal spines
are unusually numerous and are small. They completely surround the papular areas,
dividing the abactinal surface into very numerous irregular meshes. The carinal
series is pretty well marked at the base of the ray. The spinelets vary in different

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 151

individuals, being cylindrical or tapering and blunt, and rather slender, or else shorter
and thicker, with a more or less broadened, truncate summit. This character is more
marked on the median than on the lateral portions of the abactinal area. In forms
with well-marked carinal plates there are usually numerous crowded spines on each
plate. It is only a short step to the production of such forms as that shown in
Plate 63, Figure 2, which can be classed as a small spined alveolata, A few specimens
from Tongas must be placed in alveolata on account of the coarse abactinal spines
and the presence of only one or two superomarginal spines. The ordinary Tongass
acanthostoma have from three or four to six superomarginal spines, of which one is
usually larger than the rest.

Certain specimens in the Puget Sound region are similar to the Tongass variety.
(Pl. 65, fig. 1, Oreas Island, San Juan Islands.)

Type locality of forma acanthostoma.—Popof Island, Shumagin Islands, Alaska.

Remarks.—The type of Verrill’s acanthostoma was taken at Popof Island, Alaska,
and the forma is best developed in the region between Unalaska and Kodiak. In
southern Alaska, British Columbia, and the Puget Sound region it intergrades so
frequently with isacanthous forms of alveolata that it can be described as a small
spined offshoot of the isacanthous varieties of alveolata having polyacanthid super-
omarginals.

A specimen from Departure Bay which has the small pointed abactinal spines
of some specimens of acanthostoma, but only two superomarginal spines, in life was
dark bay color, the spines being Antwerp blue.

Type of Evasterias troschelii—Cat. No. 1306, U.S.N.M.

Type locality of Evasterias troschelii.—Puget Sound, Wash. (Simeahmoo), north-
west boundary survey, Dr. C. B. Kennerly.

Distribution. —St. George Island, Pribilof Islands to Monterey Bay, Calif.
Intertidal zone to 39 fathoms.

Specimens examined.—Three hundred and forty-six.

Specimens of Evasterias troschelii examined

ae EES
{. tro- | f. alreo- f.acan-|
Locality achelii lata As Remarks

i
} ‘(iebine R W mm., young.
itinNaR SPSS ee fix eos 8 $ | Albatross, 1 intermediate.

Captains Bay, Unalaska....................-.----- Ltn ago | W. H. Dall, 1872.

Dutch Harbor, Alaskn.........-------.----.-----+- bao bets I | --neeee Stanford Coll.

Mist Harbor, Nagai, Shumagin Islands a heicen eet 4 Albatross, 1893,
SOVRIenes Modine ee err Nie ee ! 1 | Albatross, 1903, seine.
Witla a 8 Finfish ans wads te weld onde BA schdpoil W. J. Fisher.

Orea, Prince William Sound, Alas! B | 1g Recs. | Stanford Coll

ea ae iat 0s oS Se i 1| 2) Albatross, 1892.
opin Bing Anais ise c sin st da sndenxd- Lelhoedn ss Bbc Albatross, 1008.
Tongnet, AMER q..nccncccnccnn cence. cede sersceee|secesees! 5) 15 | Nichols.

Ward Cove, Revillagigedo Island, Alaska. .........|--....-. } 1) 3 | T. HT. Street.
Worrebter, Meberi ANOMR0 oss cssccne ton dncnp oanin-fagsocess 1} mo Stanford Coll., H. Heath.
Herendeen Bay, Alaska... .........2.....02-.00-e)eeeeees 6) 1) Albatross, 1890.
Freshwater Bay, Alaska. ...... Roe ES | Intermediate

Insect Island, British Columbia D Lakindd. Bs ee Stanford Coll

Fort Rupert, British Columbia...... selbahebtl) veh Lcaeaas Do

Boundary Bay, British Columbia... ............... 1 fuy.

1 i
SLoit.2 Opster beds; Stanford Coll.
6)

Union Bay, Bayne Sound, British Columbia_....... 12) 1 | Albatross; shore.

152 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Specimens of Evasterias iroschelii ecamined—Continued

tro- | f. alveo- | & 1c27- . -
Locality BG f las a Remarks

Taylor Bay, Gabriola Inlet, British Columbia------|-------- Are eee eee Albatross.
Beaver Harbor, British Columbia Do.
Comox, British Columbia---.---.------------------
Departure Bay, British Columbia------------------ SO) |eaeenee 1 | acanthostoma not typical.

BSS a 6 |___.....] Stanford Coll., W. F. Thompson.

pane 113 25 2 | Calif. Acad. Sci. Coll., W. K. Fisher.
Clayoquat Sound, British Columbia- ae pee U9 | eres Stanford Coll., C. H. Gilbert.
Barclay Sound, British Columbia---- -| ljuv. 6) /4oes eee Albatross.
Victoria, British Columbia---..-----.-------------- 2d | Pa eenen 1 Do.
Straits of Fuca_...---.---- ~weassoscesesene 1 1 D.S. Jordan.
Porte hownsend a Vis Ueessss nee eae en a | ae meres Laie -| J. G@. Swan.
Puget Sond Wy PSlssee ae a eae to ee ee 6 D.S. Jordan, Oscar Sturges, C. M. Drake.
Puget Sound (Simeahmoo)-.--.--------------------- 3 8 Northwest boundary survey, C. B. Kennerly.
Puget Sound (Tacoma) --.------------------------- 5 10 F. W. Weymouth.
Oreas Island, San Juan Islands, Wash-------------- 122 | 10 Calif. Acad. Sci. Coll., Ida Oldroyd.
Friday Harbor, San Juan Islands_----------------- 6 9 Do

1

Station 4205, vicinity of Port Townsend, Wash., 26
fathoms.
Station 4222, same locality, 39 fathoms--__----------- 1

Bottom, rocks, shells; bottom temperature, 50.8° F

Pot sasee |Seeetons Bottom, gray sand, broken shells. R 355 mm.,
r46mm., br 58 mm.

C. M. Drake.

.| Stanford Coll., W. F. Weymouth.

Beatrice, Humboldt County, Calif--.-..-..--.-----
Trinidad Head, Calif.......---.------
San Francisco Bay, Calif-- Stanford Coll., W. F. Hamilton.
Monterey Bay, Calif.._-.- ps R. E. C. Stearns.

Q@armel Bay; Califio - ae 2 - <n acces encom nos eeneneee=|nanmne— 1s) eee Stanford Coll., H. Heath.

1 Many intermediates; all stages between (roschelii and alveolata.

EVASTERIAS ECHINOSOMA Fisher
Plate 59, Figures 2, 2a-2h, 4; Plates 67-69

Evasterias echinosoma FisHer, A New Sea Star of the Genus Evasterias, Proc. U.S. Nat
Mus,, vol. 69, art. 6, 1926, p. 2, pls. 1, 2.

Diagnosis. —Size large; rays five, long, tapering, stout, more or less swollen,
with a very convex abactinal, and a subplane actinal surface. Differing from E.
troschelii in having uniformly large, mostly subconical, well-spaced abactinal spines;
marginal plates unusually high on side of ray, the superomarginals being abactinal
in position and generally monacanthid; six (or five) series of actinal plates (generally
monacanthid) of which either the upper row or the inferomarginals define the margin
of ray; adambulacral plates triplacanthid, or diplacanthid and triplacanthid. Type,
R 330 mm., r 51 mm., R=6.4 r; br=about 60 mm.

Description.—The abactinal surface is armed with rather widely spaced and nearly
uniform robust spines, cylindrical at the base, the distal half conical, longitudinally
suleated, bluntly pointed, and in giant specimens with R 300 mm., about 2.5 mm.
long, by 1 to 1.5 mm. thick at the base. The distal part of the spine may be slightly
swollen so as to appear subcapitate. The spines of the distal portion of the ray are
round-tipped, and by a shortening and rounding of the terminal conical portion a
subglobose, striated tip results. A majority of spines are so formed in specimens
from stations 3281 (2), 3291 (1), 3235 (1), none of which has R greater than 200 mm.
In the specimen from station 2842 the spines are slenderer than in the type, tapering,
and pointed. The spines do not have regular arrangement. An irregular carinal

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 153

series is generally fairly well marked, the dorsolaterals standing typically singly
(but sometimes in groups or lines of two, three, or even four) on the chief nodes of the
reticular skeleton. In some of the very large specimens there are a few very delicate
terete spinelets, scarcely longer than the abactinal straight pedicellariae, scattered
over the abactinal surface. In the specimens in which the spinelets are more or less
grouped, there is rather less uniformity in size, some being of distinctly secondary
size. There is a broad and pretty definite supramarginal channel bounded abactinally
by a very irregular row of dorsal spines, which usually but not always stand closer
together than on the rest of the dorsolateral region.>

The superomarginal spines are similar in form to the abactinal spines (following
the variations of the latter) and are generally slightly smaller. Typically they stand
one to a plate, close together, forming a very well-defined series, characteristically
high on the side of the ray, so that the proximal half at least, and sometimes the whole
ray is bordered, when viewed from above, by the inferomarginals, or by the first series
of actinal spines. This character is accentuated in small examples (R 110 mm.),
in which the abactinal area is narrow. Interradially the superomarginal series ex-
tends halfway to the center of the disk. Two or three spines occur on the plates of
the proximal half of ray in specimens from stations 4796, 3235, and 3291 (one each).

There is a wide intermarginal channel (2 or 2.5 times length of inferomarginal
spines). Inferomarginal spines similar to superomarginals, but a little longer (3
or 4 mm. in giant specimens), sometimes one to a plate, sometimes two, or rather ir-
regularly one and two proximally and one distally. The series bends upward inter-
radially, and in some specimens is abactinal (or dorsolateral) in position.

In large specimens there are six series of spiniferous and one short series of spine-
less actinal plates at the base of the ray. There is considerable variation in the num-
ber of spines per plate. All plates may be monacanthid. In this case there are
eight regular, spaced, longiseries of spines, of which two are marginal series abactinal
in position. (Station 3282.) The outer three or four series are sometimes regularly
or irregularly diplacanthid and the inner two or three monacanthid; or the outer row
may be monacanthid, the next two irregularly diplacanthid, and the remaining three,
monacanthid. (Station 3281.) In a specimen from station 2842 a considerable num-
ber of plates are triplacanthid. In large specimens the sixth or inner series of actinal
spines extends one-third R measured from center of disk. The actinal spines be-
come gradually a little longer, sometimes heavier and clavate, in passing from the
outer toward the inner series. The details of the actinal spines are variable, as in
other species. The tips may be compressed and subtruncate, suleate, or tapered,
blunt, or pointed. The smallest specimen (station 3650) with R 46 mm., has four
series of actinal plates. The large specimen from Kamchatka (station 4796) with
R 265 mm. has but five series of actinal plates. Whether this is constant for large
Asiatic examples can not be determined.

The actinal channels are typically well marked, even broad in some cases, so
that the rows of spines are distinct and separated.

The adambulacral are triplacanthid and diplacanthid. In large examples most
of the plates of the proximal half of the ray are triplacanthid then irregularly dipla-
canthid and triplacanthid and finally on the distal third of the ray, mostly dipla-
eanthid. The distribution of these numbers will, of course, vary in different indi-

154 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

viduals. In general the proportion of plates occupied by three spines increases with
the size of the animal, the third spine being added on the outer side of the plate.
The combs of alternate plates are advanced further into the furrow. The first three
plates following the mouth plates are generally monacanthid; then three or four are
diplacanthid, following which, after a few plates of three and two, the regular trip-
lacanthid plates commence. The spines are slender, about as long as the inner
actinals. The furrow members are slightly tapered; the others, a little stouter,
varying from slightly tapered to cylindrical, or somewhat clavate, round tipped to
bluntly pointed. The third, outer spine may be shorter than the other two. There
are usually five pairs of united plates composing the adoral carina. The large Kam-
chatkan example is diplacanthid, and near the end of the ray, irregularly diplacan-
thid and monacanthid.

Actinostome very small. Mouth plates with two apical spines in nearly verti-
cal series, the smaller at the mouth of the furrow, the other (about as long as the
plate, and sometimes spatulate) almost directly above it (as viewed from actinal
side). The suboral spine, near outer end of plate, is about as long as first two or
three adambulacrals.

The papulae have the distribution characteristic of the genus, and are very
abundant, especially abactinally, where, in alcoholic specimens they appear to
occupy all the space between the prominent circles of crossed pedicellariae surround-
ing the spines. The size of the areas increases with age; about eight or nine areas
can be counted across the ray at base, but the dorsolaterals are very irregular. There
is a fairly regular supramarginel row. The intermarginal and actinal rows—eight in
all—are typically regular and decrease in size toward the furrow.

Straight pedicellariae (pl. 59, figs. 2, 2a-2f): There are two sorts, larger and
smaller; the larger, usually compressed ovate, wedge-shaped, with the end broadly
rounded and the tip of each jaw with two or three denticles, varies from abundant to
relatively few on the abactinal surface; they are generally abundant on the inter-
marginal and actinal integument, and a few occur on the inferomarginal, actinal,
adambulacral, and oral spines. They vary to lanceolate obtuse and lanceolate acute.
In large specimens the abactinal measure about 0.9 to 1 mm., while the actinal inter-
radial ones are 1.5 mm. long. Much smaller ones are present in variable numbers
on the actinal, adambulacral, and oral spines, and are rather sparsely scattered
along the furrow face of the adambulacral plates.

Small crossed pedicellariae (pl. 59, figs. 2g, 2h) are very abundant singly and in
groups among the papulae; in a broad zone around the abactinal and marginal
spines; and in half-wreaths on the outer side of the actinal and adambulacral spines.
The abactinal measure 0.27 to 0.3 mm. while the adambulacral measure 0.35 to 0.4
mm. (large specimens with R 270 mm. or more). Apparently there is a gradual
increase in the number of crossed pedicellariae, especially the papular, with age.

Madreporic body large, subplane, with a row of spinelets on the adcentral border;
it is situated a little less than one-third r from center of disk.

Variations.—The chief variations have been noted in the foregoing account.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 155

The example from off Petropavlovsk, Avatka Bay, Kamchatka (pl. 68, fig. 1),
suggests a representative of a slightly differentiated race. There is but one large
specimen, and therefore no chance to test the constancy of its characters. The
abactinal spines are more numerous than in typical east Bering Sea examples and are
grouped, or in lines, so that the reticulum of the skeleton is indicated, but not clearly
marked; there are but five series of actinal plates, and only four are prominent; the
proximal superomarginal plates are polyacanthid; the adambulacrals are diplacan-
thid, and the straight pedicellariae are few instead of abundant. The small speci-
men from the Okhotsk Sea, having R 46 mm., has four series of actinals, typical
abactinal spines, and very abundant straight pedicellariae.

Anatomical notes.—The carinal plates are four lobed, and strongly imbricated in
series. The dorsolaterals are irregular, in outline separated less than their own
diameter (occasionally directly overlapping by a lobe), and are connected by elon-
gated secondary ossicles on the coelomic side. These become longer toward the
marginal plates, and all underlap strongly the major plates, showing above as narrow
connecting pieces or trabeculae of the reticulum, of which the primary plates form the
disproportionately thickened nodes. The marginals and actinals are four lobed (the
transverse lobes the longer) very strongly imbricated, so that the intervals between
the plates are rather small, and the plates decrease in size from the superomarginals
to the innermost or seventh series of actinals which is spineless. The upper, or,
underlying, lobe of the actinals is the longest and is more or less compressed and
carinated.

The adambulacral plates are very short and broad, the width equaling the
length of about five consecutive plates. The ambulacrals are also very wide and short,
the width of course varying in different parts of the ray. The ambulacral pores are
quite slitlike. In large specimens there are 11 ambulacral plates in the length of 10
mm. (about opposite interbrachial angle).

The ovaries of a small specimen (R 123 mm.) are mature and open dorsally close
to the interradial line at about mid r. The gonad is roughly spindle-shaped and
consists of numerous short subequal lobulated divisions arranged rather symmetric-
ally around a central axis.

Type—Cat. No. E1237, U.S.N.M.

Type locality—Station 3278, north of the end of the Alaska Peninsula (56° 12’
30’’ N., 162° 13’ W.); 47 fathoms, fine gray sand; bottom temperature, 38.8° F.

Distribution —Southern Bering Sea, from Bristol Bay to Unalaska; the coast of
Asia from Avatka Bay, Kamchatka, to the Okhotsk Sea; 11 to 48 fathoms, fine
sand, mud, pebbles, stones; temperature range 38° to 41.2° F.

Specimens examined.—Thirteen.

156 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Specimens of Evasterias echinosoma examined

|
Sta. | | Bottom) Der of
Re Locality Depth | Nature of bottom temper- speci- | Collection
tion | ature
| | mens
1a ail ae AEs cee Ma tip ep tele
| Fathoms eRe
2842 | Off north coast Unalaska....-..-.--------- a1 OD Des sesec nee e een ee 41 1] U.S.N.M.
3235 11 | Black stones _- 1 Do.
3241 14} Black mud------ 1 Do.
3278 | North of end of Alaska Peninsula 56° 12’ 47 | Fine gray sand_--_------ oe ESRD 2 Do.
| 30” N., 162° 13’ W.
S281 |----- CD saee et ene sean ae eee eee $0) GxayisandG*-a-teee nee \oeee abe 2 Do.
3282 53 | Fine sand, green mud_-_-_---- 38. 2 1 Do.
3285 35 | Gray sand 41 1 Do.
3291 | Mouth Bristol Bay, near Alaskan Penin- 26 | Black sand 41,2 2 Do.
sula, 56° 58’ 30’ N., 159° 11’ W. |
3650 | Okhotsk Sea (to westward of Robben 28 | Brown mud, sand_---.-.----|_------- 1 Do.
| Island). | je
4796 | Avatka Bay, Kamchatka, 52° 47’ N,, 48 | Sand, pebbles, shells. __..-__|___----- 1 | Albatross, 1906.
158° 43’ BE. |

Remarks.—This well-marked species is separated from troschelii by several
definite characters: (1) The well-spaced, stout, subconical, conical, or occasionally
capitate abactinal spines; (2) the unusually numerous actinal plates, there being
seven rows in large specimens; (3) the position of the marginal plates relatively high
on the side of the ray, so that the superomarginals are abactinal in position; (4) the
triplacanthid and diplacanthid adambulacral plates.

A specimen from station 3291 (with R 162 mm. pl. 68, fig. 2) has the abactinal
spines rather more numerous, and considerably smaller than in typical specimens.
It is not to be confounded with E£. troschelii forma acanthostoma, because the other
characters are typical.

Genus STEPHANASTERIAS Verrill
Stephanasterias Verriui, Bull. Essex Inst., vol. 1, 1871, p.5. Type, Asteracanthion albulus
Stimpson.
Nanaster Perrier, Expéd. sci. Travailleur et Talisman, pp. 129, 131, 1383. Type, Stichaster
albulus (Stimpson).

Diagnosis —Small fissiparous Asteriinae with upward of nine rays, and a closely
reticulate rather irregular abactinal skeleton consisting of small, lobed or elliptical,
imbricating plates, of which the closely imbricated 4-lobed carinals form a definite
fairly regular series; actinals absent; marginals low on side of ray, and, like the abac-
tinals bearing several short delicate spinelets; papulae simple, one to several to each
area, of which there are upward of four poorly defined dorsolateral and one inter-
marginal longiseries on either side; adambulacral plates short (compressed) dipla-
eanthid or triplicanthid, without spine pedicellariae; adoral carina consisting usually
of one pair of contiguous adambulacrals increasing to three in largest specimens;
tube feet in four longiseries; large, toothed, and unguiculate straight pedicellariae;
crossed pedicellariae without enlarged terminal teeth; gonads opening low on side of
ray in interbrachial channel, intermarginally.

In addition to the type species, S. gracilis (Perrier) has been described from the
West Indian region (off Florida, Lesser Antilles, Cuba, 56 to 200 fathoms).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 157

Stephanasterias hebes Verrill from Albatross station 2766, 10.5 fathoms, off the
east coast of South America (lat. 36° 47’ S., long. 56° 23’ W.), is not a Stephanasterias,
but probably an Allostichaster.™

STEPHANASTERIAS ALBULA (Stimpson)
Plate 70, Figures 1-5; Plate 71, Figures 1, la-1g; Plate 72, Figure 5
Asteracanthion albulus Stimpson, Invert. Grand Manan, 1853, p. 14, pl. 1, fig. 5.
Asteracanthion problema Steenstrup, Vidensk. Medd., 1854, p. 240.—LUtxen, Gronlands
Echinodermata, 1857, p. 30; Vidensk. Medd., 1871, p. 300.
Asterias albula Stimpson, Proc. Acad. Nat. Sci., Philadelphia (1863), 1864, p. 142.
Stichaster albulus Verrtu1, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 351 (var. nifida,
p. 351).—Lupwia, Arktische Seesterne, Fauna Arctica 1, 1900, p. 479 (literature list).—
Von Horsten, Echinodermen des Eisfjords, 1915, p. 56, fig. 19 (distribution).
Stephanasterias albula Verritt, Bull. Essex Inst., vol. 1, 1871, p. 5; Shallow-water Star-
fishes, 1914, p. 147:

Description —The small, close-set groups of dorsolateral spinelets, upward of
five to a plate, tend to form close transverse rows, extending from the differentiated
carinal series to the inferomarginals which are low on side of ray. Carinal plates
somewhat larger than the dorsolaterals, more strongly imbricated and with five or six
divaricate spinelets. In a large specimen abactinal spinelets are 0.45 to 0.55 mm.
long and are tipped with usually three to six sharp thornlets. In aleocholic specimens
the spinelets are skin-covered, papilliform and not easy to differentiate from the
crossed pedicellariae which are scattered among them. Papulae small, single or few
to an area. Two irregular longiseries of areas nearest carinals are about the size of
the plates but two other series on dorsolateral region of ray (the outer one, supramar-
ginal) are usually considerably larger than the surrounding plates.

Superomarginal plates four lobed, closely imbricated, the descending lobe longest.
Each bears a transverse (vertical) comb of four or three spinelets similar to the abac-
tinal but sometimes a shade larger. Inferomarginal plates, on ventrolateral border
of ray bear an oblique comb of three or four spinelets of which the lower two (or
sometimes three), are considerably longer and often much thicker than the supero-
marginals. In old specimens some of them may be spatulate, or clavate and slightly
curved. Usually, however, they are all slender. They are about 1 to 1.25 mm. long.

Adambulacral plates short, so that the combs of three spines per plate (rarely
four), are close together and form a cheveux-de-frise along the furrow margin. Plates
of the outer attenuate portion of the ray frequently carry two spinelets. The spinelets
are subequal to the inferomarginals and like them are sometimes thickened in old
specimens. The middle one is often slightly to decidedly compressed and broadened
toward the tip.

Each mouth plate carries on the narrow suboral surface two to four tapered
spinelets similar to the succeeding adambulacral spinelets (of which there may be
only two on the first two or three narrowed adambulacrals). At the base of the
innermost oral spine there is a cluster of small lanceolate straight pedicellariae (sumilar
to those which fringe the furrow margin). One to several large unguiculate
pedicellariae stand at the surface of the plate.

*® West Indian Starfishes, Bull. Univ, Iowa, vol. 7, no. 1, 1915, p. 24, pl. 9, Me. 3.
63160—30——11

158 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Crossed pedicellariae (pl. 71, fig. 1b), numerous, averaging 0.25 to 0.27 mm.
with many as small as 0.225 mm. ‘They are attached at the base, sometimes on the
base, of the spinelets around which they form complete or partial circlets. They are
present sparingly on the inferomarginal spines, except the inner or lowermost which
carries them only sporadically.

Straight pedicellariae roughly of two sorts: (1) Larger, pyriform or ovoid,
dentate or unguiculate ones scattered over the surface of the body here and there
along the furrow margin, and on the oral plates. (Pl. 71, figs. le, 1f.) These appear
to be common only in the largest specimens where they measure from 0.8 to 1.2 mm.
(2) Lanceolate blunt forms with or without a few terminal teeth, and much smaller
than the above are scattered over the body and along the furrow margin. In most
specimens these are the prevalent type and vary greatly in size, from the length of
the crossed pedicellariae to 0.5 or 0.6 mm. On big specimens (R 50 mm. or more)
they intergrade in size with (1). (Pl. 71, figs. 1d, le.)

Madreporic bodies usually two, in opposite interradii near the margin of disk.
Some but not all small, 6-rayed examples have one madreporite. Ordinarily fission
takes place on a plane between the two madreporites, but there are a number of
exceptions, as for instance the large 7-rayed example, Plate 70, Figure 1, station 3548.
Specimens in process of acquiring new rays usually have the second madreporite
undeveloped on the regenerating part of the disk.

Type locality —Grand Manan, New Brunswick (at mouth of Bay of Fundy).

Distribution —On the Atlantic coast of North America “it is common from low-
water mark to 100 fathoms in the Bay of Fundy and off the coast of Nova Scotia.
Dredged at more than 100 stations between north latitude 46° 50’ and 35° 12’ 30’’.
Off Cape Hatteras and off South Carolina it is common in 16 to 50 fathoms. Common
south of Marthas Vineyard in 50 to 150 fathoms. In depth its range is 0 to 229
fathoms; in one case recorded from 435 fathoms, off Delaware, and once from 1,253
fathoms.” (Verrill, 1914, p. 148.) The species has been taken on the east, south,
and west coasts of Greenland as far north as latitude 80°; from Ellesmere Land,
Jones Sound, Cumberland Gulf; Iceland; Jan Mayen; Spitzbergen; Murman coast;
Barents Sea; Kara Sea and Kara Strait; Matochkin Strait, Nova Zembla. (See
Von Hofsten, 1915, p. 57.)

In the north Pacific region the species has been taken in 33 to 350 fathoms from
southern Bering Sea to Lynn Canal, southern Alaska.

It has been recorded by Marenzeller (according to Von Hofsten, 1915), from
1,000 to 1,200 meters, Japanese Sea.

The species is probably completely circumpolar in distribution, but it is to be
noted that no specimens have been taken in northern Bering Sea, nor from the Arctic
Ocean just north of Bering Straits—regions which have yielded other species of arctic
sea stars.

Specimens examined.—Seventy-four.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 159

Specimens of Stephanasterias albula examined

i | }
i | Bottom) N®™ | num
a = Locality Depth | Rottom temper- —- ber of
: ature | mens | ™*
| i Fathoma| oF
5223 | Bering Sea, S4* 20’ 15’ N., 165° 32 W._.. 2... & | Bleck pebbles..................- 2 7
39

3257 | Bering Sea, 54° 40’ N., 165° 32’ W.................... j 81 | Gray sand, gravel. ..............

3311 | Bering Sea, 53° 59’ 36” N., 106° 20’ 43” W__.
3331 | Bering Sea, 54° 01 40’ N.., 166° 48’ 50” W....

348 | Bering Sea, 4° 44’ N., 165° 42” W..........-..-...--

i |
4258 | Vicinity Funter Bay, Lynn Canal, Alaska.......... | 300-313 | Mud | 412)

OD me ee ee ee ee ee ee ee
ewmargaraewtweananeeao- ee

Sli asad itt dea tabiels i ‘
| i

4292 | Shelikof Strait, Alaska (off Cape Uyak). ............ 102-04 | Blue mud, fine sand........ ----| 30.8)
4777 | Petrel Bank, Bering Sea, 52° 11’ N., 179° 40’ E...... | 5243 | PEPE IN esac oeerees |
4778 j....- MDA ctiiqwesiindntantasibapinsetinaten tin nithaitehe ndheis 43-33 , Fine black gravel...............).-.----- |
4779 |... eee oe ga sere et aie amg &4-8 | Broken shells, pebbles, sand...../........|
4787 | Off Copper Island, Commander Islands... .........- 66-07 | Gremivand ss Lee BS |
|
4788 |..... UBS ak be ele 8 a 57-86 |....- aa dada edoeacianc ale Anas
4700 | Of Cape Monati, Bering Island, Commander 1h 1A oe aaa ee cali ceeck |
Islands.

4701 j..... I Seneca teal thd treres ots pee ie caper toaet estes ti | AI AEE Entnantsntenaehwaembecbual ces neste 10 5

WROTE oO aS ae BG IS a) an) et We ee ea Ca "4

2 7

| “ s

a tia as ee eae ion d uted prasliinee Rien ameail dieidvuaanvetnentivaucasiicowsDcnsahdnte tes } 1° ®

| Unalaska (Captains Harbor, Dall).................. |ovssteste|eneeneneneecneeenesseenenencereene|teeneene | 1 7

! Divided.

Genus APHANASTERIAS Fisher

Aphanasterias Fisner, Ann. Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 601.—Type, A.
pycnopodia Fisher.

Diagnosis.—Resembling a large 5-rayed Stephanasterias in general appearance,
but differing in having compound papulae, very regularly diplacanthid adambulacrals,
diplacanthid inferomarginals, monacanthid superomarginals and abactinals, extremely
compressed ambulacrals, and in lacking fissiparity. No actinal plates (except pos-
sibly a few interradial rudiments); marginals small, on ventrolateral border of ray;
abactinal skeleton irregularly reticulate, the carinals arcuately and rather irregularly
four sided, and imbricated in a definable series; dorsolaterals very irregular trans-
versely elongate; adambulacrals and ambulacrals much compressed, the adoral
carina well developed, with four to six pairs of plates involved; gonads opening inter-
radially and dorsally; crossed pedicellariae in circlets around the abactinal and
superomarginal spinelets; in tufts on the inferomarginals; conspicuous unguiculate
straight pedicellariae; no pedicellariae whatever on the adambulacral spines,

Remarks.—To the differences separating this genus from Stephanasterias might
well be added “‘a better developed adoral carina.” The small specimens of Stephan-
asterias ordinarily found in collections have only the first postoral pair of adambu-

160 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

lacral plates in contact interradially, but a large specimen which I have (station 3548,
Bering Sea) has three pairs in contact.

One of the best differences seems to be the compound structure of the papulae,
to which the finely papillose appearance of the interspinal spaces is due. In Ste-
phanasterias the papulae are simple papillae, not at all subdivided.

Aphelasterias japonica superficially resembles Aphanasterias pycnopodia but its
rays are stouter. Its spinelets are more like those of Stephanasterias. There are
three or four inferomarginal, and the same number of superomarginal spines in ver-
tical (transverse) combs, the abactinal spinelets are in small groups, and the papulae
are simple. The straight pedicellariae are inconspicuous and lanceolate, and the
crossed pedicellariae are quite small (0.16 to 0.18 mm. long; pl. 72, fig. 5). As in
Aphanasterias the adambulacral plates are regularly diplacanthid, the ambulacrals
are much compressed, and the adoral carina is narrow and in large specimens com-
posed of three to five pairs of plates. But the resemblance of this feature to that of
Aphanasterias pycnopodia is not strong, since the carina is much more constricted in
Aphelasterias and the first and second adambulacrals are longer than the succeeding.

In Stephanasterias the gonads open at the base of the ray, between the supero-
marginal and inferomarginal plates; in Aphelasterias they are attached to the inner
surface of the superomarginals a short distance from the base of the ray (a trifle less
than r), but the ducts open just above the superomarginals. In the present genus the
attachment point is close to the interbrachial angle, above the superomarginals (which
turn upward rather abruptly as they approach the angle) and at the dorsal entrance
of interbrachial or axillary channel. In the other two genera the aperture is low on
the side of the ray.

APHANASTERIAS PYCNOPODIA Fisher

Plate 70, Figures 6, 7; Plate 72, Figures 1-3
Aphanasterias pycnopodia Fisopr, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 601.

Diagnosis.—Rays five; R 76 mm., r 7 mm., R=10.8 r; breadth of ray at base,
9 or 10 mm.; disk very small, rays very slender and gradually tapering to an atten-
uate extremity; abactinal surface well arched; marginal plates on the ventrolateral
margin; tube feet very crowded, sometimes five or six across the furrow; actinos-
tome small, sunken; adoral carina well developed; abactinal spinelets small, solitary,
scattered, except the carinals which form regular series; superomarginals forming
one, inferomarginals two, and adambulacral spinelets two regular longiseries; large
unguiculate straight pedicellariae; crossed pedicellariae in small circles around the
abactinal and superomarginal spinelets, and in tufts on the inferomarginals; gonads
opening dorsally.

Description.—The ray has few trenchant features which may be used as recog-
nition characters. The well arched abactinal surface is beset with short, conical or
conico-cylindrical, sharp or bluntly pointed, spaced spinelets, one to a plate and
averaging about 0.5 to 0.6 mm. in length. Those of the carinal series form a definite
line and are spaced about their own length apart, and are stouter than the adjacent
dorsolateral spinelets, which increase slightly in size toward the superomarginal
series. The dorsolateral spinelets are more widely spaced, do not form regular series,
and about four or five can be counted in the width of each dorsolateral area, at base

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 16]

of ray. There is one superomarginal spine, a little longer than the carinals, and just
below it two slightly longer, tapered inferomarginal spinelets, the upper the shorter.
The three spines form three longitudinal series, the two inferomarginal being on the
ventrolateral margin of the ray, the superomarginals spaced above them by about
the length of a spine. Each abactinal and superomarginal spinelet is surrounded by
a collar of crossed pedicellariae borne on a sheath while the inferomarginals carry a
small cluster, that of the outer being the larger. It is often absent from the inner or
lower spine. Scattered irregularly among the abactinal and marginal spines are
unguiculate straight pedicellariae sometimes longer than the spinelets. A relatively
huge one is found in each actinal interradial area.

The abactinal skeleton is of the irregularly reticulate type. The carinals are
irregularly and the marginals fairly regularly four lobed and directly imbiicated
except in the case of the proximal carinals which may have an internal connecting
ossicle. The dorsolaterals are quite irregular. Actinal platesabsent. (PI. 72, fig. 1.)

Papulae compound, four to eight small papillae to a cluster and two to four
clusters to a mesh, irregularly distributed. Each cluster represents a subdivided or
compound papula. Intermarginal areas with one cluster. The papulae fill in the
spaces between the circumspinal crowns of pedicellariae and give the surface a
minutely papillose appearance.

Adambulacral plates juxtaposed closely to inferomarginals, very short, so that
the consecutive pairs of spines are closely in contact. The spines, which are slender,
slightly tapered, and blunt, are nearly equal, a little shorter but very much slenderer
than the lower inferomarginal spine, and form two regular longiseries on the furrow
margin. The proximal spines measure 1.25 mm. A cordon of large pedunculate
lanceolate straight pedicellariae (proximally 0.95 mm. long) occupies the furrow face
of the adambulacrals. They sometimes extend between the furrow spines, but the
latter carry no pedicellariae whatsoever.

The actinostome is small and constricted, and the adoral carina rather well
developed, consisting of about five postoral pairs of adambulacrals joined on the
interradial line. Mouth plates small, compressed, each with two rather stout,
flattened, blunt spinelets, the inner close to the mouth. At the base of the inner
spinelet is a lanceolate straight pedicellaria, and frequently a larger unguiculate one
(with two claws to a jaw) occupies a position between the suboral and actinostomial
spinelet (one to each oral angle).

Ambulacral ossicles very much compressed; ambulacral pores very narrow slits,
the two mesial rows not exactly straight, so that the tube feet appear in more than
four series, in the broadest portion of the ray.

Crossed pedicellariae, distributed as indicated above, are of fairly uniform size
(about 0.3 mm.), though small ones (0.22 mm.) are found scattered here and there
among the larger. They lack enlarged teeth on the terminal expanded portion but
have numerous small denticles on the shank.

The straight pedicellariae of the abactinal and lateral portions of rays and disk
are very obtuse as seen in profile, and from two to three times as long as broad. The
jaws are scarcely broadened distally and end in one to three prominent claws. Often
one claw fits between two of the opposing jaw. The largest abactinal are about 1
mm. in length. The large actinal intermediate pedicellaria (1.5 to 1.7 mm.) has

162 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

heavier, distinctly spatulate jaws with two or three interlocking claws. The adam-
bulacral pedicellariae attached by fleshy peduncles to the furrow face of the plates are
lanceolate, without teeth, except incipiently, on the adoral carina. In life the pedun-
cles are probably capable of considerable extension.

Madreporic body 3 mm. in diameter, with pores rather than striae, and situated
a little nearer to interbrachial angle than midway of r.

Gonads small, opening in interbrachial angle above the superomarginal plates.
Ampullae single, pyriform, but somewhat two lobed when contracted.

Type—Cat. No. E 1505, U.S.N.M.

Type locality—Station 2847, vicinity of Shumagin Islands (55° 01’ N., 160°
12’ W.), 48 fathoms, fine gray sand, bottom temperature 42° F.; one specimen.

Remarks.—This species resembles a large, sparsely spined Stephanasterias albula
with two instead of three inferomarginal spines, one instead of three or four supero-
marginal spines, and two instead of three adambulacral spines. The abactinal
spinelets of Stephanasterias are in little groups which tend to form transverse rows,
in large specimens especially. Most North Atlantic examples of Stephanasterias
albula are quite small, but I have a specimen from 91 fathoms, Bering Sea (station
3548) in which the longest rays are 58 mm. The large straight pedicellariae of
Stephanasterias have more numerous teeth and are of different form.

Genus PISASTER Miiller and Troschel

Asterias (part) Branpt, Prodromus, 1835, and authors.

Pisaster Miituer and Troscuet, Archiv f. Naturgesch. 6 Jahrg, vol. 1, 1840, p. 367; System
der Asteriden, 1842, p. 20. Type, Asteracanthion margaritifer Miller and Troschel
(=Asterias ochracea Brandt).—A. AGassiz, Mem. Mus. Comp. Zodl., vol. 5, 1877, p.
96 (citation) —Fisurr, Smithsonian Mise. Coll., vol. 52, 1908, p. 89; Zool. Anz., vol.
33, 1908, p. 358.—VeErriut, Amer. Journ. Sci., vol. 28, 1909, p. 63; Shallow-water
Starfishes, ete., 1914, p. 67.—Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 556.

Asteracanthion (part) Mii~umr and Troscuxt, System der Asteriden, 1842, p. 20.

Calliasterias Fewxers, Bull. Essex Inst., vol. 21, 1889, p. 38. Type, Asterias exquisita de
Loriol (= Pisaster giganteus young).

Diagnosis —Large, heavy-ossicled, monacanthid Asteriinae haying an irregularly
articulate abactinal skeleton; few to numerous short subcapitate abactinal spines;
two to five longiseries of spiniferous actinal plates; a small, deeply sunken actino-
stome; long, upeurved adoral carina; extremely compressed ambulacral ossicles;
unique furcate straight pedicellariae, each jaw of which ends in one long and one
short hyaline blade; no pedicellariae on adambulacral spines; gonads opening
dorsally.

Description.—Abactinal skeleton irregularly reticulate; the carinals and mar-
ginals at first four lobed; one or more intermarginal ossicles; in old specimens second-
ary ossicles are developed between consecutive inferomarginal and actinal plates as
well as between the plates of the transverse series; normally two inferomarginal and
one actinal spine to a plate, each with a thick pad of crossed pedicellariae on outer
side; abactinal spines variable, but uniformly short, subconical to subglobose, the
more or less specialized distal portion longitudinally scored; in old specimens there
may be more than five longiseries of actinal plates, and the skeleton is strengthened
by the addition of new ossicles on the coelomic side. Actinostome small, deeply
sunken, the adoral carina composed of upward of 15 pairs of contiguous adam-

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 163

bulacrals; proximal end of furrow petaloid in form; ambulacral ossicles very
much compressed so that the tube feet stand in six rows at the widest part of
furrow; interbrachial septum strongly calcified, and extending about two-thirds
toward center of disk; aperture from arm coelom into disk small (about twice
diameter of madreporite in ochraceus); a series of transverse, vertical, superactinal
partitions or buttresses connects the adambulacral and inferomarginal plates, the
spaces between them forming a series of coelomic cavities above the actinal plates.
Papulae numerous, slender, in clusters, the actinal very long; the clusters originate
from a common hernialike swelling of the skin which, with bases of papulae, is
armed with tiny lanceolate pedicellariae. (Pl. 77, fig. 5.) Stomach eversible;
pyloric coeca long, bulky; intestinal coecum with slender irregular lobes; gonads,
extensive racemose tufts, opening dorsally.

Small fureate straight pedicellariae (usually mingled with the coresed) are
peculiar to the genus and consist of a basal piece, longer than broad, and two curved
jaws each ending in two thin hyaline blades, characteristic for each species. One
blade is always much longer and broader than the other. Large sessile, ovoid,
*stone-hammer”’ pedicellariae with narrow to broad ovate jaws and terminal inter-
locking teeth are found scattered over the body but especially in the intermarginal
and in the actinal interradial channels; very small lanceolate dermal straight pedicel-
lariae are common especially among the papulae and in long pedunculate clusters
attached to the furrow margin, often surrounding one or more large lanceolate,
toothed pedicellariae. Crossed pedicellariae, in dermal tufts or surrounding spines,
are numerous and lack any conspicuously enlarged terminal teeth. (Actinostomial
ring, pl. 76, fig. 1.)

Remarks.—The most characteristic features of this genus are the furcate pedicel-
lariae, which are unique; the broad actinal area; and the long adoral carina bent
upward to the deeply sunken, small, actinostome. These are all indications of high
specialization, as are the very numerous, crowded, ambulacral ossicles and small
mouth plates. If we assume as primitive a type similar to Hydrasterias, with
open oral angle (that is, without adoral carina), with biserial tube feet and
uncrowded ambulacral ossicles, no actinal plates, and few, regular, dorsolaterals,
then Pisaster is one of the most specialized genera in the family. It is moreover
restricted in habitat, as specialized genera are likely to be, being confined to the
Pacific coast of North America.

Verrill (1914, p. 67) lays considerable emphasis upon the large unguiculate
pedicellariae as peculiar features of this genus, but this type is widely distributed
through the family and is in no way characteristic of Pisaster alone. If these
pedicellariae were diagnostic of Pisaster they would exhibit peculiarities for each
species as do the unique fureate pedicellariae. Nothing similar to the latter have
been described for any sea star. No mention of them is made by Verrill, though
they are very numerous in his Pisaster papulosus (=brevispinus) as well as in other
forms, except southern California races of P. ocharaceus and P. giganteus where they
are rather scarce.

The species are extremely variable. As is often the case when abundant mate-
rial is available, specific distinctions become less distinct. Three species are sharply
differentiated, without the least indication of intergradation. All the other tax-

164 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

onomie divisions have been arranged under one or another of these three—ochraceus,
giganteus, brevispinus.

In the left column are the forms recognized in this synopsis and in the right
column the names which the same forms bear in Verrill’s Shallow-water Starfishes.

Pisaster ochraceus forma ochraceus_---------------- Pisaster ochraceus (p. 69).
Pisaster fissispinus (p. 76).
Pisaster ochraceus f. nodiferus.-.------------------ Pisaster ochraceus var. nodiferus (p. 72).
Pisaster ochraceus f. confertus.--.-..--.------------ Pisaster confertus (p. 73).
Pisasier ochraceus segnis..----------==----+--~=---= Pisaster ochraceus.
Pisaster brevispinus f. brevispinus__-------- Red SN Pisasler brevispinus (p. 77).
Pisaster brevispinus f. paucispinus__--------------- Pisaster papulosus (p. 91).
Pisasler paucispinus (p. 98).
Pasaster Giganleus ae ee ene Pisaster liitkenii (p. 83).

Pisaster litkenii var. australis (p. 88).
Pisaster giganteus (p. 97).

Pagasters Qi ganteus (COP iUQULS eee Pisaster capitatus (p. 81).

Pisaster grayt (p. 97).

Pisaster grayi Verrill is a phantom species, the origin of which is traceable to a
mistake in Perrier’s Révision des Stellérides, page 68. Perrier there described the
types of Gray’s Asterias katherinae (=Leptasterias polaris katherinae) as having
monacanthid adambulacrals, whereas the types (which can be fixed with perfect pre-
cision) all have diplacanthid adambulacrals. Verrill accepted Perrier’s description,
but inferred that these supposedly monacanthid specimens were a second set of
6-rayed asterids which Perrier had mistaken for the types of Asterias katherinae. No
such second set exists, however. Pisaster grayi is therefore without status. The
name must be regarded as a synonym of Leptasterias polaris katherinae.

PISASTER OCHRACEUS (Brandt)

Asterias ochracea BranpT, Prodromus, 1835, p. 69 (Sitka).—Sriwrson, Boston Journ. Nat.
Hist., vol. 6, 1867, p. 527, pl. 23, fig. 2—Vzernitt, Trans. ‘Conn. Acad. Sci., vol. 1,
1867, pp. 325, 326—Prrrinr, Rév. Stell., 1875, p. 70—Wuttraves, Trans. Royal
Soe. Canada, vol. 4, 1887, p. 116—Agassiz, Mem. Mus. Comp. Zodl., vol. 5, 1877,
p. 96, pl. 11, figs. 1-7—StapENn, Challenger Asteroidea, 1889, pp. 566, 826.—H. L.
Crark, Bull. Mus. Comp. Zoél., vol. 51, 1907, p. 67, pl. 6, fig. 3.

Asterias janthina Branpt, Prodromus, 1835, p. 69 (Sitka).

Asteracanthion margaritifer MwuLuER and TroscHEeL, System der Asteriden, 1842, p. 20
(Sitka 2).

Asterias conferta Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 263 (Puget Sound).—
VeRRILL, Trans. Conn. Acad. Sci., vol. 1, 1867, p. 326—Prrrier, Rév. Stell., 1875, p.
71.—S.iapeEn, Challenger Asteroidea, 1889, pp. 566, 820.—Brtu, Proc. Zool. Soc., 1881,
p. 494.—WuitrAves, Trans. Royal Soc. Canada, vol. 4, 1887, p. 116—Der Lortot,
Mém. Soe. Phys. et Hist. Nat. Genéve, vol. 32, pt. 2, 1897, p. 171, pl. 3, figs. 1-lg.

Asierias fissispina Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 264 (Shoalwater
Bay, Wash.).—VERRILL, Trans. Conn. Acad. Sci., vol. 1, 1867, p. 326.

P{isaster) ochraceus VERRILL, Amer. Journ. Sci., vol. 28, 1909, p. 63.

Pisaster ochraceus Fisher, Smiths. Misc. Coll. vol. 52, 1908, p. 89—Vxrrritt, Amer. Nat.
vol. 43, 1909, pp. 542, 544, 548; Shallow-water Starfishes, 1914, p. 69, pl. 21, figs. 1, 2;
pl. 49, figs. 3-3d; pl. 56, figs. 3, 3a—H. L. Cuark, Bull. Amer. Mus. Nat. Hist., vol. 32,
1913, p. 203 (San Diego).—Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 557,
pl. 19, fig. 1; pl. 20, figs. 1, 2, 4.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 165

Pisaster ochraceus var. nodiferus Vexnitt, Shallow-water Starfishes, 1914, p. 72, pl. 56,
figs. 3, 3a (Monterey to Sitka).
Plisaster] confertus Venniti, Amer. Journ. Sci., vol. 28, 1909, p. 63.
Pisaster confertus Verniii, Shallow-water Starfishes, 1914, p. 73, pl. 38, figs. 1, 2; pl. 53,
fig. 2.
Balentap fissispinus Vernity, Shallow-water Starfishes, 1924, p. 76, pl. 39, figs. 1, 2.
Diagnosis. —A usually thick-rayed Pisaster with numerous small subcapitate,
striated abactinal spines arranged in a very irregular reticulate pattern on dorsal and
lateral surface of rays, or in detached convex groups, especially on outer part of ray.
Actinal spines longer than the abactinals, clavate, striate, in upward of six longitudi-
nal series, Furcate pedicellariae with the undivided basal portion of jaw about
equal in length to the longer of the two hyaline blades (which has the upper edge
decidedly uneven in most cases). (Pl. 74, figs. 1, 2.
Description.—The species is so variable that a composite description is impossi-
ble. There are three fairly well marked formae which freely intergrade. Two of
these have been heretofore ranked as species.

Forma OCHRACEUS (Brandt)

Plate 73, Figure 1; Plate 74, Figures 2, 2a; Plate 75, Figures 1-4, 5a, 5b; Plate 76, Figures 1, 4, 5, 5a,
5b, 6, 6a; Plate 79, Figures 1,6; Plate 80; Plate 81, Figure 3; Plate 83, Figure 1; Plate 85, Figure 1

This form stands between confertus, with small spinlets, and nodiferus character-
ized by stout subglobose abactinal tubercles in more or less isolated convex groups or
heaps. It is assumed to be the typical form, and it was so regarded by Stimpson when
he described confertus as a distinct species. The only specimens of ochraceus from
Sitka, the type locality, examined by me (pl. 81, fig. 3) are referable to this forma.
But Asterias janthina Brandt, also described from Sitka, is almost certainly the com-
mon purple phase of confertus so conspicuous at Departure Bay, British Columbia.

An average specimen from Monterey Bay measures R 140 mm., r40 mm., breadth
of ray at base, 40 mm. Specimens reach twice this length of ray; rays may be rela-
tively broader. Sitka example: R 115 mm., r 36 mm., breadth of ray at base, 43 mm.

The disk is normally convex or rather high arched owing to the deeply sunken
actinostome; the rays are arched, normally subcireular in cross section; the actinal
surface, viewed from below, slopes downward to the border of the narrow ambulacral
furrow, which broaden aburptly at base of ray. The abactinal skeleton extends low
on the side of the ray, the superomarginal plates being near the actinal surface and
separated from the inferomarginals by a relatively narrow but well-defined channel.
The entire skeleton is very robust and rigid, the dorsal forming an irregular reticulum
on either side of the rather irregular carinal series, The skeletal ridges are made
up of very numerous small irregular plates of a great variety of shapes and sizes,
several layers thick in mature specimens, bound together by extremely tough tissue.
The intervals are sunken, and are subdivided by much lower trabeculae not externally
visible even in dried examples unless treated with caustic potash. Between the sec-
ondary trabeculae, defining the actual intervals of the skeleton, emerge the numerous
clusters of papulae, very conspicuous in life. Among these and surrounding them on
the slopes of the higher ridges are very numerous clusters of crossed and furcate,
straight pedicellariae, the latter predominating in most cases. Along the meandering

63160—30——12

166 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

summit of the higher ridges, above this closely packed mass of papulae and pedicell-
ariae, are the stout small spines with subglobose striate ends (pl. 75, figs. 5a) arranged
usually in single file, sometimes by twos on the carinal series, or forming small groups
at the nodes. These tubercles are usually nearly uniform but variations occur in
which there is considerable disparity in size.

The variations in the pattern formed by the abactinal spines are as numerous
as the individuals of the species, typical examples being figured.

The superomarginal spines are not different from the abactinal though occa-
sionally a little larger than those directly above them. The plates carry usually one
spine, sometimes two, especially at nodes. Some variations have a considerable num-
ber of the plates with two marginal spines, or one accompanied by two smaller spine-
lets. A few accessory spinlets appear sometimes on the intermarginal plates.

Inferomarginal plates with usually two spines in an oblique series, with upward
of six similar actinal spines between them and the adambulacral system. Actinal
spines one to a plate. These spines are longer and more clavate than the abactinal,
the ornate fluting being interrupted on the outer side for the attachment of a dense
cluster of crossed pedicellariae. The actinal armature is dense and efficient. In life
very long slender pointed papulae extend far beyond the spine tips, to such an extent
as to nearly hide the armature on the proximal part of the ray.

Adambulacral plates inconspicuous, narrow, with a single slender, tapering, blunt,
terete or flattened spinelet. The spinelets are fairly regular, very closely placed,
about as long as the inner actinals and they increase in length over the area where the
furrow is broadened, at base of ray, to decrease again on the long adoral carina.

The adoral carina along its inner half is narrower than the mouth plates and is
composed of upward of 10 pairs of plates. It bends downward (as viewed from the
oral surface) to the oral angle. (PI. 75, fig. 2.) The very small actinostome is guarded
by the stout tapering oral spines, which are stouter than the adjacent adoral spines.
One or two stand on the free margin and a longer suboral on the middle of the surface.
(Pl. 75, fig. 2; pl. 79, fig. 1.) Less often there are three flattend actinostomial spinules.
Length and thickness is extremely fariable even in specimens from the same region.

Crossed pedicellariae (pl. 73, figs. 1, 1a): Abundant, mixed in varying proportions
with fureate straight, on abactinal plates and, practically without furcate straight, in
dense clusters on inferomarginal and actinal spines. They are smaller than in typical
nodiferus being 0.27 mm. long but differ in no important details (Sitka, Monterey).

Straight pedicellatiae: (2) The characteristic Pisaster bifid furcate pedicellariae
are very numerous abactinally, and sometimes intermarginally. Clusters composed
almost entirely of them are found in the bottom of the hollows among the abactinal
papulae. Near the bases of the spines they accompany the crossed form. They
measure 0.27 mm. broad by 0.22 high (Sitka); 0.25 by 0.25 mm; 0.29 by 0.3 (Monterey)
(Pl. 74, fig. 2; pl. 79, fig. 6.)

(b) Undifferentiated lanceolate, of very small size (many as small as 0.13 mm.
long), are scattered over all the papular areas. These are similar to Plate 76, Figure
6a. Clusters of tiny lanceolate pedicellariae surrounding a large one, at the end of long
peduncles, are distributed all along the furrow face of the adambulacral plates—not
on all the plates, but here and there. (PI. 76, figs. 6, 6a.) In some dried specimens
these appear, superficially, to be attached to the furrow spines. In no instance has

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 167

a pedicellaria been found attached to a furrow spine. The central pedicellaria is com-
monly 1 mm. long and is sometimes more robust than Figure 6.

(c) On all surfaces of the body but especially actinally and intermarginally are
found the heavy, subconical, denticulate pedicellariae whose exact form is shown on
Plate 76, figures 4, 5. Figures 2, 3, and 7, although representing forma confertus are
also typical of some specimens of ochraceus. The largest are found on the actinal
interradial angle and commonly measure 1.5 mm. in length. The abactinal are
commonly 0.5 to 1 mm. long. In some examples, without apparent reference to
locality (Monterey, Crescent City, Calif.) the denticulate pedicellariae are rather
scarce abactinally.

Madreporic body, conspicuous, situated about one-third r from center of disk;
striae fine, branched, irregularly centrifugal.

Color in life —There are commonly three principal color phases—brown (yellowish
brown to dull sepia); brownish purple to dull purple; rather bright ochraceous yellow.
The brown phase varies to olive, mottled with dull gray green, and also intergrades
through brownish purple with the dull bluish purple phase. The latter is less vivid
than the color of forma confertus. The yellow phas? is less common at Monterey than
the brown and purple-brown.

Distribution —This typically intertidal form has been taken as deep as 48 fathoms
off Monterey, Calif. It is the typical form of the open ser coast, from Sitka to Point
Sal, Calif., where confertus appears never to occur. It occurs less commonly, along
with confertus, in the protected waters of Puget Sound and tributaries, and in the
many quiet bays and inlets opening upon the remarkable channel which separates .
Vancouver Island from the mainland. In these waters confertus is the dominant form
but intermediates frequently occur. It is absolutely impossible to allocate all speci-
mens since the differences between the formae are relative only.

Remarks.—The type of Stimpson’s Asterias fissispina, from Shoalwater Bay,
Wash., has been carefully examined. The furcate straight pedicellariae are typical
for Pisaster ochraceus. The specimen was dried without first being preserved in aleo-
hol so that the rays have become unnaturally flattened and the spinebearing ridges
of the abactinal skeleton appear to outline rather large areas. But in P. ochraceus
there is wide variation in this respect. The ventral spines have a deep groove on the
outer side. Some of these and some of the dorsolaterals are slightly bifid or mal-
formed at the tip, not an uncommon occurrence in old animals. There are no grounds
for regarding this other than as an individual variant of ochraceus—one reason per-
haps that no other specimens have been found. (For figures see Verrill, pl. 39.)

Dr. A. Djakonov, of the museum of the Academy of Sciences, U. R. S. S., kindly
undertook to find the type of Brandt’s Asterias ochracea. This involved reviewing
all the specimens of Pisaster in the museum. Doctor Djakonov writes that the real
type is not in existence. There are several specimens collected by Wosnessenski,
including two resembling nodiferus from California, 1841; and three of forma
ochraceus, without locality. There are also two specimens well preserved in alcohol
of a variety near nodiferus, labeled by Brandt “ Asterias margaritifera—Siteha.”
There is no type existing of Asterias janthina Brandt.

168 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Forma CONFERTUS (Stimpson)
Plate 73, Figure 2; Plate 74, Figures 1, 1a, 1b; Plate 75, Figure 5; Plate 76, Figures 2, 3, 7; Plate 81,
Figures 1, 2; Plate 82, Figures 2, 3, 4; Plate 86, Figures 2, 4, 7
Asterias conferta STIMPSON.

Pisaster confertus VERRILL.
Pisaster ochraceus forma confertus Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 559,

pl. 20, fig. 1.

The type of Stimpson’s Asterias conferta was taken at Simeahmoo, Puget Sound,
by Dr. C. B. Kennerly (No. 1275 U.S. Nat. Mus.). I have examined it several times
and it is figured by Verrill. (1914, pl. 38.) A number of my specimens from near
Tacoma, Wash., practically duplicate the type, but others from Departure Bay,
Union Bay (Bayne Sound), and Boundary Bay, British Columbia, have the char-
acteristics of the variety in an accentuated form. In these the small dorsal spinules
are much more numerous and are three or four, sometimes upward of seven or eight,
to a plate along the carinal series and on the stellate apical system of the disk. weil:
81, fig. 1; pl. 82, fig. 2.)

Confertus intergrades with ochraceus (Departure Bay, British Columbia, the
sounds of Washington) and it also develops acervate varieties less distinct but parallel
to nodiferus. (Pl. 82, fig. 3.)

In forma confertus the abactinal spinelets are smaller and frequently more numer-
ous than in ochraceus (pl. 75, fig. 5), the majority only 0.8 or 0.9 mm. long and rela-
tively slender. Some specimens (as those from Departure Bay and Boundary Bay,
British Columbia, pl. 81, fig. 1; pl. 82, fig. 2) carry three to eight spinelets on most of
the carinal and primary apical plates. Other specimens have the spinelets in single
file, or with the slight irregularity in number normal to ochraceus. (Pl. 82, fig. 4.)
The superomarginal plates usually carry three or four spinelets giving the armature
a different and more crowded appearance than that of ochraceus. In some cases the
actinal spines are slightly slenderer than those of ochraceus; in others there is no
difference, and I am unable to find any constant actinal features by which confertus
can be distinguished from ochraceus.

Crossed pedicellariae (pl. 73, fig. 2): These are similar in size, form, and distribu-
tion to those of ochraceus.

Straight pedicellariae: (a) The furcate pedicellariae are very numerous but
appear to be exactly the same as those of ochraceus. That shown on Plate 74, Figure
la, is intermediate between the furcate and the ordinary broadly lanceolate, undif-
ferentiated type. The larger measure 0.28 by 0.28 mm.

(b) The undifferentiated lanceolate pedicellariae are very numerous, rather
more so than in ochraceus, especially abactinally.

(c) The denticulate pedicellariae are similar to those of ochraceus but often more
conspicuous owing to the smallness of the spines. They are usually more numerous,
but there is no constant difference inform. Especially large ones (1 to 1.5 mm. long)
are found on the actinal interradial angle, and along the intermarginal channel. (PI.
76, figs, 2, 3, 7.)

As in ochraceus, both thicker and slenderer rayed specimens occur. At Depar-
ture Bay, British Columbia, where the form is very common, mature examples range
from 100 to 170 mm. major radius. Larger examples of course occur in favorable
situations.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 169

Color in life-—The prevalent color at Departure Bay, British Columbia, is purple.
This analyzes as follows: The small spines, whitish; adjacent ruffs of small crossed and
fureate pedicellariae rich aster purple or pansy purple, this area widening as the violet
papulae are contracted. When the papulae are fully extended the animal is bluer;
when contracted, the aster purple predominates. This color extends to edge of fur-
row. Tube feet, pale yellow.

Another less common phase—vinaceous rufous, reticulated lighter along the
spiniferous ridges, paler below. Sometimes varied with orange rufous.

Another less common phase—madder brown, papulae very dark maroon, the
color extending to furrow; tube feet, pale yellow.

Specimens from Union Bay, Bayne Sound, British Columbia, are labeled “ bluish
purple.”

A specimen from Boundary Bay, British Columbia, intermediate between the
specimen figured (pl. 81, fig. 1) and the Sitkan example of ochraceus (pl. 81, fig. 3)
was pansy violet with white spines (Ff. W. Weymouth).

Distribution —In the list of localities it will be seen that this form is practically
confined to the quiet bays and sounds of Washington, British Columbia, and southern
Alaska. It is difficult to escape the inference that the characteristic small spinelets
of the abactinal area are correlated with quiet water, but that this is not the only
factor is evidenced by the presence, along with confertus, of forma ochraceus and
nodiferus, the latter found on open coasts and also in deep water (Monterey Bay).
There is nothing approaching a sharp line of distinction between ochraceus and
confertus.

Type of forma confertus—Cat. No. 1275, U.S.N.M., Puget Sound (Simeahmoo),
Wash.

Forma NODIFERUS Verrill

Plate 73, Figures 3, 7; Plate 75, Figure 5c; Plate 82, Figure 1; Plate 86, Figures 1, 3, 5, 6, 8, 9
Pisaster ochraceus var. nodiferus VERRILL, 1914, p. 72, pl. 56, figs. 3, 3a.
Pisaster ochraceus forma nodiferus Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 560.

Abactinal spines stout, globose, or depressed-globose, finely striated, larger than
in forma ochraceus (pl. 75, fig. 5c) and congregated in well separated convex groups
or heaps, three or four to a dozen unequal or subequal tubercles to a group. In the
center of the disk there is usually a well defined stellate palisade of spines surrounding
a central group of tubercles and from the points of the star a carinal series of tubercles
often extends half the length of the ray but as often is entirely absent, or represented
here and there by unequal tubercles. The spaces between the groups of tubercles
are usually entirely free from spines, or in intermediate forms have more or less reticu-
lation at base of ray and more complete isolation and prominence of the groups distally.

The actinal surface is similar to that of forma ochraceus.

Crossed pedicellariae (pl. 73, fig. 3) are similar in form to those of ochraceus
proper but a little larger. Forma nodiferus, Monterey, 0.30 to 0.36 mm.; Tongass,
Alaska, 0.31 mm. Forma ochraceus, Monterey to Sitka 0.27 mm. (some as smal!
as 0.25 mm.),

Straight pedicellariae: a. The furcate type resemble those of ochraceus and vary
somewhat in abundance. On a specimen (No. 18146) from Puget Sound, near
Tacoma, they are much more abundant than the crossed; Tongass, Alaska, less

170 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

abundant; Sausalito and Monterey, Calif., much less abundant. In this form there
is a tendency to lose the furcate pedicellariae with decrease in latitude.

b. The lanceolate form appears to be about as numerous as in ochraceus.

c. The large denticulate pedicellariae are very numerous in the northern exam-
ples, less numerous in the southern. They are rather variable in form; some of the
proximal intermarginals develop 8 to 10 teeth whereas the usual number is 4 to 6.

Color of forma nodiferus follows closely that of ochraceus. A number of speci-
mens from Monterey Bay were colored: Ground tint grayish olive to tea green and
light artemisia green, with large patches of rich olive brown, raw umber, and Van
Dyke brown.

Distribution of forma nodiferus.—Southern Alaska to northern Lower California
(south of Point Conception as a prevalent form of subspecies segnis).

Type locality of Pisaster ochraceus—Sitka, Alaska.

Distribution of Pisaster ochraceus——Southern Alaska to Point Conception, Calif.

Specimens eramined.—One hundred and seventeen in addition to many hundreds
from Monterey, and very numerous living examples at Departure Bay, British
Columbia.

Specimens of Pisaster ochraceus examined

f. och- | f. con- | f. nodi-

ae Locality | Taceus | fertus | ferus Remarks
Sibi, AUNGiAt 7S SUAT TLE 3d 2 ee RR FRG Se [rears
Revillagigedo Island, Alaska ._.....--.--..---.----- | Ward Cove; T. H. Streets.
Guels Island), Alaska. ---===s--22>-c=sas=es----2-5=2 2 | Near ochraceus; Albatross, 1894.
Tongass, Alaska-_-_..- bee = | | Nichols.

Naha Bay, Alaska
Boundary Bay, British Columbia

l intermediate; Albatross, 1905.
Stanford Coll.; 1 aberrant, F. W. Weymouth, W.

| Laban Thompson.
Union Bay, Bayne Sound, British Columbia---_----|_------- BO) oR Seance
Taylor Bay, Gabriola Inlet, British Columbia --__- 3 Sips 1 intermediate.

Departure Bay (Nanaimo), British Columbia.1____ 4 24 | 17 | U.S.N.M. Coll.; Stanford Coll; Cal. Aead. Sci.
| Coll.; W. K. Fisher, W. F. Thompson; Albatross.

Port Renfrew, Straits Fuca, British Columbia Stanford Coll., J. C. Brown.

Barclay Sound, Vancouver Island_--_-.------- Albatross.

Port: Ludlow>) Wash i22 2059 ene sande ee p! S. Bailey.

Oreas, San Juan Islands, Wash.?_...--...---------- 4 Cal. Acad. Sei. Coll., Ida S. Oldroyd; confertus
coarse spined, near ochraceus.

Friday Harbor, San Juan Islands, Wash___-_.___-_]__--__-- Ghee Cal. Acad. Sei. Coll., Ida S. Oldroyd; rather coarse
spines.

Puget Sound (Simeahmoo)----...._..___--_---__--- 1 | Dr. C. B. Kennerly.

Puget Sound (near Tacoma) -....-_--..-.--------.. 5 Frank Russell, Chas. M. Drake, F. W. Weymouth

BHOSIWALAD Bay, WHSNt-oecc—. oo coerce eee oe Dr. J. G. Cooper, type of fissispinus.

Crescent City, Calif_.__.__-
Trinidad, Calif.......-_._.-

Stanford Coll., J. O. Snyder.
Stanford Coll., F. W. Weymouth.

Humboldt Bay, Calif. ...-__- Chas. M. Drake.
Drake's Bay, Calif-.-.--------- & Albatross.
San Francisco Bay, Calif....__- = = Dr. John Hornung.

San Francisco Bay (Sausalito) -~.......-......2-__- Albatross; seine.
Farallone Islands; Calif:..--.0. 2 52... LkbcLke C. H. Townsend.

Santa Cruz, Calif................ 3 A. Forrer.

Monterey Bay (vicinity Point Pinos Hopkins Marine Station.

Monterey Bay, station 3131_.......- 4 miles north of New Monterey, 48 fathoms, rocks;
intermediate with nodiferus.
EOUUSAl, Oats cen enscteesccreane = aaceneeeeeeeee Carl Hubbs.

1 At Departure Bay, where the Canadian Biological Laboratory is situated, the commonest forma is confertus. There are
fewer, coarser spined specimens indistinguishable from ochraceus, and others, in which the spines form very prominent groups
on the outer part of the ray, near nodiferus. 6 of the young examined are referable to nodiferus; 5 others are intermediate between
nodiferus and confertus.

? The specimens from Orcas are difficult to classify by formae; with the exception of the acervate forma they are usually neither
clearly confertus nor typical ochraceus.

4 The type of confertus.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS~—~ FISHER 171
PISASTER OCHRACEUS SEGNIS Fisher

Plate 73, Figures 4, 8; Plate 75, Figure 6; Plate S4

Pie enn segnis Fisuer, Ann. Mag. Nat. Hist. ser. 9, vol. 17, 1926, p- 560, pl 20,
g. 3.

Diagnosis.—Differing from typical ochraceus in having relatively few furcate
straight pedicellariae and conspicuously larger crossed pedicellariae. Type, R. 143
mm., r 44 mm.; f. nodiferus (Venice, Calif.) R. 124 mm., r 35 mm.

Description—-This is the southern California representative of Pisaster
ochraceus and it parallels exactly Pisaster giganteus capitatus, which also has larger
crossed pedicellariae and much fewer furcate pedicellariae than its northern relative.

Both forma ochraeeus and nodiferus extend continuously to southern California,
the latter apparently predominating at Laguna Beach, Orange County. The differ-
ences enumerated in the diagnosis apply to both formae.

The type of segnis corresponds to forma ochraceus but there is a tendency toward
nodiferus in the arrangement of spines on the outer half of the ray. Exactly com-
parable specimens are common at Monterey. In general appearance there are no
distinguishing features, until the small pedicellariae are closely scrutinized, when the
scarcity of the fureate sort, so numerous in northern specimens, becomes apparent.
The dentate stone-hammer pedicellariae are not very large, nor very numerous except
proximally in the interradial channel. There are five or six actinal spines in a trans-
verse series, in addition to the two inferomarginals.

All of the seven specimens from Laguna Beach are well developed forma nodiferus
or perhaps more properly speaking the southern subforma of this. These specimens
have large subglobular spines (pl. 75, fig. 6) in very convex groups, with wide generally
spineless intervals between. Superficially there is little to distinguish them from the
Tongass specimen of nodiferus which has unusually heavy tubercles. These southern
nodiferus have heavier tubercles than Monterey specimens of forma nodiferus. They
correspond to the heavier spined form of Pisaster giganteus capitatus. These dried
specimens from Laguna Beach, as well as a similar one from Venice, Calif., have
smaller clusters of ‘“‘minor”’ pedicellariae than comparable examples from Monterey
to Alaska, since in southern specimens the furcate pedicellariae, so important in
northern examples, are rather scarce. On this account the spaces between the
groups of spines appear less well armed, although the individual crossed pedicellariae
are larger. This feature will distinguish the southern specimens from the Tongass
example (pl. 82, fig. 1) which was unusually heavy tubercles. The stone-hammer
pedicellariae are stouter than in the type specimen; about as in Plate 76, Figures 5, 50;
occasionally even a little broader. Small, undifferentiated lanceolate pedicellariae
are scarce.

Crossed pedicellariae: As in typical ochraceus there is a difference in size in the
two formae. Type, length of well-developed ped. 0.35 to 0.36 mm.; nodiferus, Laguna
Beach, 0.36 to 0.41 mm, (PI. 73, fig. 4; compare with fig. 3, Tongass, Alaska, no-
diferus.) Comparable specimens of forma ochraceus Monterey to Sitka, 0.25 to 0.27
mm.; forma nodiferus, 0.30 to 0.36 mm.

Fureate straight pedicellariae (pl. 73, figs. 8, Sa) have a diameter of about 0.28
or 0.29mm. The form of the blade (type specimen) is rather nearer that of northern

172 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

nodiferus than northern ochraceus. The undivided basal part of each jaw is relatively
high, as in the northern race. Figure 7, same plate, is a blade from a specimen of
forma nodiferus, Monterey. In segnis the upper edge of jaw blade varies in minor
details, as evidenced by the two specimens figured.

Type.—No. E 1238, U.S.N.M.

Type locality Sixteen miles south of international boundary, Lower California.

Distribution —From type locality to Santa Barbara, Calif., intergrading north
of this point with typical ochraceus.

Specimens examined.—Fifteen.

Laguna Beach, Orange County, Calif., seven, forma nodiferus. Prof. W. A.
Hilton.

Venice, Los Angeles County, Calif., one, forma ochraceus, one forma nodiferus.
Prof. A. B. Ulrey.

Lower California, 16 miles south of international boundary, six forma ochraceus
and ochraceus-nodiferus. 1. F. Ricketts.

Remarks.—It is interesing to note that in the southern extension of their ranges
both ochraceus and giganteus run to heavier spines, larger and often fewer crossed
pedicellariae, and much fewer furcate pedicellariae.

Mr. Paul O. Greeley observed that on the ocean side of the Government break-
water at San Pedro, Calif., segnis predominated to the extent of about 80 per cent
while on the harbor side P. giganteus capitatus was the more abundant, there being
only 1 or 2 segnis to 50 capitatus (November, 1925).

PISASTER GIGANTEUS (Stimpson)

Plate 73, Figures 5, 5a, 5b; Plate 74, Figures 3, 3a-3c; Plate 77, Figures 1, 2, 2a, 2b, 3, 3a—3c, 5, 6,
6a-6d, 7, 7a; Plate 78, Figures 5, 5a, 5b; Plate 83, Figure 2; Plate 85, Figure 2; Plate 86, Figure
11; Plate 87

Asterias gigantea Stimpson, Boston Journ. Nat. Hist., vol. 6, 1857, p. 528, pl. 23, figs. 4, 5, 6
(Tomales Bay, Calif.) —Verritu, Trans. Conn. Acad., vol. 1, 1867, p. 327.

Asterias liitkenii Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 265 (California or
Oregon).—VeERRILL, 1867, p. 32.—PrrnrierR, Rév. Stell., 1875, p. 70.—BrEt1, Proc. Zool.
Soc., London, 1881, p. 495.—Stapen, Challenger Asteroidea,- 1889, pp. 566, 824.—Dr
Lortot, Mém. soc. phys. et hist. nat. Genéve, vol. 32, 1897, p. 15, pl. 2 [17], figs. 1-1h
(Vancouver Island).

Asterias exquisita pr Lortot, Recueil Zool. Suisse, vol. 4, 1887, p. 408, pl. 18, fig. 2 (young
specimen, Santa Cruz, Calif.).

Calliasterias exquisita Fewkes, Zoological Excursions. 1. New Invertebrata from the Coast
of California, Bull. Essex Inst., vol. 21, 1889, p. 33.

P{isaster] liitkenii Verrityt, Amer. Journ. Sci., vol. 28, 1909, p. 63; Amer. Nat., vol. 43,
1909, p. 543.

Pisaster liitkenii Fisher, Smithsonian Misc. Coll., vol. 52, 1908, p. 89.—V»ERRILL, Shallow-
water Starfishes, 1914, p. 83, pl. 40, figs. 1, 2.

Pisaster liitkenti var. australis VeRRILL, Shallow-water Starfishes, 1914, p. 88 (off Pacific
Grove, Calif., not San Diego).

P{isaster) giganteus Verritt, Amer. Journ. Sci., vol. 28, 1909, p. 63.

Pisaster giganteus VERRILL, Amer. Nat., vol. 43, 1909, p. 545; Shallow-water Starfishes, 1914,
p. 89, pl. 37, figs. 1, 2—Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 561,
pl. 19, figs. 2, 2a, 2b; pl. 21, figs. 1, 2.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 173

Diagnosis.—Rays five or six; size sometimes very large with R over 300 mm.
A typical Pisaster, differing from ochraceus in having less numerous and longer,
solitary, terminally swollen, subconic to subglobose coarsely striated spines, rather
uniformly spaced over the abactinal surface, never in close ranked lines forming a
recticulate pattern, nor in acervate heaps; base of spine surrounded by a zone of
bare skin, blue in life, fringed by numerous crossed and fureate pedicellariae; stone-
hammer-shaped, large pedicellariae abundant to scarce. The giant type measures R
about 304 mm.,r 73 mm., R=4.1 r. A characteristic intertidal example from Mon-
terey measures R 123 mm., r 30 mm., R= 4.1 r; breadth of ray at base, 31 to 33 mm.
Disk rather high; ray, subcylindrical in section, slightly flattened actinally, and capable
of slight inflation at base.

Description.—The abactinal spines are numerous, solitary, spaced, with swollen
coarsely striated or furrowed, subconical or acorn-shaped ends. (PI. 77, figs. 2, 3.)
Each stands on a prominence of its plate, in the middle of a subcircular disk of bare
integument (blue in life) which is fringed with a usually thick cordon of crossed and
furcate pedicellariae. This blue sheath can rise nearly to the end of the spines,
which are white, or can be retracted to their bases. The spines do not typically form
either transverse or longiseries but stand irregularly. In some examples the carinal
series can be traced with ease. One can count across the ray at base 10 to 15 spines
in a zigzag way (or upward of 25 in big specimens). Small examples are found,
however, with only about five irregular longiseries and these are easily mistaken for
capitatus.

There are really two extremes of formae with even more numerous intergrades.
One has heavier, more widely spaced, less numerous abactinal spines. (PI. 87, fig. 2.)
The spines are usually more obtuse or subglobose terminally and it is this type
which sometimes predominates in capitatus, at the south. The other, or typical
form (pl. 87, fig. 1; pl. 85, fig. 2) has numerous slenderer spines with more elongate,
swollen tips, acorn-shape, subglobose, obtuse, conical or even subcylindrical. Their
coarse longitudinal furrows give them the appearance of tiny drills, and they vary
widely in form even in the same ray.

The spaces between the collars of pedicellariae are filled with the clusters of deli-
cate papulae which, when expanded, arise from swellings of thin integument above
the skeletal intervals. On the actinal surface the papulae are long and fill the spaces
between the spines. The bases of the papulae are armed with very tiny lanceolate
straight pedicellariae. (Pl. 77, fig. 5.)

Superomarginal series situated low on side of ray, often irregular proximally.
The spines, one to a plate, are very similar to the superadjacent abactinals, Inter-
marginal channel well-marked.

Inferomarginal spines two to a plate (occasionally three in big specimens),
clavate, longitudinally striate, the tip sometimes obliquely beveled, or near the base
of ray, irregularly eroded, suleate, or pitted. Most of the spines, as well as the similar
actinals, may have a sulcus on the external side, but this is sometimes lacking.
(Pl. 77, fig. 1.)

Actinal spines normally one to a plate, similar to the inferomarginals, but those
of the adjacent row often a little thicker and longer; those of the innermost row gener-

174 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

ally slenderer. The spines are longitudinally channeled or striated, the furrows
varying in depth. They are less evident in the type than in some large examples
from Monterey. There are ordinarily three or four longiseries of actinal spines in
the common medium-sized examples found in tide pools. Two specimens of the
heavier-spined form with R 55 and 80 mm. have two series. A large specimen of
the typical form from Monterey (R 215 mm.) has three, while one with R only 120 mm.
has four. The type (Tomales Bay) has six actinal spines in a transverse series on the
broadest part of ray and four at the middle. These actinals are clavate with usually
a sulcus on the external face. Verrill (1914, p. 87) mentions a specimen of “‘liitkenit”’
from off Monterey having R 284 mm., in which

The proximal inferomarginal and most of the actinal plates bear two, three, or even more, stout,
often divergent spines, so that one can count eight to ten of these spines on each irregular trans-
verse series. But one ray (doubtless a reproduced one) is only about two-thirds as long as the rest,
though nearly as stout; on this the ventral spines form only four or five rows, though similar in
size. On the distal half of all the rays the ventral rows decrease to about four to six, the plates
bearing only two spines, or but one.

Adambulacral spines, similar to those of ochraceus, in a single close series. In
the typical form the spines are slender, very slightly flattened and tapered, or mixed
tapered and untapered, round-tipped (blunt) with or without a depression at the
tip suggesting an incipient groove. In the larger-spined form the spines are more
evidently flattened with a slight groove on the outer side toward the tip. On the
narrow adoral carina the spines usually lengthen, become slenderer, and near the
mouth plates stand single file. On the ray the adambulacral spines stand very close
to the inner series of actinals, to which they are about equal in length.

The mouth plates have the usual Pisaster armature of a longer, usually tapering,
subacute, blunt, or subtruncate actinostomial spine, with a very short lateral com-
panion; and a prominent, tapered suboral, somewhat stouter than the innermost
adambulacrals. The Monterey specimens present numerous variations in the rela-
tive size and shape of the spines.

Madreporic body large, exposed, situated less than one-half r from center of
disk; striae fine, irregular.

Crossed pedicellariae (pl. 73, figs. 5, 5a) are smaller than those of capitatus and
typically more numerous. The majority occur in the circumspinal wreaths of the
abactinal and inferomarginal spines and in the external clusters of the inferomarginals
and actinals, but occasionally small clusters of them are found between the dorsal
spines, though not so commonly as in ochraceus. The pedicellariae of both large
and medium-sized specimens from Monterey Bay measure 0.27 to 0.31 mm. in length,
while those of the type (which is very large) vary from 0.27 to 0.34 mm., with a few
as large as 0.4 mm.

Straight pedicellariae: (@) Very small broadly lanceolate pedicellariae (pl. 73,
fig. 56) about 0.15 mm. long are abundant on the thin integument of the papular
swellings and on the bases of the papulae themselves. Similar ones form conspicuous
pedunculate clusters or festoons here and there along the furrow face of the adam-
bulacrals, usually at the base of a conspicuous, often quite large, toothed pedicellaria.
The small ones are 0.15 to 0.2 mm. long and the large one about 1.5 mm., sometimes
smaller or larger.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 175

(6) The bilaminate furcate pedicellariae (pl. 74, figs. 3, 3a-3c) are abundant
abactinally, brigaded with the crossed. They are not so plentiful on the actinal
spines. An average one measures 0.21 mm. broad by 0.2 high. They are smaller
than those of ochraceus and differ in form.

(c) The stone-hammer-shaped denticulate pedicellariae are very variable in size
and shape, and are similar to those of capitatus. (PI, 77, figs. 6, 6a-6d). They are
sometimes abundant abactinally, but again may be practically absent, except from
the proximal part of the intermarginal channel and among the actinal spines. The
largest (1.6 mm. long) are found in the actinal interradial channel. The jaw varies
from broadly lanceolate to broadly oval and the number of teeth from four to eight
This type of pedicellaria appears to be too variable to be of use in distinguishing
species although there are average differences,

Color in life: Abactinal spines white, surrounded by a conspicuous zone of
bright to dull blue integument; general tint brown, the papular areas raw Sienna;
actinal surface including inferomarginal spines, yellowish white with pale grayish
green papulae; tube feet Capucine buff. Young specimen: General tint cinnamon,
with spots of reddish brown between the collars of pedicellariae; spines pale pink,
violet, and blue; zone of blue skin surrounding spine pale blue. Another variety,
tawny olive mottled with snuff brown or sepia.

Variations —The type specimen has six rays and was dried without first being
hardened so that the abactinal surface has collapsed. It is a veritable giant, the
radii being 304 and 73 mm. The abactinal spines are very numerous, fairly equi-
distantly spaced, and the swollen end is elongate, tapered and slightly swollen at the
rounded tip. (Pl. 77, figs, 2, 2a, 26.) The large denticulate ‘‘stone-hammer”
pedicellariae are very numerous abactinally and are exactly like those of Monterey
specimens. As mentioned above, there are six actinal plus two inferomarginal spines
in a transverse series on the broadest part of the ray; on the middle 4+2; one series
of actinals reaches to within two or three inferomarginal plates of the terminal shield.
The inferomarginals carry two spines throughout the series.

At Monterey giant specimens of both the slenderer and heavier spined formae
occur, usually offshore or on the piles of wharves. They have never been taken in
tide pools. Occasionally a 6-rayed specimen is met with. In the giant specimens
of the heavier-spined form, the abactinal spines are rather more widely spaced, are
sometimes quite unequal, and the end is more subglobose than conical. Scarcely
two examples of these ‘‘overfed” individuals are alike. Verrill has noted a specimen
with an extraordinary development of actinal spines (1914, p. 87).

The medium-sized specimens occasionally found in tide pools at Monterey Bay
are also very variable in respect to the size, shape, and spacing of the spines and in
the number of ‘‘stone-hammer”’ pedicellariae present, as indicated in the description.
It is probable that the form with fewer, heavier spines has become dominant in the
southern part of California; or conversely, what is known as the race capitatus has
developed from this sparsely spined forma of giganteus.

Two specimens rather below medium size, from 48 fathoms, off Monterey (sta-
tion 3131), vary nearly as widely in respect to robustness of spines as do ordinary
intertidal examples. Both have very numerous “‘stone-hammer” pedicellariae

176 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Verrill (1914, p. 88) introduces a new variety from San Diego—Pisaster litkenii
var. australis. The description does not state the differences which separate the
variety from typical litkenii (that is, giganteus) but simply lists the characters of a
large specimen supposed to have been collected by Dr. Edward Palmer. This speci-
men, which I have examined, bears the number 4029 and also a very small label
pasted on, with ‘““W. R. Coe, 1901.” Doctor Coe tells me that he believes this
specimen to be one of several large ones which he collected at Pacific Grove, Mon-
terey Bay, in 1901. The loose label with “San Diego, Dr. Edward Palmer” by an
unfortunate circumstance became associated with this Monterey specimen. The
type of var. australis is therefore from Monterey Bay and represents simply an
individual variant.

Young.—Small specimens resemble capitatus in having few, relatively large,
abactinal spines. A specimen with R 27 mm. has a zigzag longiseries of dorsolateral
spines and a very irregular series of carinals. The stone-hammer pedicellariae are
numerous and thicker than the spine-tips in most cases. (Pl. 86, fig. 11.) The
superomarginal plates have a conspicuous area of hyaline bosses. Crossed and
fureate pedicellariae, numerous. Monterey.

Anatomical notes —The skeleton is essentially like that of ochraceus, a very irreg-
ular reticulum of higher spiniferous ridges inclosing spaces subdivided by lower, spine-
less trabeculae into the irregularly circular or oval papular areas (skeletal intervals).
The vertical intermarginal pillars between the supero and infero marginal plates
appear to be like those of ochraceus. These increase in size with age. At first, in
young specimens, there is simply a single ossicle tying the supero and infero marginal
plates. Secondary marginal and actinal plates develop with growth. As the animal
grows older the general skeleton is strengthened by the addition of elongate ossicles on
the coelomic surface, especially in the form of buttresses on the sides of the ray, and
across the actinal area, between the supero marginals and lower end of the ambulacral
plates. These actinal buttresses, composed of numerous small ossicles, separate a
series of rather deep pits all along the ray, at the outer end of the ambulacral plates.
The older the specimen the deeper the pits. Through these extensions of the coelom
the fluid of the body cavity reaches the extensive system of actinal papulae.

The ambulacral ossicles are very compressed and thin. The furrow broadens at
the base to narrow quickly toward the small actinostome. At the widest part there
are eight longiseries of tube feet in well-developed specimens.

The actinostome is only about one-fourth r in diameter and is characteristically
sunken so that it lies close to the abactinal body wall. The animal everts its stomach
when eating. It is the large very numerous and powerful proximal tube feet which
open mussel shells, or lift barnacles and limpets from rocks. The long adoral carina
is much narrower at the inner end than the oral plates. Occasionally plates of one
side of carina are overdeveloped so that the median suture lies on the side of the
carina, or follows a rather sinuous course.

Type.—No. 1283, U.S.N.M.

Type locality —Tomales Bay, Calif.; Mr. Samuels.

Distribution —Southern part of Vancouver Island to Monterey Bay, intergrad-
ing to the south with P. giganteus capitatus. Low tide to 48 fathoms (Monterey Bay).

Specimens examined.—Very numerous examples from Monterey Bay, Calif.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 177

Station 3131, Monterey Bay, Calif., 36° 41’ 30’ N., 124° 54’ 10°’ W., 48 fathoms,
mud, rocks, two specimens.

The type of Asterias litkenii, “California or Oregon.”

The type of Asterias gigantea.

Remarks.—This species is very distinct from both ochraceus and brevispinus
and can be recognized by its usually brown or purplish-brown color and evenly distri-
buted, whitish, dorsal spines surrounded by a zone of blue skin, often very vivid in
luster.

It is regrettable that the name giganteus falls to this beautiful species by the
fortunes of nomenclature, since the common tide-pool form is by no means gigantic
for the genus. In favorable localities, however, the species does reach a large size, as
evidenced by the type with R 304 mm., and by specimens caught offshore in
Monterey Bay.

I have not examined specimens from north of California unless the type of
litkenii is from Oregon. Verrill records a specimen from Vancouver Island (Canadian
Geological Survey) and de Loriol one from Saanich Inlet, north of Victoria on the
east side of Vancouver Island. De Loriol’s figure * shows a specimen with very
small, tapered, usually not capitate spines, with occasionally two or three on the
dorsolaterial plates and usually two or three on the carinals. This specimen may
represent a forma comparable to confertus in ochraceus. It certainly is not typical

giganteus.
PISASTER GIGANTEUS CAPITATUS (Stimpson)

Plate 73, Figures 6, 6a; Plate 77, Figures 4, 4a; Plate 85, Figure 3; Plate 8S

Asterias capilata Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1862, p. 264.—Vexnrint,
Trans. Conn. Acad. Sci., vol. 1, 1867, p. 327.—Perrnrier, Rév. Stell., 1875, p. 71.—
Stapen, Challenger Asteroidea, 1889, p. 566, 820.

P {isaster) capitatus Verrity, Amer. Journ. Sci., vol. 28, 1909, p. 63.

Pisaster capitatus C. F. Baker, First Annual Report of Laguna Marine Laboratory (1912),
p. 89.—Verritt, Shallow-water Starfishes, 1914, p. 81, pl. 36, figs. 3, 4; pl. 56, fig. 4.

Pisaster giganteus capitatus Fisner, Ann. Mag. Nat. Hist., ser. 9, vol. 17, 1926, p. 562.

Diagnosis —Differing from Pisaster giganteus in having typically fewer and
generally stouter abactinal spines, larger crossed pedicellariae, and very few furcate
bilaminate straight pedicellariae. Rays five, rarely six. R 190 mm., r 52 mm.; rays
inflated, abactinal spines stout, widely spaced, subconical; larger cireumspinal collars
8 to 10 mm. broad. Narrower-rayed form (Venice, Calif.), R 170 mm., r 40 mm.,
width of circumspinal collars, 5 mm.

Description —The two formae of P. giganteus extend into southern California
and are thus included in the race capitatus. The somewhat heavier spined form (to
which the type belongs) has even fewer spines than its northern equivalent but there
are scarcely two specimens alike. One from San Diego with R 140 mm. has only
two dorsolateral series of spines irregularly aligned (about five longiseries between the
superomarginals, while a specimen from Laguna Beach (R 180 mm.) has the equiva-
lent of nine series without serial alignment. The spines are widely spaced with
broad collars of bare skin fringed by a circular ruff of crossed pedicellariae. The

© Mém, soc. phys. ot hist. nat. Gemdve, vol, 92, i807, pl. 2, fe. t.

178 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

space between the spines may reach 10 or 12mm. The spines are strongly capitate,
robust, with ornately grooved, subglobular to acorn-shaped ends, and are seated
rigidly on convex plates of the very irregular skeleton. The dorsolaterals are some-
times heavier than the carinals and superomarginals, sometimes subequal to them.
(Pl. 77, fig. 4.) In between the spines are conspicuous clusters of papulae guarded by
small lanceolate pedicellariae, attached to the papulae, and in most cases by few to
many much larger stone-hammer denticulate straight pedicellariae similar to those of
giganteus. There are very few isolated clusters of crossed and furcate pedicellariae
among the papulae.

The other extreme has narrower rays and more numerous, generally slenderer
sometimes quite unequal abactinal spines. There is little to distinguish it from
some specimens of true giganteus (with which Verrill confused it) except the characters
of the pedicellariae enumerated in the diagnosis. One may count from 9 to 15
spines across the ray between the two superomarginal series, but there is never
regular serial alignment. The collar of blue skin around each spine is generally
narrower than in the type forma. The usually slenderer rays cause the spines to be
less widely spaced. There is complete intergradation between the two formae in a
good series of large specimens from Venice, Calif., contributed by Prof. A. B. Ulrey.
The Laguna Beach specimens are all the type form.

The superomarginal spines are similar to the dorsals, but generally more numerous
than the outer dorsolaterals, and form a straight series, low on the side of ray. A
specimen from San Diego has 23 superomarginal spines corresponding to 47 infero-
marginal plates.

Inferomarginal spines smaller than the superomarginals, two to a plate, sub-
clavate, finely grooved and with a well marked, external, often deep sulcus. There
are usually two or three longiseries of actinal spines similar and subequal to the in-
feromarginals, sometimes a little heavier on the proximal part of the ray. They all
carry external clusters of crossed pedicellariae. As in typical giganteus certain speci-
mens, on some of the inferomarginal and actinal plates of the proximal half of ray have
two to four spines, greatly increasing the spine count and destroying any regular
alignment. Such a specimen may have 8 or 10 spines in an irregular transverse series.

Adambulacral spines slender, flattened, subtruncate or blunt, slightly tapered or
not, sometimes with a shallow groove on the outer half, deepening toward the tip
(Laguna Beach). In such a specimen the spines range from slightly tapered, blunt,
to slightly spatulate. The adoral carina is very narrow, composed of about 10 or 12
pairs of plates, and back of the mouth plates the first two or three pairs of plates
have the spines in single file (some of the spines being suppressed).

Each mouth plate has two actinostomial spines (the latter quite small) and one
suboral, tapered and pointed, similar to but often stouter than the adjacent adam-
bulacral spines.

Crossed pedicellariae in circumspinal wreaths abactinally and in clusters on
outer side of inferomarginal and actinal spines. They are less numerous than in
typical giganteus which sometimes has clusters between the spines. Pedicellariae
average larger than in giganteus; length 0.5 to 0.46 mm. (Laguna Beach; San Diego;
Lower California). (Compare pl. 73, figs. 5 and 6.) In form they adhere closely to
the generic type and are very little different from those of giganteus. Note relative

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS— FISHER 179

length of side of terminal lip. In Lower Californian specimens there are fewer teeth
in the median vertical series of the shank of jaw.

Straight pedicellariae: (a) There are a few to numerous tiny lanceloaté dermal
pedicellariae, similar to those of giganteus, guarding the papulae (actinal and abacti-
nal). Along the furrow face of the adambulacrals are pedunculate clusters, here and
there, of very small lanceolate pedicellariae, usually at the base of a large toothed one
(as in giganteus).

(6) Small furcate pedicellariae, so numerous in giganteus, are relatively few and
occur with the crossed; diameter 0.225 mm. (PI. 73, fig. 6a.)

(c) Heavy subconical “stone-hammer” pedicellariae with small terminal teeth
vary greatly in number and present no constant differences from those of giganteus.
When almost absent from the abactinal surface they may be found intermarginally
and in the interbrachial channel. The jaws are likely to be broader in the thicker
spined specimens.

Madreporic body subcircular, conspicuous, less than one-half r from center of
disk; striae fine, irregular.

Young.—Two young specimens from Laguna Beach, Orange County, Calif.,
measure R 16 and 24mm. Each has a single series of well spaced dorsolateral spines
and a single series of actinals. In the smaller there are two pairs of contiguous adam-
bulacrals behind the mouth plates; in the larger, four. (PI. 85, fig. 3.)

In contrast to giganteus of approximately the same size, the abactinal spines are
fewer and more widely spaced while the furcate pedicellariae are very scarce; in
giganteus these are abundant.

The skeleton of the smaller specimen is constructed essentially as that of young
ochraceus, but shows larger areas of hyaline beads on the superomarginal plates,
which are practically covered. They are also on the primary dorsolaterals and
carinals (absent in ochraceus).

Type.—No. 1280, U.S.N.M.; Dr. J. S. Newberry.

Type locality—San Luis Obispo Bay, Calif.

Distribution —N orthern Lower California to San Luis Obispo Bay, Calif.; inter-
tidal and shallow water.

Specimens examined.—Twenty-eight.

Specimens of Pisaster giganteus capilatus examined

Locality of speci- Remarks
mens |

ee a SP Uy Cask can ctpodsdshvenoncucenscuneucadncon Type. Dr. J, 8. Newberry.
Santa Barbars, Calif_................ oe 2) D.8. Jordan, 1880.
Venice, Los Angeles County, Calif.................. owodube eve 12 A. B. Ulrey, Stanford Coll.
Laguna Beach, Orange County, Calif wmiciisceetsiglihe mt 5 W. A. Hilton, C. F. Baker, Stanford Coll
Lower California (16 miles south of international boundary)........ 4 E. F. Ricketts.
ee SE ee ceria leaned cc anicasanccsieccumetinsoancnen ed 3) BE. C. Starks, Stanford Coll

Kn Fadia Caio a cs wdcts -tciicesdetacens we ccencsccsceemsccoscessaucaces 1) 30888, U.S.N.M.
!

180 BULLEYIN 76, UNITED STATES NATIONAL MUSEUM

Remarks.—By a curious mistake the type locality is stated by Stimpson to be
“San Diego, Calif., Colorado expedition, Dr. J. S. Newberry.’’ The type specimen
is No. 1280, U.S.N.M., and is labeled in Doctor Stimpson’s well-known handwriting
“ Asterias capitata, Stm., San Luis Obispo, Doctor Newberry.” This specimen is
really intermediate between capitatus and giganteus, as the locality would lead one to
infer. It has not the characters of the race capitatus developed to the extreme
degree that one would wish to see in a ‘‘type”’ and it is very unfortunate that the type
specimen is not from San Diego, or from some locality well south of Point Conception.

Dr. Mary J. Rathbun, in this connection, showed me documentary evidence that
Doctor Newberry did not collect at San Diego. In 1883, Dr. Richard Rathbun
published the following in Descriptive Catalogue of the Collection Illustrating the
Scientific Investigation of the Sea and Fresh Waters (Great International Fisheries
Exhibition, London, 1883, U. S. America), page 12:

Explorations and surveys west of the one hundredth meridian, under the direction of Gen.
A. A. Humphreys, Chief of Engineers, by Lieut. George M. Wheeler in charge, from 1872-1879.
The naturalists of this survey were Dr. H. C. Yarrow, Mr. H. W. Henshaw, Professor Newberry,
Mr. Charles E. Aiken, Dr. J. T. Rothrock, and Oscar Loew, and their field operations included the
fresh-water lakes and rivers of Utah, Colorado, New Mexico, Arizona, and western and south-
western Nevada, Salt Lake, Utah Lake, and other salt-water lakes, and the Pacific coast in the
vicinity of Santa Barbara, Calif.

The type has R 81 mm., r 21 to 23 mm. The abactinal spines are of the large,
capitate, well-spaced sort characteristic of capitatus from further south, but the
crossed pedicellariae are too small and the furcate pedicellariae are too numerous.
The larger crassed pedicellariae measure 0.36 mm. in length while the Laguna Beach
and San Diego examples measure 0.4 and 0.45 mm. (giganteus, 0.26 to 0.31 mm.).
The smaller pedicellariae of the type measure 0.31 mm. in length. The furcate
pedicellariae are not so numerous as in typical giganteus, but are not so scarce as in
the extreme form of capitatus.

Although the type of capitatus is frankly an intermediate it is superficially more
like the southern than the northern race.

The record of capitatus from Monterey Bay appears to be based on the form of
giganteus having unusually stout, well spaced spines. I have a young specimen which
might pass for capitatus but it has abundant furcate and small crossed pedicellariae.

PISASTER BREVISPINUS (Stimpson)

Plate 74, Figures 4, 4a—4e; Plate 76, Figure 8; Plate 78, Figures 1-4; Plate 79, Figures 2-5; Plate
86, Figures 12-16; Plates 89-93

Asterias brevispina Stimpson, Boston Journ. Nat. Hist., vol. 6, 1857, p. 528, pl. 23, fig. 3
(San Francisco Bay, Calif).

Asterias paucispina Srimpson, Proc. Boston Soe. Nat. Hist., vol. 8, 1862, p. 266 (Puget
Sound).—Prrrier, Réy. Stell., 1875, p. 60.

Asterias (Pisaster) papulosa Verritt, Amer. Journ. Sci., vol. 28, July, 1909, p. 63 (Puget
Sound).

P{isaster] brevispina Verriwyt, Amer. Journ. Sci., vol. 28, 1909, p. 63.

Pisaster brevispinus VERRILL, Shallow-water Starfishes, 1914, p. 77, pl. 41, figs. 1, 2; pl. 44,
figs. 1, 2; pl. 45, fig. 1; pl. 69, fig. 3; pl. 76, figs. 1-10.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS FISHER 181

Pisaster papulosus Vennitt, Shallow-water Starfishes, 1914, p. 91, pl. 42, fig. 1; pl. 43, fig. 1;
pl. 60, fig. 1; pl. 76, figs. 22d; pl. 80, fig. 4.
Pisaster ? paucispinus Vernity, Shallow-water Starfishes, 1914, p. 98, pl. 36, figs. 1, 2.

Diagnosis.—Difiering from P. ochraceus in its rose-pink coloration, variegated
with gray-green or maroon-purple papular areas; dorsolateral spines not forming
a reticulated pattern (except occasionally in forma brevispinus) but standing singly
or in small groups, often few in number; furcate pedicellariae of characteristic form,
with upper margin of jaw smooth, not crenulate. Rays five. A giant specimen of
forma brevispinus (pl. 90, fig. 2), R 320 mm., r 66 mm.; one of forma paucispinus
(Verrill’s papulosus) R 315 mm., r 60 mm. This species is one of the largest star-
fishes known.

Description. —The most obvious variation in this species is in the abundance of
abactinal spines. There is a complete series of intergrades between the extremes.
It is convenient to recognize two formae equivalent to the old species brevispinus
and paucispinus. Each of these is divisible into several subformae; but in view of
the fact that there are about as many intergrades as representatives of the subformae
no service is rendered by recognizing them formally by name.

The species brevispinus is easily distinguished in life from its congeners by its
light rose-pink color, mottled with the gray-green or maroon papular areas. This
color is rather striking and no variation of ochraceus or giganteus approaches it,
Moreover, the pink color varying in intensity and shade is relatively constant, as
compared to the wide range in ochraceus from yellow ochre to purple.

Forma BREVISPINUS (Stimpson)

Plate 74, Figures 4c, 4d; Plate 76, Figure 8; Plate 78, Figures 1, 2, 2a, 2b, 4, 4a, 4b; Plate 79, Figures
2, 2a, 2b, 3c, 4; Plate 89; Plate 90, Figures 1, 2; Plate 91, Figure 1; Plate 92; Plate 93, Figure 4

To this forma belong those individuals having dorsolateral spines typically in
small groups or clusters, which are spaced without regular order between the carinals
and the superomarginals. There is great variation in the number of spines per
group as well as in the number of groups. In extremes such as Plate 89, Figures 1
and 2, there are 3 or 4 to 8 or even 10 capitate spines per group, sometimes arranged
in short transverse series. The groups are somewhat acervate (pl. 89, fig. 2; pl. 90,
fig. 2), especially on the outer part of the ray. In these extremes the carinal spines
are crowded in transverse groups on the plates so as to form a conspicuous radial
band. The primary apical plates of the disk are similarly heavily armed, the spines
forming a rude star. These accentuated forms of brevispinus have been taken at
Puget Sound, Wash.; Crescent City, Bolinas, and Monterey Bay, Calif.

The less pronounced phase of brevispinus is shown on Plate 92. Figure 1 is com-
parable to Stimpson’s type (see Verrill, pl. 41) from San Francisco Bay. In these,
the dorsolateral spines stand singly or in small groups of two to five. They are
slenderer and the end is frequently subconical with a subacute or rounded tip. The
carinal spines are one to three to a plate. In some specimens there is but a single
series of carinal spines while in others, not materially different as to dorsolateral
armature, the spines are two or three ranked on the proximal carinals or on nearly all.

182 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

In some specimens of the less spinous subforma the abactinal spines are slender,
tapered, subacute, without, or with only a slightly swollen, subcapitate tip. They
stand usually singly on their plates and their collars of pedicellariae are generally
conspicuous. Plate 92, Figure 2, represents a specimen intermediate between
Figure 1 and this weak-spined variant (Monterey Bay). The giant specimen (pl. 91)
has tapered pointed spines and belongs on the border of brevispinus, as does Plate 93,
Figure 4. The latter has fewer and slightly stouter spines than specimen Plate 92,
Figure 1, and is an intermediate between the well-developed phase of brevispinus and
paucispinus. The loss of the adradial series of spines would transform this specimen
into one of forma paucispinus. This remark is true of the great specimen, Plate 91.
Compare it with Plate 90, Figure 2, a representative of the heavier spined subforma.
Both are from San Juan Islands, Wash.

Superomarginal spines, one to three to a plate, are not of much value for diag-
nostic purposes. The number varies on different rays of the same specimen. Speci-
men, Plate 89, Figure 1, has two (sometimes one) on plates of proximal two-thirds
of ray and one on the distal third; Figure 2, the same; Plate 90, Figure 2, giant
specimen, has three sometimes with a small fourth, at base of ray, then two or three,
and finally one near tip; Plate 91, figure 1, giant specimen, two proximally and one
distally; Plate 92, Figure 1, irregularly two or one on a few proximal plates and one
on rest of series. Superomarginals are similar in size and form to the abactinals
but in some specimens are slightly gouge-shaped on outer side of tip.

The inferomarginal and actinal spines are characteristic of the genus and present
no especial peculiarities. There are two clavate to subspatulate, usually externally
furrowed, inferomarginals, followed in a transverse series by three or four similar
actinals, all much stouter than the superomarginals. In well-developed specimens
the two outer actinals are generally a little larger than the inferomarginals. There
is considerable variation in the form of the spines and their robustness. In very large
specimens proximally the tips become flattened as if pinched while soft. A shallow
sulcus is normally present on the outer side in most of the specimens. (PI. 78, fig. 1.)

Adambulacral spines slender, slightly tapered, round-tipped, forming a single
crowded series. Here and there a spine is squeezed a little in advance of its neighbors
on the furrow edge. Where the series bends upward toward the actinostome the
spines gradually lengthen and thicken, on the approach to the adoral carina, upon
which they are generally conspicuously stouter and longer. This carina is unusually
well developed and in mature specimens is composed of 12 to 15 pairs of conjoined
plates. (Pl. 79, fig. 2a.) It curves upward to the actinostome, which is sometimes
quite close to the dorsal wall of the well-arched disk. The first few plates of the
carina lack spines, although these same plates are spiniferous in young specimens.
In fully adult specimens the first adambulacral plate is conspicuously shorter than
the second; in young specimens they are subequal to or larger than the following
plates.

The mouth plates widen with age and sometimes become somewhat distorted.
They are, as is usual in Pisaster, considerably broader than the narrow neck of the
adoral carina but there is little specifically characteristic. Ordinarily there are two
unequal flattened blunt marginal (actinostomial) spines and a longer and stouter,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS~— FISHER 183

variously formed, but usually tapered and blunt suboral spine, not in the same
position on companion plates, owing to crowding.

Papulae abundant, grayish green in life, arising from various sized hernialike
elevations of thin integument carrying numerous very small broadly lanceolate
pedicellariae, which also occur on the base of the papulae themselves. ‘These clusters
are of unequal size, some containing a large number of papulae and they are closely
spaced among the spines. Actinal areas each with few, very long papulae, very
numerous and conspicuous in the aggregate.

Crossed pedicellariae (pl. 79, figs. 2b, 4) of the usual Pisaster type are fairly
abundant, mixed with the fureate type, in circumspinal wreaths and in numerous
small clusters, independent of spines, among the papulae. They occur also in
clusters on the outer side of the inferomarginal and actinal spines. Abactinal pedi-
cellariae from a giant specimen (pl. 90, fig. 2) measure 0.24 to 0.27 mm. in length
(San Juan Islands, Wash.), while on a large Monterey specimen (pl. 89, fig. 2) they
measure 0.31 mm. Actinal and inferomarginal pedicellariae are larger (0.35 mm.).

Straight pedicellariae: (a) The Pisaster bifid furcate pedicellariae are character-
istic in form (pl. 74, figs. 4c, 4d) and differ from those of ochraceus and giganteus in
having the shorter blade of the jaw in a rudimentary form. The larger blade is
more like that of giganteus but is uniformly larger. Its upper border is smooth, not
scalloped as in ochraceus. The pedicellariae have a diameter of 0.27 to 0.32 mm. and
are remarkably uniform in size, irrespective of size of specimen and locality. They
have the usual distribution and in some specimens outnumber the crossed pedicel-
lariae abactinally, especially on the clusters between the spines.

(6) Very small, broadly lanceolate pedicellariae are usually abundant abactinally
as well as actinally, especially on and around the bases of papulae. Actinally they .
are frequently of more diverse and larger sizes. Very similar ones form conspicuous
pedunculate clusters on the furrow surface or margin of the adambulacrals. These
clusters, or sometimes long festoons, generally carry one or several very large, termi-
nally toothed pedicellariae, about which the smaller ones cluster in masses. (PI. 76,
fig. 8; pl. 78, figs. 2, 4.)

Large dentate pedicellariae of numerous sizes up to 2.5 mm. (exceptionally
4 mm.) long are sometimes very abundant. (San Juan Islands, pl. 90, fig. 2.) There
is such wide variation in form (the jaws varying from a tapered form with narrow
tip and few teeth (pl. 78, figs. 3, 4, 4a-44) to a broad rounded end and 6 or 8, excep-
tionally 10 or 12, teeth (pl. 78, fig. 2a) that it is difficult to find features of precise
diagnostic value. The shape varies widely on the same specimen, being generally
longer and slenderer actinally (fig. 3d), where the longest occur in the axillary channel.
Specimens occur in which there are few abactinal dentate pedicellariae. The relative
development of spines has no relation to the abundance of pedicellariae or vice versa.
The large specimen from Lopez Island, San Juan Islands (pl. 90, fig. 2), has a most
extraordinary equipment of this type of pedicellariae, the other sorts being very
numerous but less conspicuous (pl. 78, figs. 2a, 24). Among the examples of forma
paucispinus are two from Nanaimo having a minimum of spine development and the
abactinal surface armed with a multitude of these “major” pedicellariae which are
more conspicuous than the spines. (PI. 91, fig. 2; pl. 78, figs. 3, 3a-3d.)

184 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The madreporic body is large, subcircular, and situated about one-third r from
center of disk.

Color in life: Rose pink, the papular areas dark sage green or maroon purple.
Some specimens are brighter or more intense pink than others. Those with purplish
papular areas are more striking than those with the more prevalent greenish areas.

Forma PAUCISPINUS (Stimpson)

Plate 74, Figures 4, 4a, 4b, 4e; Plate 78, Figures 3, 3a-3d; Plate 79, Figures 3, 3a, 3b, 4a, 5; Plate 86
Figures 12-16; Plate 90, Figure 3; Plate 91, Figure 2; Plate 93, Figures 1-3, 5

The type of Stimpson’s Asterias paucispina represents the immature phase and
that of Verrill’s Pisaster papulosus a mature stage of this form.

The principal distinction lies in the number of dorsolateral spines. In typical
examples these stand singly, or in twos and occasionally threes, in one fairly direct to
decidedly zigzag longiseries. At the end of the ray a few extra spines are added on
either side. These spines or small groups are widely spaced from each other and from
the carinals and superomarginals. Certain variants add spines here and there outside
the series. Plate 90, Figure 3, represents the primitive form. Similar specimens,
having R upward of 120 mm. are not uncommon at Monterey Bay. Plate 93,
Figures 1 and 3, is the variation having the spines occasionally in groups of two or
three, and with a few scattered ones near the marginals. This intergrades with
specimens similar to Plate 93, Figure 4, which is an intergrade with forma brevispinus.

In forma paucispinus (pl. 79, figs. 3, 3a-3c) the spines are usually stout with
subconical, or acorn-shaped, grooved, differentiated tips. A subforma (pl. 91, fig. 2)
with very few, rather inconspicuous dorsolateral and carinal spines but with a mul-
titude of large dentate pedicellariae (often more robust than the spines) is found at
Nanaimo, British Columbia, and represents the extreme of difference from the mul-
tispinous subforma of brevispinus. This small spined variety is represented by a
giant specimen from station 4219, vicinity of Port Townsend, having R 315 mm. It
also has very numerous “‘major’’ pedicellariae which are not so robust as the spines.
A specimen (pl. 93, fig. 2) from soft green mud, Monterey Bay, also has very few,
rather weak spines, but only the usual equipment of pedicellariae. A variation with
spines similar to this, but with almost no larger straight pedicellariae occurs at
Bradley Lagoon, British Columbia. In forma brevispinus nearly as wide a variation
in the number of straight pedicellariae is found.

Papulae are normally very numerous and, on account of the fewer spines, are
rather more conspicuous than in brevispinus. They occur in clusters interspersed
with groups of crossed and furcate pedicellariae, with which, in alcoholic specimens,
it is easy to confuse them. As in other species of Pisaster the ventral papulae are
long, and increase in number with size of animal. In large specimens they form dense
masses between the ventral spines. Verrill’s Pisaster papulosus, characterized by
few spines and very numerous papulae, is the normal adult of paucispinus of the ex-
treme type.

Superomarginal spines one, or sporadically two, to a plate, sometimes smaller
sometimes larger than the dorsolaterals. In the curious subforma from Nanaimo
(pl. 91, fig. 2) several consecutive superomarginal (and carinal) plates may lack

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 185

spines. The available lime seems to have gone into pedicellariae. There are indi-
cations that in some cases the spines have been absorbed.

Inferomarginal spines two (sporadically three). Actinal spines in two, rarely
three, longiseries. The inner spine, when present, is generally slenderer than the
others. The big specimen from station 4219 has sometimes two, sometimes three
spines in the cross series of the actinal system, or four and five counting the infero-
marginals. Forma brevispinus has usually three or four, exceptionally five longiseries
of actinals at base of ray. There is no constant chatacteristic difference distinguish-
ing the spines in the two formae.

The adambulacral and oral armature is similar to that of forma brevispinus.

The pedicellariae of forma paucispinus are like those of brevispinus and vary in
number within about the same limits. Specimens from Nanaimo have an extraor-
dinary equipment of large dentate pedicellariae. The bouquets of adambulacral
straight pedicellariae are often very voluminous and can be extended in life to the
ends of the papulae which are adjacent to the adambulacral plates. They apparently
afford some protection to these papulae and to the tube feet, among which the long
peduncles of the bouquets twine. (PI. 74, figs. 4, 4a, 4b, 4e; pl. 78, fig. 3, 3a-3c;
pl. 79, fig. 5 (inferomarginal).)

Madreporic body large, exposed, situated as in brevispinus.

Color in life: Rose pink with sage green papular areas. There is apparently
no constant difference in color from f. brevispinus. A specimen from station 4503:
Abactinal spines light rose purple; general surface olive buff; papular areas dark
grayish green; actinal surface, light rose purple.

“oung.—Plate 86, Figures 12-16. These figures are intended to facilitate com-
parison between the young of the three species of Pisaster.

The young, by reason of the paucity of dorsolateral spines, are all referable to
forma paucispinus. Forma brevispinus passes through this stage and adds spines,
while the adults of paucispinus maintain the juvenile arrangement of one dorsolateral
series. Figure 12 has R 15 mm. and one actinal series of spines for half the length of
ray. The furcate and crossed pedicellariae are spaced, in circles, around the spines
and form groups of two or three in between. Figure 13 has R 30mm. _ The actinals
extend nearly to tip of ray. Both have relatively few dentate straight pedicellariae,

One means of distinguishing the young of the three species, especially brevispinus
from ochraceus, is the furcate pedicellariae, the jaws of which are characteristic.

Skeleton —There are no striking points of difference between the skeletons of
brevispinus and ochraceus. In brevispinus the prominent groups of dorsolateral spines
are seated upon plates much larger and more convex than the others. The same
development is seen in ochraceus forma nodiferus. In both species the skeleton is
strengthened, during the growth of the species, by the addition of internal ossicles.
This is especially observable in a series of transverse superactinal partitions or but-
tresses which pass from the ambulacral across to the superomarginal systems.
Between these partitions are deep recesses or wells into which the highly important
actinal papulae open.

The mouth plates appear to be more deeply sunken in this species than in the
other two, but as considerable movement is possible, variation is apparent. When the
actinostome is near the upper surface of the disk, as it frequently is, the adoral carina

186 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

is at right angles to the substratum upon which the animal is crawling. There are
fewer longiseries of actinal plates (which are a little larger) than in ochraceus.

In young specimens (pl. 79, fig. 4a) the skeleton has a series of wide intervals or
spaces adjacent to the superomarginals and the primary dorsolaterals are relatively
big. As in ochraceus the carinals lose their 4-lobed contour later in life.

Very young specimens (R 14 mm.) have, on the superomarginal plates, a very
small area of hyaline bosses. These disappear in specimens with R 18 mm. In
young of P. giganteus capitatus the superomarginals have relatively more surface, are
more triangular in shape, and these specialized areas cover nearly the whole surface.
They are well-marked in the young of ochraceus, having R 22 mm.

Type of Pisaster brevispinus—Cat. No. 1285 U.S.N.M.

Type locality—Near mouth of San Francisco Bay, Calif., 10 fathoms, sand.

Distribution Sitka, Alaska (Verrill) to Santa Barbara, Calif.; lowest tide to
56 fathoms, sand, or mud and sand.

Forma brevispinus has been taken in the Puget Sound region, Washington, and
from Crescent City to Monterey Bay, Calif.

Forma paucispinus (type locality, Puget Sound) ranges from Sitka to Santa
Barbara.

Specimens examined.—One hundred and fifteen, in addition to many from Mon-

terey Bay.
Specimens of Pisaster brevispinus examined

ae Locality Depth Nature of bottom vet eae Remarks
Fathoms
3110 | Vicinity Half Moon Bay, Calif-__.-.-._-- 89) || FRocky <2 52k =o aoe eco ane
BILIC | eenee COE be A EE ee ee TEs 901)\Grayisand: ee et ee 2 4 intermediate.
3122 | Between Point Afio Nuevo and Santa 88ie| | Gravisand, muds ssesseke = spe lesenae ce 2 |
Cruz, Calif. |
3147 |____. COREA: Secon See eee eee eens 86:9 Browmimud’=s see csoe ee eeeeneee 1)
3153 6255 (Green wmud! Se oo eee eRe 1
3154 20) Black sand, mudte =. .2-oe=. |eoeseeee 3 |
4219 | Vicinity Port Townsend, Wash___._---._- 16) )'Greenwmud,:sand==--ses-2==s|-==ee eee 1
4487 | 3.7 miles southeast Santa Cruz Point, 18% | Hardigrayisand -22--2225--2> yee
Calif. (Monterey Bay).
4492 | Monterey Bay, 7 miles southeast Santa 26" | Soft'green mud.-.-=2=4-..-=4|223. 22. 1
Cruz.
S005 Montereyi Bay ~-sseeo ceee eee tone ee 12 | Hard coarse sand_.-..------- 7 6 | 2 intermediate.
4503 | Monterey Bay (off Soquel)_-.---__________ iy | eaxayysandeete 2 te eet ce 4 25
4519 | Monterey Bay (off Moss Landing) _____--- 3b, ||| Hard pray'sand-# =~ 2) 8 = alee See 1 intermediate.
4560 | Monterey Bay (off Soquel Point) __ es 10 | Fine gray sand_-__-__- 1 3
M561 ||‘=3.3 Gort 22 Sess oe 15 | Coarse sand, shells____ dl eeeeeer 4
4562 | Monterey Bay (8 miles southeast Santa 10) baktard sand, rocks: 22-25-24 |-5- ou | 6
Cruz).
4563 | Monterey Bay (off Soquel Point)________- 8) Rocky #21 4H. 26 Seow io, ee 3
5718 | San Francisco Bay (middle) Ae 779) (Graymudie. Se heal eo 1
5762 | San Francisco Bay (lower) _----.--_-______ 6. ide coc ee ee eae 1
| Kasaan Bay, Prince of Wales Island, south- |. .-_____- bs 535 ee ae 12 See ee 2/| T. H. Streets.
east Alaska.
jppxadleyWeeoon, British Columbia. oe e-toc sane | oo eee ee en ee ee | esto! 1} W. F. Thompson.
Barclay Sound, Vancouver Island, British |.-.......|.-.....-..-.--..-----.-.--.---|_------- 2
Columbia.
Nanaimo, Vancouver Island, British Co- |Wharves|._._..-..__........__.-.-----_|_--__--- 3
lumbia.
| Departure Bay, British Columbia...._.__- 110:| Soft sand... 25-65, eeo ache eeeeseee 2
| Lopez Island, San Juan Island, Wash_---- 18-25 | ooo ooo ee ae ee 2) 2 Miss M, Bush.

1 Feet.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS~-FISHER 187

Specimens of Pisaster brevispinus eramined—Continued

= f. brert- 1. pauel-
Depth Nature of bottom spinus | opinus Remarks
| Fethome
loons 1 1 Mus. Comp. Zoit
a eee a Rings ‘ Do.
Piss} eda duanp sbpuphnaddcdhde suse 2 1 | J. 0, Snyder
BS eran citates diohs 1 | ©. M. Drake
(————E————E—————— 2 | Intermediate.
indians alos och quthe dies dgecaeddusd «bho A lena :
Cg. a a i 1 PF. W. Weymoath
PY CRs SCG TROIEO PO clan eiepericlenepepyaquiuvepncurspgaree= ; 1 \........) W. F. Hamilton.

San Francisco Bay....... vntbeleintatoe
PR Se ee De einen Intermediate.
Monterey Bay, Calif..... | . Sand... rt] Hopkins Marine Sta-
on; Mus. Comp.
Zoi. No, 1821.

ee RUE ER CHEB ers oerecinetinrehiass taied enopeeaed kieditiedunwaiipmmnans tek ninth asiomnebaianiaiets
Santa NiMMiaial gene ee ee delta een 2) Mus. Comp. Zoi.

Remarks.—Verrill’s Pisaster papulosus is the fully grown phase of the immature
specimens to which Stimpson gave the name Asterias paucispina. These are figured
by Verrill (1914, pl. 36). Figure 1 (Cat. No. 1287, U.S.N.M.) is the cotype, not
type; Figure 2 (No. 1286) is the type. Both are somewhat museum worn, but I
have numerous specimens, almost identical, as well as smaller and larger ones.
Apparently forma brevispinus always begins as paucispinus and adds spines to the
few well-spaced ones of the single dorsolateral series. I have seen only one speci-
men of brevispinus as small as R 53 mm.; most of them have R 100 mm. or more.

Verrill was in doubt concerning the generic position of paucispinus because
only one series of actinal spines is present in the type. This is only a juvenile
condition.

In connection with Verrill’s account of brevispinus (p. 81) the record of Santa
Cruz Island, Calif., is an error. The specimen (1821, M. C. Z.) came from Monterey
Bay, off Santa Cruz. The figures representing the crossed pedicellariae of papulosus
(Verrill, 1914, pl. 76, fig. 2b) are wholly inaccurate, being unlike any crossed pedi-
cellariae in Pisaster.

CHECK LIST OF ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS

The following check list contains the names and ranges of all species, sub-
species, and formae found within the geographical limits covered by the present
treatise. After each name is given a reference to part 1 or part 2 of this work,
where the species is treated in detail. In order to avoid repetition, the species
treated in this concluding installment—namely, the Asteriinae—have not been
listed since the names may be found by reference to foregoing pages. Appropriate
citations are given for the few species not treated in parts 1 and 2. References to
Verrill’s Shallow-water Starfishes have been inserted wherever his views or treat-
ment have materially differed from my own. In the Phanerozonia there are few
such citations. In the Forcipulata the very wide differences in treatment are not
apparent in this check list. But the history of each species is covered by the

18 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

oO

synonymy which may be found on the page cited. In order to lessen the confusion
which exists in the Coscinasteriinae and Asteriinae a list of Verrill’s names has been
appended to this check list. In a parallel column is given my interpretation of the
particular species or variety. Further notes will usually be found in the account
of that species or of its formae.

Order PHANEROZONIA Sladen
Suborder PAXILLOSA Perrier
Family PORCELLANASTERIDAE Sladen

EREMICASTER TENEBRARIUS (Fisher), pt. 1, p. 24.
Southern Alaska to the Galapagos Islands; 1,569 to 2,259 fathoms; ooze and
soft mud.
EREMICASTER PACIFICUS (Ludwig), pt. 1, p. 29.
Bering Sea to Gulf of Panama and vicinity of Galapagos Islands; 859 to 1,879
fathoms; ooze, soft mud, fine sand.

Family GONIOPECTINIDAE Verrill

CTENODISCUS CRISPATUS (Retzuis), pt. 1, p. 31.
Bering Sea to Sea of Japan and south along the American coast to Gulf of Pan-
ama; Arctic Ocean; North Atlantic.

Family ASTROPECTINIDAE Gray

LEPTYCHASTER ARCTICUS (Sars), pt. 1, p. 43.

Circumpolar; North Atlantic; Bering Sea, south to Japan.

LEPTYCHASTER PACIFICUS Fisher, pt. 1, p. 45.

Leptychaster millespina Verrill, 1909, p. 553.

Leptychaster pacificus Fisher, Verrill, 1914, p. 326.

Southern part of Bering Sea to Vancouver Island; 56 to 238 fathoms; sand,
pebbles, soft mud.

LEPTYCHASTER ANOMALUS Fisher, pt. 1, p. 48.

Glyphaster anomalus Verrill, 1914, p. 328.

Bering Sea (vicinity Pribilof Islands and west of St. Paul) to southeast Alaska;
on the Asiatic side to the Sea of Japan; 32 to 688 fathoms; fine gray or black sand,
green mud or pebbles.

LEPTYCHASTER INERMIS (Ludwig).

Parastropecten inermis Ludwig, 1905, p. 76; pl. 4, fies cal, 22s ple oil teeters
pl. 22, fig. 126.

Leptychaster inermis Fisher, 1911, p. 53; Clark, 1913, p. 188 (Monterey Bay to
Ballenas Bay).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 189

Monterey Bay to Galapagos—Panama region; 659 to 871 fathoms; gray sand,
green mud. The small specimen of anomalus recorded by me, 1911, p. 52, from
795 to 871 fathoms, Monterey Bay, was probably inermis.”

LEPTYCHASTER PROPINQUUS Fisher, pt. 1, p. 54.

Vicinity of Commander Islands, Bering Sea; 54 to 72 fathoms; green sand.
TROPHODISCUS ALMUS Fisher.

Bull. U. S. Nat. Mus., vol. 52, 1917, p. 368, pls. 28-30.

Southeast coast of Kamchatka, 51° 16’ N., 157° 48’ W.; 96 fathoms, black
sand; northern part of Okhotsk Sea (Djakonov).

TROPHODISCUS UBER Djakonov.
Ann. Musée Zool. Acad. Sci. U. R. S. S., vol. 27, 1926, p. 310, pl. 21, figs. 1, 2.
Southern part of Okhotsk Sea; 30 to 60 fathoms; sand and mud.

DIPSACASTER EXIMIUS Pisher, pt. 1, p. 86.

From north of Monterey Bay to vicinity of San Diego, Calif.; 206 to 525
fathoms; green, black, or yellow mud.
DIPSACASTER BOREALIS Fisher, pt. 1, p. 91.

Bering Sea and south of the Aleutian Islands; 121 to 351 fathoms; sand, green
mud.
DIPSACASTER LAETMOPHILUS Fisher, pt. 1, p. 95.

South of Alaska Peninsula (55° 26’ N., 155° 26’ W.); 695 fathoms; mud.
DIPSACASTER ANOPLUS Fisher, pt. 1, p. 97.

Washington to vicinity of San Diego, Calif.; 300 to 800 fathoms; mud, fine
sand.
PSILASTER PECTINATUS (Fisher), pt. 1, p. 72.

Bering Sea to Bay of Panama; 1,033 to 1,625 fathoms; green ooze, green mud
sand, and hard bottom.
BLAKIASTER RITTERI (Verrill).

Bunodaster ritteri Verrill, Amer. Nat., 1909, p. 554; 1914, p. 322, pl. 84, figs. 1,
la; pl. 104, figs. 1, 2; pl. 105, figs. 1, la; text figs. 15, 16. As Blakiaster, see pt. 1,
p. 40 footnote; also Fisher, Bull. Mus. Comp. Zodl., vol. 54, no. 4, 1911, p. 164.

Although the type is said to be from off San Francisco, it is more likely to have
been taken off San Pedro, Calif. I have seen the type which is very closely related
to Blakiaster conicus Perrier.
ASTROPECTEN ARMATUS Gray, pt. 1, p. 56.

Astropecten siderealis Verrill, 1914, p. 317, pl. 50, fig. 6.

Astropecten braziliensis armatus Déderlein, Die Gattung Astropecten, ete., 1917,
p. 84.
Astropecten erinaceus Clark, 1913, p. 188.
Ecuador to San Pedro, Calif.; shore to 30 fathoms; sand, dark brown mud.

™ [ have examined a specimen of inermis dredged in 425 fathoms of Panama (R 12mm.r6mm.). Ingeneral form it resembles
L. propinqguus, but has broader superomarginal plates, Inermis has six or seven furrow spinelots, amomalus, three or four. For
discussion of the generic position of inermis, seo Fisher, 1911, p. 583. Tho practical difficulty of recognizing Purastropecten for the
short-rayed species of Leptyehaster is there pointed out. Nevertheless Verrill later attempted to maintain two genera (1914, p. 325),
although for Parastropecten he used Glpphaster Verrill, 1909. The generic diagnosis does not cover the known variations of L.enomeles
nor does it apply to the closely related L. propinguus.
63160—30——13

190 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

I have maintained the species in a broader sense than Déderlein who recognizes
erinaceus (including érstedii Liitken) as a distinct subspecies of braziliensis, an
Atlantic species. The type of armatus came from Ecuador, which is also the type
locality of erinaceus (the range of which is given as from Ecuador to Lower Cali-
fornia). A. erinaceus appears to be a forma of armatus rather than a distinct sub-
species. In any event the name armatus can not be restricted to southern Cali-
fornia specimens, if these are really distinct from the race called erinaceus, since the
type of armatus is from Ecuador. Nor does it seem wise to make armatus a subspecies
of a tropical Atlantic form, which is widely separated geographically with a broad
land barrier intervening.

Verrill reports (1914, p. 319) a specimen from the cold water off San Francisco,
which is obviously an error and leads me to think that his Blakiaster ritteri and
Astrometis californica are also from the warm area of southern California.

This species is far too variable to split into races on the strength of trivial char-
acters.

ASTROPECTEN ORNATISSIMUS Fisher, pt. 1, p. 67.

Vicinity of Guadelupe and Cerros Islands, Lower California, north to Santa
Catalina Island and San Pedro, in southern California; 47 to 207 fathoms; fine or
coarse sand, green mud.

ASTROPECTEN VERRILLI CALIFORNICUS Fisher, pt. 1, p. 61.

Astropecien californicus Fisher, 1906c, p. 299; 1911, p. 61.—Verrill, 1914, p. 319.

Astropecten verrilli de Loriol, Déderlein, 1917, p. 85.

From north of Point Reyes, Calif., to northern Lower California; 10 to 244
fathoms, usually on fine sand, but occurring also on coarse sand, and on mud.

Astropecten verrilli seems, biologically, to be a warm-water offshoot of californicus,
which in a typical form inhabits cold water (47° to 52° F.). A. verrilli has a single
longiseries of small tubercles along the superomarginal series. Déderlein classifies
it in his braziliensis group.

THRISSACANTHIAS PENICILLATUS (Fisher), pt. 1, p. 79.

Washington to Los Coronados Islands, Lower California; 277 to 822 fathoms;
green mud or fine sand.
DYTASTER GILBERTI Fisher, pt. 1, p. 101.

Off San Diego, Calif., 2,196 to 2,228 fathoms, gray mud; undoubtedly inter-
grading with D. gilberti demonstrans Ludwig of the Panama region.

Family LUIDIIDAE Verrill

LUIDIA FOLIOLATA Grube, pt. 1, p. 106.

Southeast Alaska to San Diego, Calif., and probably to Mazatlan, Mexico;
10 to 189 fathoms, usually in less than 80 fathoms, and on fine sand or mud; some-
times on hard sand.
LUIDIA LUDWIGI Fisher, pt. 1, p. 113.

Monterey Bay, south to San Pedro, Calif., in 15 to 50 fathoms; mud, sand;
gray sand, shells.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 191

LUIDIA ASTHENOSOMA Fisher, pt. 1, p. 116.
From north of Monterey Bay, Calif., to Los Coronados Islands, Lower California;
11 to 339 fathoms; mud, fine sand, broken shells.

Suborder NOTOMYOTA Ludwig, 1910

Myonola Verrill, 1914, p. 310.
Family BENTHOPECTINIDAE Verrill

PECTINASTER AGASSIZI EVOPLUS (Fisher), pt. 1, p. 123.

Pectinaster agassizi Clark, part, 1913, p. 191.

Off southern California, 984 to 1,059 fathoms, gray and green mud.
LUIDIASTER DAWSONI (Verrill), pt. 1, p. 128.

Bering Sea south along the coast of Alaska to Queen Charlotte Islands; 56 to 159
fathoms; pebbles, sand.

NEARCHASTER ACICULOSUS (Fisher), pt. I, p. 133.

From south of the Alaska Peninsula to Cedros Island, Lower California, in
about 300 to 800 fathoms, usually in about 500 to 700 fathoms; principally on
green mud.

NEARCHASTER VARIABILIS (Fisher), pt. 1, p. 137.

Southern Bering Sea to southeastern Alaska; 108 to 351 fathoms; gray sand,

mud; in shallower water than aciculosus.
NEARCHASTER PEDICELLARIS (Fisher), pt. 1, p. 138.

South of Unimak Island, Alaska, 280 fathoms, green mud, rocks.
MYONOTUS INTERMEDIUS (Fisher), pt. 1, p. Ml.

Monterey Bay, Calif., 958 to 755 fathoms, very soft mud.
BENTHOPECTEN ACANTHONOTUS Fisher, pt. 1, p. Ld.

Southern California, 984 to 1,059 fathoms, mud.

BENTHOPECTEN MUTABILIS Fisher, pt. 1, p. 147.

Off Prince of Wales Island, British Columbia, 1,569 fathoms, gray ooze.
BENTHOPECTEN CLAVIGER Fisher, pt. 1, p. 150,

Southern Bering Sea to Oregon; 987 to 1,064 fathoms; green mud or ooze.

Suborder VALVATA Perrier
Family ODONTASTERIDAE Verrill

PERIDONTASTER CRASSUS (Fisher), pt. 1, p. 154.

Odontaster crassus Fisher, 1911, p. 154.

Peridontaster crassus Koehler, Australasian Ant. Exp. 1911-1914, vol. 8, pt. 1,
1920, p. 195.

Monterey Bay to San Diego, Calif.; 92 to 243 fathoms; gray sand, mud, shells.

Family RADIASTERIDAE Fisher *

" For a discussion of this family see Fisher, Starfishes of the Philippine Seas, ete., 1919, p. 215

192 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Subfamily PRIAMASTERINAE Fisher

GEPHYREASTER SWIFTI (Fisher), pt. 1, p. 175.
From the end of the Aleutian Chain to Washington; 34 to 188 fathoms; rocks.

Family GONIASTERIDAE Forbes

Subfamily PSEUDARCHASTERINAE Sladen

PSEUDARCHASTER PARELII (Diiben and Koren), pt. 1, p. 180.

Bering Sea to Kodiak Island and Sea of Japan; 70 to 351 fathoms; sand, mud,
gravel. Also North Atlantic.
PSEUDARCHASTER PARELI ALASCENSIS Fisher, pt. 1, p. 185.

Oregon to southern Alaska, 68 to 1,064 fathoms; rocks, sand, mud.
PSEUDARCHASTER PUSILLUS Fisher, pt. 1, p. 187.

North of Monterey Bay, Calif., to San Cristobal Bay, Lower California; 54 to
382 fathoms; mud, sand, sand and pebbles.
PSEUDARCHASTER DISSONUS Fisher, pt. 1, p. 192.

Bering Sea to Oregon; 784 to 1,064 fathoms; green mud.

Subfamily GONIASTERINAE Verrill

MEDIASTER AEQUALIS Stimpson, pt. 1, p. 198.

Alaska Peninsula (Chignik Bay) south to northern Lower California; 9 to
160 fathoms; rocks and shells, hard sand, gravel, clay, pebbles, green mud.
MEDIASTER TENELLUS Fisher, pt. 1, p. 202.

Southern California, 291 to 510 fathoms; mud, rocks.

M. transfuga Ludwig is probably a form of this species.

CERAMASTER PATAGONICUS (Sladen), pt. 1, p. 214.

Ceramaster granularis (Retzius), Verrill, 1914; p. 290.

Southern Alaska to the southern part of Bering Sea; 41 to 134 fathoms; sand,
gravel. Also vicinity of Cape Horn; a specimen from Carmen Island, Gulf of Cali-
fornia; one from off Cape San Lucas, 491 fathoms (Clark, 1913, p. 193).

Verrill’s citation of granularis is based on the record of Whiteaves, Straits of
Georgia, 40 fathoms. Whatever west coast material he may have seen was too
imperfect to figure and hence not suitable for critical study. A record by Ives from
Marmot Island, Alaska, is equally open to question, since the same author reported
granularis from Monterey, Calif., where it certainly does not occur (probably
Mediaster aequalis).

Verrill’s figures are all from Atlantic material.

CERAMASTER JAPONICUS (Sladen), pt. 1, p. 206.

Japan to southern Bering Sea, thence south along the American coast to Oregon;
106 to 786 fathoms; green mud, gray sand.

CERAMASTER LEPTOCERAMUS (Fisher), pt. 1, p. 210.

From Point Conception, Calif., to San Cristobal Bay, Lower California; 284 to
594 fathoms; green mud, sand.

CERAMASTER CLARKI Fisher, pt. 1, p. 217.

Southern Bering Sea to southern California; 334 to 600 fathoms; greenish brown
sand.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 193

CERAMASTER ARCTICUS (Verrill), pt. 1, p. 21%.
Tosiaster arcticus Verrill, 1914, p. 292, pl. 50, figs. 3, 3a; pl. 99, figs. 1, 2.
From Bering Island along the Aleutians to Kodiak Island, Alaska; low tide to
102 fathoms.
CLADASTER VALIDUS Pisher, pt. 1, p. 219.
Amukta Pass, Aleutian Islands; 283 fathoms; black sand, rocky.
HIPPASTERIA SPINOSA Verrill, pt. 1, p. 224.
From southern California to Alaska (Kodiak); 27 to 121 fathoms; gray sand,
mud, rocks and mud.
HIPPASTERIA SPINOSA KURILENSIS Fisher, pt. 1, p. 226.
Kuril Islands; 229 fathoms; black sand, pebbles.
HIPPASTERIA LEIOPELTA Fisher, pt. 1, p. 227.
Sea of Okhotsk to southern Bering Sea; 48 to 69 fathoms; sand, pebbles.
HIPPASTERIA LEIOPELTA ARMATA Fisher, pt. 1, p. 230.
Kuril Islands, 229 fathoms (with kurilensis).
HIPPASTERIA HEATHI Fisher, pt. 1, p. 231.
Clarence Straits, Alaska; 206 to 248 fathoms; coarse sand, rocky.
HIPPASTERIA CALIFORNICA Fisher, pt. 1, p. 233.
Southern California to Washington, 266 to 847 fathoms.
CRYPTOPELTASTER LEPIDONOTUS Fisher, pt. 1, p. 238.
Santa Cruz Island, California, to Tres Marias Islands, Mexico (Hippasteria
pacifica Ludwig); 266 to 680 fathoms; sand, gravel.

Family LINCKIIDAE Perrier

LINCKIA COLUMBIAE Gray, pt. I, p. 242.
Southern California and Lower California; Colombia, west coast; Galapagos
Islands; shore to 40 fathoms.

Family PORANIIDAE Perrier

DERMASTERIAS IMBRICATA (Grube), pt. 1, p. 209.
Including Dermasterias imbricata var. valvulifera Verrill, 1914, p. 308, pl. 97,
figs. 2, 2b.
Sitka, Alaska, to Monterey Bay, Calif.; at and just below low tide, rocks.

Order SPINULOSA Perrier
Family ASTERINIDAE Gray

PATIRIA MINIATA (Brandt), pt. 1, p. 254.

Asterina miniata Fisher, 1911, p. 254.

Patiria miniata Verrill, 1914, p. 264.

Sitka, Alaska, all the way to San Diego, Calif., thence to the Gulf of California,
north of La Paz; low water to 165 fathoms; rocks, shells, gravel, hard sand.

194 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Family ECHINASTERIDAE Verrill ®

PORANIOPSIS INFLATA (Fisher), pt. 1, p. 261.

Oregon to San Diego, Calif.; 26 to 159 fathoms; rocks, sand, green mud.

This species undoubtedly ranges far south of California, and in deep water it
gradually changes to flexilis.

PORANIOPSIS INFLATA FLEXILIS Fisher, pt. 1, p. 261.

Central and southern California; 334 to 600 fathoms.

This form probably ranges to the Galapagos Islands where it has been described
as P. mira (Alexandraster mirus Ludwig). P. mira (Ludwig) preoccupied in Porani-
opsis by de Loriol’s P. mira (Fisher, 1911, p. 261).

HENRICIA SANGUINOLENTA (0. F. Miiller), pt. 1, p. 271.

Bering Sea south to the vicinity of the Kuril Islands and on the American coast
to Washington. Circumpolar and North Atlantic. In the north Pacific region, 30
to 229 fathoms; sand, mud, gravel.

The above includes variety pectinata Verrill (Cribrella pectinata Verrill), 1914, p.
230, Eastport, Me. Variety rudis Verrill, 1914, p. 233, Point Franklin, Arctic
Ocean, 13% fathoms, sand.

There seems to be little use in naming the legion of variants of this species. It
is questionable whether pectinata of Eastport, Me., is the same as the Bering Sea
variation resembling it.

HENRICIA SANGUINOLENTA forma TUMIDA Verrill. .

Henricia tumida Verrill, 1909, pp. 554, 555, fig. 5; 1914, p. 234, pl. 12, figs. 1, 2.

Henricia sanguinolenta eschrichtii Fisher, 1911, p. 276, pl. 67, figs. 1-3; pl. 68,
figs. 1-2.

Henricia tumida borealis Verrill, 1914, p. 236, pl. 12, figs. 3, 4.

Henricia arctica Verrill, 1914, p. 239, pl. 87, figs. 3-8c, text fig. 13.

Arctic Ocean north of Bering Strait south to Kodiak; westward along Alaskan
Peninsula and Aleutian Islands to Bering Island and Kamchatka, to Sea of Okhotsk
(Brandt), and south to Simushir, Kuril Islands. Low tide to 53 fathoms.

This form is exactly equivalent to eschrichtii of part 1. Verrill’s name has
been adopted since the type is from Unalaska, while that of eschrichtii was from
Greenland; and since some doubt exists that the two forms are precisely the same.
They are however analogous stouter rayed shallow-water phases of the wide ranging
sanguinolenta.

Verrill applied three names to this forma—tumida, borealis, arctica. He attributed
some importance to the temporarily arched disk of brooding females. His H. tumida
borealis applies to specimens with slenderer rays. Both stout and slenderer rayed
forms occur in the same tide pool and are in several lots of specimens, as for instance,
Nikolski, Bering Island. The narrower rayed variants intergrade with sanguino-
lenta. Rarely, for reasons unknown, the rays are short, broad, and flaccid. (Attu
Island, Fisher, 1911, pl. 68, fig. 2.) Verrill attributes this specimen to Cape Lis-

” Verrill described Echinaster robustus (1914, p. 207), from Sooke, Vancouver Island. I have examined the type; and Dr. H.
L. Clark, a photograph of the type furnished by Dr. W. R. Coe. Neither Clark nor I believe that robustus came from Vancouver
Island nor from a locality within a thousand miles of Vancouver Island. If “Echinaster robustus” is not a genuine new species
from Mexico, or south, it is likely an extra ‘fat’ Echinaster sentus from Florida. Much of Verrill’s material was without attached
labels and nothing would have been easier than an inadvertent shift. This actually occurred in the case of “‘ Pisaster lutkenii’”’ var,
australis. Equally unlikely are all records of Echinaster tenuispinus from California and north.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 195

burne, Arctic Ocean, and refers it to his H. arctica, based on a worn specimen. In
my opinion HZ. arctica is but a sporadic variant of tumida with the plates and spinelets
smaller than usual. The Attu Island example shows that the variation may occur
far from the Arctic. Verrill distinguishes arctica also by the large number of adam-
bulacral spinelets, in ‘‘several’’ transverse rows. But his Figure 3a, Plate 87, shows
only two transverse rows. Although young specimens of tumida often have only one
transverse series of spinelets, large examples generally have two. Two specimens
from the same lot as Verrill’s type of arctica were recorded by me (1911, p. 278, Cape
Lisburne, Henry D. Woolfe) and I find no reason to consider them specifically different
from the very variable, unstable tumida.

Verrill also listed typical tumida as sanguinolenta, namely, the specimen from
Bering Island, N. Grebnitsky (Verrill, 1914, p. 229). I have an example of the same
lot which is referable to tumida.

HENRICIA LEVIUSCULA (Stimpson), pt. 1, p. 280.

Including Henricia leviuscula var. lunula Verrill, 1914, p. 218, pl. 88, figs. 2—-2c;
var. inequalis Verrill, 1914, p. 219, pl. 88, figs 1, la; var. spatulifera Verrill, 1914,
p. 224, pl. 5, fig. 1; pl. 14; text fig. 12; other synonyms as given in pt. 1, p. 280.

Aleutian Islands to Monterey Bay, Calif., in a typical form, and thence south to
the Santa Barbara Channel and San Diego, Calif., the southern forms not typical.
The typical form is found along shore at low tide, or in comparatively shallow water.
Low tide to 53 fathoms.

HENRICIA LEVIUSCULA forma DYSCRITA Pisher.

Henricia leviuscula dyscrita Fisher, 1911, p. 289, pl. 74, figs. 1-5.

Henricia leviuscula var. dyscrita Verrill, 1914, p. 223, pl. 12, fig. 6.

Middle and southern California, offshore, 21 to 80 fathoms; sand, stones, mud,
and rocks.

This may eventually prove to be a good deep water race of leviuscula. It is
however rather unstable and a more conservative treatment is to consider it a forms
of leviuscula in some respects intermediate with spiculifera.

HENRICIA LEVIUSCULA SPICULIFERA (Clark).

Cribrella spiculifera Clark, Proc. Boston Soc. Nat. Hist., vol. 29, 1901, p. 328,
pl. 2; pl. 4, fig. 1.

Henricia leviuscula multispina Fisher, 1911, p. 286, pl. 72, figs. 1-4; pl. 73,
figs. 1, 2.

Henricia spiculifera (Clark), Fisher, 1911, p. 295.

Henricia leviuscula spiculifera Verrill, 1914, p. 220.

Henricia leviuscula var. multispina Verrill, 1914, p. 222.

From Oregon to Bering Sea, Bering Strait, the Aleutians, and south on the
Asiatic side to Simushir, Kuril Islands; low tide to 238 fathoms.

Plate 72, Figures 1 and 2, and Plate 73, Figures 1 and 2, of part 1, give an
excellent idea of this relative of leviuscula. Verrill unfortunately figured a Bering
Island specimen (of which I have duplicates), instead of Clark’s types, which he held
for many years. These types came from Puget Sound and are evidently not extreme
examples of the race, but are probably nearer to leviuscula than is the type of multi-

196 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

spina. The race as a whole reaches its best development along the Aleutian chain,
the type locality of multispina.

Typical examples are readily distinguished from leviuscula by their delicate and
more numerous, rather hyaline spinelets, ending in several delicate thorns. Varia-
tions with larger or smaller abactinal plates and slender to stout rays occur. I do
not think it is possible to distinguish a northern multispina from a southern spiculifera
and have accordingly united the two.

HENRICIA ASPERA Fisher, pt. 1, p. 293.

Bering Sea (Bering Island, Pribilofs, Aleutian Islands) south along the coast to
the Santa Barbara Channel, Calif.; 26 to 313 fathoms; mud, sand, pebbles, shells,
rocks.

HENRICIA ASPERA forma ANNECTENS Fisher.

Henricia leviuscula annectens Fisher, 1911, p. 291, pl. 70, fig. 2d; pl. 71, figs. 1, 3.

Henricia leviuscula var. annectens Verrill, 1914, p. 224.

Aleutian Islands to Santa Barbara Channel, Calif., 21 to 135 fathoms; rocks,
shells, sand, mud.

This form in most respects is intermediate between aspera and leviuscula but
resembles aspera more than leviuscula. Specimens referable to this forma have been
taken also at station 3220, near Unalaska, 34 fathoms; and station 4784, near Attu,
Aleutian Islands, 135 fathoms. The range is probably coextensive with that of
aspera.

HENRICIA ASTHENACTIS Fisher, pt. 1, p. 297.

Vicinity of Santa Barbara Island, Calif.; Shumagin Islands; Bering Sea (Bowers
Bank and off Kamshatka); 178 to 682 fathoms; green and gray sand, green mud,
black mud.

HENRICIA LONGISPINA Fisher, pt. 1, p. 299.

From vicinity of Vancouver Island to southern Alaska; 41 to 134 fathoms; soft

green mud, gravel, sponges.

HENRICIA LONGISPINA ALEUTICA Fisher, pt. 1, p. 300.
Off Attu Island, Aleutian Islands; 135 fathoms; coarse pebbles.

HENRICIA POLYACANTHA Fisher, pt. 1, p. 302.
Cyllaster polyacantha A. H. Clark, Proc. Biol. Soc. Washington, vol. 29,1916, p.61.
Off San Diego, Calif.; 359 fathoms; mud, fine gray sand.

HENRICIA CLARKI Fisher, pt. 1, p. 303.

Cyllaster clarki A. H. Clark, Proc. Biol. Soc. Washington, vol. 29, 1916, p. 61.

Santa Cruz Island, Calif., to the Revillagigedo Islands, Mexico; 124 to 822
fathoms; black sand, fine gray sand, green mud, black mud.

Family SOLASTERIDAE Perrier
SOLASTER ENDECA (Linnaeus), pt. 1, p. 307.
Solaster endeca and Solaster galaxides Verrill, 1914, pp. 244, 248; grlaxides, pl.
46, figs. 2, 2a; pl. 87, figs. 5-5c; pl. 89 fig. 2.
Bering Sea south to Victoria, British Columbia, and San Juan Islands, Wash.;

12 to 238 fathoms; gray sand, gray green mud, coarse sand. In the North Atlantic
and Arctic Ocean. (See von Hofsten, 1915, p. 39.)

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 197

Verrill’s galarides was founded upon three specimens from the vicinity of Vie-
toria, British Columbia, and Puget Sound, Wash. As pointed out by me in 1911
(p. 308) there is no constant difference between endeca and galazides, the distinctions
listed by Verrill being rather of the nature of minor individual variations and not
constant,

It is probable that the type is now the only one of Verrill’s three specimens
which is extant.

SOLASTER STIMPSONI Verrill, pt. 1, p. 207.

Solaster stimpsoni Verrill, 1914, p. 254, pl. 10, figs. 1, 2; pl. 11, figs. 1, 2; pl.
15, figs. 1, 2; pl. 46, figs. 1, 1c; pl. 94, fig. 2; pl. 95.

Solaster constellatus Verrill, 1914, p. 257, pl. 46, figs. 3, 4; pl. 90, fig. 2; pl. 93;
pl. 94, fig. 1.

Southern Bering Sea (Commander Islands), and Kodiak, south to Oregon, in
shallow water; low tide to 33 fathoms; rocks and shells.

Since the publication of part 1, I have observed abundant living material at
Departure Bay (Canadian Biological Station). I have also examined the type of
constellatus kindly forwarded by the Provincial Museum, Victoria. In spite of
Verrill’s vigorous defense (1914, p. 258) of constellatus, the fact remains that the
alleged species was based upon a single specimen from Puget Sound, from which
general region I have handled probably ten times the amount of material available
to Verrill. The type does not differ sufficiently from the variable stimpsoni (many
of which have nine rays) to constitute even a good forma. The larger pseudopaxillae
upon which Verrill lays stress are apparent only, due to their having dried with the
spinelets fully spread apart. They are well within the normal variation of stimpsoni.
It is simply a question of adequate material,

SOLASTER DAWSONI Verrill, pt. 1, p. 213.

Monterey Bay to the Aleutian Islands, thence to the Kuril Islands; low tide to
229 fathoms; rocks (Monterey), sand, shells, black sand, green mud.

SOLASTER DAWSONI ARCTICA Verrill.

Verrill, 1914, p. 253, pl. 87, figs. 6, 6a; text fig. 14.

Solaster dawsoni part, Fisher, 1911, p. 315 (Point Franklin); pl. 85, fig. 1
(topotype).

Point Franklin, 10 miles west, 13 fathoms, sand.

Verrill’s type is one of the specimens collected by the Point Barrow expedition
(John Murdoch) and is of the same lot as the two recorded by me. It seems
probable that this is an arctic race of dawsoni.

SOLASTER PAXILLATUS Sladen, pt. 1, p. 315.

From south of Yokohama, Japan, to Bering Sea, thence to vicinity of Kodiak
Island, Alaska; 56 to 350 fathoms; gray sand, pebbles, mud.

It is possible that Clark’s record of this species (1913, p. 196) from off Santa
Rosa and Santa Cruz Islands, Calif., 534 to 640 fathoms, refers to S. exiguus.
SOLASTER EXIGUUS Fisher, pt. 1, p. 319.

Off Anacapa Island, Calif.; 603 fathoms, green mud.

63160—30——14

198 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

SOLASTER BOREALIS (Fisher), pt. 1, p. 320.

From off San Diego, Calif., to Bering Sea, thence to Honshu, Japan; 225 to
1,044 fathoms; blue, green, brown, gray, black mud.

SOLASTER HYPOTHRISSUS Fisher, pt. 1, p. 324.

Shumagin Islands to Attu, Aleutian Islands; 135 to 625 fathoms; green mud,
coarse pebbles.

CROSSASTER PAPPOSUS (Linnaeus), pt. 1, p. 325.

Solaster papposus, Fisher, 1911, p. 325.

Circumpolar, through Bering Strait, into Bering Sea; thence south along the
west coast of North America to Washington; on the Asiatic side to the Sea of Okhotsk.
North Atlantic. (See von Hofsten, 1915, p. 32.) In the north Pacific region,
lowest tide to 283 fathoms; from a variety of bottom including gravel, rocks and
shells, sand, mud.

HETEROZONIAS ALTERNATUS (Fisher), pt. 1, p. 331.

Washington to San Diego; 316 to 603 fathoms; green and black mud, fine sand,
LOPHASTER FURCILLIGER Fisher, pt. 1, p. 334.

From south of the Alaska Peninsula to southern California, thence to the Gala-
pagos Islands; 192 to 1,100 fathoms; green mud, sand, globigerina; also rocks, shells,
fine gray sand.

LOPHASTER FURCIFER VEXATOR Fisher, pt. 1, p. 338.

Lophaster furcilliger verator Fisher, 1911, p. 338.

Southern Bering Sea to northern California; 75 to 350 fathoms, typically in
less than 200; black sand, mud, gravel.

Family KORETHRASTERIDAE Danielssen and Koren

PERIBOLASTER BISERIALIS Fisher, pt. 1, p. 341.
Southern Bering Sea and off southern California; 57 to 440 fathoms; green
and gray mud, fine gray sand, pebbles, rocky.

Family PTERASTERIDAE Perrier

PTERASTER MILITARIS (0. F. Miller) pt. 1, p. 346.
Pteraster militaris Verrill, 1914, p. 272.
Pterasterides aporus (Ludwig) Verrill, 1914, p. 277.
Bering Sea to Washington; 41 to 344 fathoms; green mud, gray sand, gravel.
Arctic; North Atlantic. (See von Hofsten, 1915, p. 47.)
PTERASTER TRIGONODON Fisher, pt. 1, p. 348.
Off Santa Cruz Island, Calif.; 447 to 510 fathoms; black mud, rocks.
PTERASTER JORDANI Fisher, pt. 1, p. 350.
From northern Lower California to Washington; 266 to 984 fathoms; mud or
sand.
PTERASTER MARSIPPUS Fisher, pt. 1, p. 352.
Bering Sea, 52 to 351 fathoms; gravel, sand.
PTERASTER COSCINOPEPLUS Fisher, pt. 1, p. 354.
Off southern and central California; 281 to 1,062 fathoms; mud, sand.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 199

PTERASTER TEMNOCHITON Fisher, pt. 1, p. 255.

Aleutian Islands, Attu to Unalaska; 56 to 135 fathoms; pebbles.
PTERASTER PULVILLUS Sars, pt. 1, p. 355.

Near Bering Island, Bering Sea; 72 fathoms; pebbles. Also Arctic Ocean and
North Atlantic (off Maine to Newfoundland); Norwegian coast, from 60° to Fin-
mark; Barents Sea; Kara Sea.

PTERASTER TESSELATUS Ives, pt. 1, p. 959.

Pteraster tesselatus Ives, Proc. Acad. Nat. Sci. Phila., 1888, p. 421.—Fisher, 1911,
p. 359, pl. 104, figs. 1-5.—Verrill, 1914, p. 268, pl. 32, figs. 1, 2; pl. 86, figs. 4~—4e;
pl. 97, fig. 1.

Pteraster multispinus Clark, Proc. Boston Soc. Nat. Hist., vol. 29, no. 15, 1901,
p. 328, pl. 3, figs. 1, 2—Fisher, 1911, p. 359.—Verrill, 1914, p. 271.

? Pteraster gracilis Clark, Proc. Boston Soc. Nat. Hist., vol. 29, 1901, p. 326,
pl. 3, figs. 3, 4.—Fisher, 1911, p. 349.—Verrill, 1914, p. 271.

Pteraster tesselatus hebes Verrill, Amer. Journ. Sci., vol. 28, 1909, p. 61; 1914,
p. 270, pl. 96, figs. 1, 2.

Bering Sea (Bering Island, Aleutian Islands, Pribilofs, Hagemeister Island)
south to Strait of Fuca and Strait of Georgia; gravel, sand, mud.

Verrill has given some notes on the types of Pteraster multispinus and gracilis,
both young specimens; neither, in my opinion, are valid species. It is possible that
‘gracilis is the young of militaris. Verrill’s hebes is a dried and distorted tesselatus
from Departure Bay, British Columbia, where typical tesselatus has been collected.
PTERASTER TESSELATUS ARCUATUS Fisher, pt. 1, p. 363.

Monterey Bay, 56 to 46 fathoms, rocks.

PTERASTER OBSCURUS (Perrier), pt. 1, p. 363.

Pteraster obscurus (Perrier), Fisher, 1911, p. 363 (synonymy as given).—Verrill,
1914, p. 274.

Pteraster obscurus var. octaster Verrill, 1914, p. 276.

Bering Sea, from Bering Straits south to Bering Island, Kamchatka, and Una-
laska; 17 to 85 fathoms; mud, black sand, fine gray sand. Also 50 to 145 fathoms
in the North Atlantic, whence it ranges into the Arctic (Spitzbergen, Greenland).
Evidently cireumpolar. (See von Hofsten, 1915, p. 50.)

DIPLOPTERASTER MULTIPES (Sars), pt. 1, pl. 371.

In the north Pacific from San Diego, Calif., and Suruga Gulf, Japan, north to
southern part of Bering Sea; 81 to 640 fathoms; mud, sand, gravel. Also in the
North Atlantic, on the west side from 35° to 45° north latitude, and on the east side
from Norway and Barents Sea; 50 to about 580 fathoms.

HYMENASTER KOEHLERI Fisher, pt. 1, p. 373.

Southern Bering Sea (between Unimak Island and the Pribilofs); 1,771 fathoms;
blue ooze.

HYMENASTER PERISSONOTUS Fisher, pt. 1, p. 376.

From off Rosario Bay, Lower California, to Bering Sea; 225 to 1,771 fathoms;
gray mud, blue and green ooze.
HYMENASTER QUADRISPINOSUS Fisher, pt. 1, p. 389.

Bering Sea to Rosario Bay, Lower California, and probably south to the vicinity
of Panama; 1,059 to 1,771 fathoms; green mud, gray and blue ooze.

200 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Order FORCIPULATA Perrier

Suborder BRISINGINA Fisher
Family BRISINGIDAE Sars

CRATEROBRISINGA SYNAPTOMA Fisher, pt. 2, p. 10.

Off British Columbia (52° 39’ 30’’ N., 132° 38’ W.); 1,588 fathoms; gray ooze,
coarse stones.
BRISINGELLA EXILIS (Fisher), pt. 2, p. 13.

Off southern California from San Diego to Santa Barbara Island; 448 to 1,059
fathoms; green mud.
BRISINGELLA PUSILLA Fisher, pt. 2, p. 16.

Off southern California from San Diego to Santa Cruz Island; 301 to 1,059
fathoms; green and black mud.
BRISINGELLA PANNYCHIA Fisher, pt. 2, p. 18.

Bowers Bank, Bering Sea; 771 fathoms; green mud.
FREYELLASTER FECUNDUS (Fisher), pt. 2, p. 21.

Off Point Pinos, Monterey Bay, Calif.; 755 to 1,062 fathoms; soft gray mud,
hard sand, and mud.
FREYELLA MICROPLAX (Fisher), pt. 2, p. 25.

British Columbia to central California; 861 to 1,588 fathoms; mud and sand.
FREYELLA INSIGNIS Ludwig, pt. 2, p. 27.

Gulf of Panama to southern California; 1,772 to 2,228 fathoms; on gray mud,
green mud, brown mud, and globigerina ooze.
ASTROCLES ACTINODETUS Fisher, pt. 2, p. 29.

Off British Columbia (55° 20’ N., 136° 20’ W.); 1,569 fathoms; gray ooze.

Suborder ASTERIADINA Fisher
Family ZOROASTERIDAE Sladen

ZOROASTER OPHIURUS Fisher, pt. 2, p. 34.

Bering Sea to Lower California (off Ballenas Bay and San Rosario Bay); 879 to
1,217 fathoms; mud and fine black sand.

ZOROASTER ACTINOCLES Fisher, pt. 2, p. 37.

Northwest of Yunaska Island, Aleutian Islands; 1,217 fathoms; fine black sand.
ZOROASTER EVERMANNI Fisher, pt. 2, p. 40.

Off southern California, San Diego to Santa Cruz Island; 216 to 510 fathoms;
green mud, black mud, gray sand.

ZOROASTER EVERMANNI MORDAX Fisher, pt. 2, p. 43.

From Washington to southern California; 239 to 760 fathoms; in south only in
over 600 fathoms; green mud.
MYXODERMA SACCULATUM Fisher, pt. 2, p. 45.

Bowers Bank, Bering Sea, and off central California, undoubtedly intergrading
with the southern form from off southern California; 550 to 766 fathoms; green mud,
sand.

MYXODERMA SACCULATUM ECTENES Fisher, pt. 2, p. 49.

Off southern California from the vicinity of Santa Cruz Island to Los Coronados

Islands; 510 to 100 fathoms; gray mud, green mud, black mud.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 201

MYXODERMA PLATYACANTHUM (1H. L. Clark), pt. 2, p. 82.
From Point Conception, Calif., to San Cristobal Bay, Lower California; 205 to
287 fathoms; green mud.
MYXODERMA PLATYACANTHUM RHOMALEUM Fisher, pt. 2, p. 54.
From off Oregon (43° 46’ N.) to central California; 277 to 296 fathoms; gray
sand. Intergrades off San Luis Obispo County with platyacanthum.

Family ASTERIIDAE Gray

Subfamily PepICELLASTERINAE Fisher
PEDICELLASTER MAGISTER Fisher, pt. 2, p. 59.

Southern Bering Sea, from the Commander Islands to Unimak, and north to 57°;
south along the Alaskan coast to Kasaan Bay, Prince of Wales Island; 12 to 121
fathoms; gravel, black and gray sand; green mud.

PEDICELLASTER MAGISTER MEGALABIS Fisher, pt. 2, p. 64.

From vicinity of San Diego, Calif., to Washington; 284 to 530 fathoms; green
and yellow mud, or black sand and gravel.
ANTELIASTER COSCINACTIS Fisher, pt. 2, p. 70.

Off Santa Cruz Island, Calif.; 447 to 510 fathoms; black mud, rocks.
ANTELIASTER MICROGENYS Fisher, pt. 2, p. 73.

South of Santa Cruz Island, Calif.; 48 fathoms; fine gray sand.

ANTELIASTER MICROGENYS NANNODES Fisher, pt. 2, p. 74.

Bowers Bank, Bering Sea; 247 fathoms; gray sand, green mud.
TARSASTER ALASKANUS Fisher, pt. 2, p. 78.

Off Indian Point, vicinity Naha Bay, Behm Canal, Alaska; 240 to" 108 fathoms;
rocky.

AMPHERASTER MARIANUS (Ludwig), pt. 2, p. 81.

From Tres Marias Islands, Mexico, to Washington; 277 to 676 fathoms; black,
green, yellow mud, gray sand, rocks.
AMPHERASTER CHIROPLUS Fisher, pt. 2, p. 84.

Off Santa Cruz Island, Calif.; 447 to 510 fathoms; black mud, rocks.
AMPHERASTER ATACTUS Fisher, pt. 2, p. 8%.

Off Los Coronados Islands, vicinity of San Diego, Calif.; 266 to 323 fathoms;
gray sand.

Subfamily LABIDIASTERINAE Verrill
RATHBUNASTER CALIFORNICUS Pisher, pt. 2, p. $9.

Off California, from vicinity of San Diego to Point Arena; 207 to 369 fathoms;

mud, gray sand, black sand, mud and shale.

Subfamily COSCINASTERIINAE Fisher

STYLASTERIAS FORRERI (de Loriol), pt. 2, p. %.
Southern Alaska to San Diego, Calif.; 16 to 291 fathoms; usually on rocky
bottom.
DISTOLASTERIAS NIPON (Dédertein), pt. 2, p. 103.
Japan Sea (Peter the Great Bay); Japan; Hong Kong; low tide to 81 fathoms.
SCLERASTERIAS HETEROPAES Pisher, pt. 2. p. 112.
Half Moon Bay to south of San Diego, Calif.; 27 to 85 fathoms; fine sand,
rocks, mud.

202 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

ASTROMETIS SERTULIFERA (Xantus), pt. 2, p. 119. : :
Gulf of California to Santa Barbara, Calif.; very low tide to 33 fathoms.

ASTROMETIS CALIFORNICA (Verrill), pt. 2, p. 126. ,

Said to be from off San Francisco, but more likely from off San Pedro, Calif.
Probably only a variant of sertulifera.

LETHASTERIAS NANIMENSIS (Verrill), pt. 2, p. 132.

Strait of Fuca and Strait of Georgia; 25 to 40 fathoms; gray sand, pebbles, mud.
LETHASTERIAS NANIMENSIS CHELIFERA (Verrill), pt. 2, p. 134.

Saghalien to Bering Strait, thence to Gulf of Alaska off Kodiak Island (Kam-
chatka, Commander Islands, Aleutian Islands, Bering Sea, Alaska Peninsula); low
tide to 93 fathoms; sand, gravel, sand and mud, clayey mud.

ORTHASTERIAS KOEHLERI (de Loriol), pt. 2, p. 139.

Yakutat Bay, Alaska, to Santa Rosa Island, Calif.; lowest tide and subtidal zone

to 125 fathoms; rocks, rocks and sand or mud.

forma KOEHLERI, pt. 2, p. 145.
Distribution of the species.

forma LEPTOSTYLA Fisher, pt. 2, p. 145.

Strait of Fuca; Behm Canal, Alaska; 39 to 100 fathoms; green mud, pebbles,
rocks.

—— forma BIORDINATA Verrill, pt. 2, p. 146.

Vancouver Island; low tide; rocks.

—— forma MONTEREYENSIS Fisher, pt. 2, p. 146. :
Monterey Bay; subtidal zone; very rocky situations.

Subfamily PYCNOPODIINAE Verrill

LYSASTROSOMA DESMIORA Clark, pt. 2, p. 151.

Gulf of Tartary.

PYCNOPODIA HELIANTHOIDES (Brandt), pt. 2, p. 154.

Unalaska, Aleutian Islands, to San Diego, Calif.; intertidal, in rocky situations,
to 238 fathoms; sand.

Lee Boone * has recorded from Espiritu Santo Island, Gulf of California, under
the name Heliaster kubinii (pl. 4), a specimen of Pycnopodia helianthoides which
can not be located in the Bingham Oceanographic Collection. In a letter, Mr.
A. E. Parr, in charge of the collection, informs me that some of the material of this
expedition was inadequately labeled. Locality highly improbable for Pycnopodia.

Subfamily AsrermNak& (Verrill, part)

All species and formae of this subfamily believed to be valid are treated in this
volume and may be taken, seriatim, for a check list.

4 Echinoderms from the Gulf of California and the Perlas Islands, Bulletin Bingham Oceanographic Collection, vol. 2, art. 6,
Dec., 1918, p. 7, pl. 4,

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 203

LIST OF SPECIES, SUBSPECIES, AND VARIETIES OF ASTERIIDAE IN VERRILL’S
SHALLOW-WATER STARFISHES, 1914, WITH THEIR EQUIVALENT IN THE
PRESENT MONOGRAPH ®

Pisaster ochraceus, 69............------ P. ochraceus.
P. ochraceus segnis.
——— ochraceus var. nodiferus, 72.......- P. ochraceus forma nodiferus.
an ONT NIC le ice one ee P. ochraceus forma confertus.
—— fissispinus, 76..........-..-.----- P. ochraceus forma ochraceus.
—— brevispinus, 77-.-..-.-.--..---.-- P. brevispinus forma brevispinus.
ames CR ULEEUD OU sep pre ug ween lg, sl P. giganteus capitatus (part).
P. giganteus (part).
SOI I ase i es aii wih Ss wah Do.
—— litkeni var. australis, 88._......._- Do.
—— giganteus, 89..._..........._..--- Do.
—— papulosus, 91. _.....-..-.......--. P. brevispinus forma paucispinus.
ee, ee a ee Pee Leptasterias polaris katherinae.
—— poucispinus, 98...-..----...-.-.-- P. brevispinus forma paucispinus.
Marthasterias sertulifera, 100_.......---- Astrometis sertulifera (part).
Asterias victoriana, 102._.......-..-.--- Evasterias troschelii forma troschelii.
—— polythela, 104. ............--....- Leptasterias polaris acervata forma polythela.
—— nanimensis, 105_.-.-..--.-------- Lethasterias nanimensis.
mee IIE WOTE TIONG alas rca tow see wets ar Leptasterias polaris acervata forma acervata.
—— katherinae, 112..............--.-- Leptasterias polaris katherinae.
—— mulliclava, 114. ..........-..-..-- Leptasterias camtschatica.
L. camtschatica dispar forma nitida.
Leptasterias inaequalis, 117_......------ Evasterias troschelii, young.
— leptalea, 119_._.......-.....--..- L. leptalea.
GPOED RAO oe ios oa aes se tem ne L. arctica
am GE Le So ee ee eee SAT! L. coei (part).
cae CIID A Mg dk Wate lel ate lingo Evasterias troschelii.
—— vancouveri, 125..................- L. hexactis vancouveri.
eee UNE GUO ds nce cape an ans does ach na caro i L. hexactis forma heractis.
L. pusilla.
—— aequalis, 127.........-.-.-------- L. aequalis forma aequalis.
—— aequalis var. compacta, 130__.._---- Do.
—— — var. concinna, 132_....._.---- Do.
— — var. nana, 132___.___.......- L. aequalis forma nana.
—— epichlora, 132. ...........--..---- L. hexactis (part).
L. alaskensis (part).
—— epichlora alaskensis, 136.........-- L. alaskensis (part).
—— — alaskensis var. carinella, 137 __- Do.

—— — miliaris var. regularis, 139_... L. hezactis forma regularis.
—— —— miliaris var. subregularis, 139 *

—— —— miliaris var. subnodulosa, 139_. Evasterias troschelii, young.
—— —— plena, 140__..._......-..-.. L. hexactis forma plena.
—— —— pugetana, 142 ___._...._.-..- L. hezactis forma regularis.

% These are given in the order of their occurrence; the figure following the name is the page upon which the description occurs
In the right column is the equivalent form.

* This “race” probably includes at least two distinct forms of which the six rayed may be young J. Aesectis forma plene while
the five myed is not improbably young Erasterias troschelii forma alevolata or acanthostoma, No definite type was indicated and no
Specimen can be found. The measurements indicate » young specimen (RK 16 mm., r 5 mm.).

% Type lost. This is another young specimen, probably an individual variant of some form of L. Acrectis; possibly it ls a young
six-tayed Eresterias as two actinal rows of spines are indicated—very unusual for a small Herasterias.

204 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Leptasterias dispar, 142... ________-_.-- L. camtschatica dispar forma dispar.
———oblectas 1444 0S Se ee L. groenlandica forma groenlandica.
Stenasterias macropora, 145_----------- _ §. macropora.
Stephanasterias albula, 147_.---------- _ S. albula.
Ctenasterias cribraria, 148_--_----- ae. Leptasterias groenlandica forma cribraria.
Evasterias troschelii, 151... ------------- E. troschelit.
— troschelii var. rudis, 158_---------- E. troschelii forma troschelii (part).
—— troschelii var. densa, 161__--------- E. troschelii forma acanthostoma, young.
- troschelit var. alveolata, 162-------- E. troschelit forma alveolata (part).
— troschelii var. subnodosa, 163____-_- E. troschelit forma troschelii (part).
—— troschelti var. parvispina, 163_------ E. troschelii forma alveolata, young.
——— gcanthostoma, 165. 28-2252 -2=2=555" E. troschelii forma acanthostoma.
Orthasterias columbiana, 168------------ O. koehleri.
—piorainata Wiohen ene e a eee O. koehleri forma biordinata.
— californica, 174..2---2- -. -=_ __ Astrometis californica.
= roehlent wl Dee ee ee O. koehleri.
———=dawsont, V5: = 325 = ote eee Astrometis sertulifera (part).
— MENLO MIU wll eae ee en Leptasterias coet (part).
Leptasterias polaris katherinae (part).
ST OTR ENUM Omen etn ee ens ee Stylasterias forreri.
— forreri forcipulata, 180_------------ Do.
—— leptolena, 182. - 5... 22-2 s=scs == Do.
= WJONOIENG, LO4-= 2 aan mae ee Astrometis sertulifera (part).
Distolasterias chelifera, 185------------- Lethasterias nanimensis chelifera.
Parasterias albertensis, 187_-_----------- Asterias amurensis.
Allasterias rathbuni, 189_._____-__--_---_ Do.
— rathbuni nortonensis, 191_________-- Do.
—ATUOMALL, Lote ae eee Asterias amurensis forma anomala.
Pycnopodia helianthoides, 198___ .-----_- P. helianthoides.

SUMMARY OF THE ASTERIINAE, NOTASTERIINAE, AND NEOMORPHASTERINAE
OF THE NORTHERN HEMISPHERE ®

This summary, like that for the southern hemisphere, is not a revision but a
review. It will perhaps prove useful as a background for the foregoing detailed
account of the north Pacific fauna. To avoid repetition of matter, all genera and
species which have been treated in the main body of this work are followed by the
page on which fuller information may be found.

Subfamily ASTERTINAE (Verrill, part)
Genus APHANASTERIAS Fisher, p. 159

1. APHANASTERIAS PYCNOPODIA Fisher, p. 160.
Genus APHELASTERIAS Fisher

Aphelasterias Fisner, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, p. 602. Type, Asterias
japonica Bell.

Diagnosis —Diplacanthid Asteriinae resembling 5-rayed Stephanasterias; abac-
tinal plates small, numerous, imbricated, with small papular areas, not longiserially
arranged but in somewhat irregular transverse series, on either side of a definite
carinal series; no actinal plates; abactinal spinelets small (surrounded by circlets of

+s Genera arranged alphabetically; for key, see page 3.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 205

small crossed pedicellariae) isolated or in small groups, the general surface having a
neat appearance; superomarginals and inferomarginals regular, larger than abac-
tinals, each bearing a transverse comb of three or four spines (the inferomarginals
the larger) which carry groups of crossed pedicellariae; papulae compound; not
fissiparous; gonads opening just above superomarginals at a distance from inter-
brachial angle; straight pedicellariae small, compressed, lanceolate; no adambula-
eral spine pedicellariae.
1. APHELASTERIAS JAPONICA (Bell).
Asterias japonica Bet, Proc. Zool. Soc. London, 1881, p. 515, pl. 48, figs. 6, 6a-6b.— Dévrn-
LEIN, Zool. Anz. vol. 25, 1902, p. 335. (Asterias japonica Stimpson is a nomen nudum.)
Asterias torquata Suapen, Challenger Asteroidea, 1889, p. 570, pl. 102, figs. 1-4.

I have examined specimens from the following Japanese localities: Otaru
(Hokkaido), Same (Rikuoku), Ayukawa, Tsuruga (Echizen). A single specimen
from Albatross station 4808, Tsuruga Strait, 47 fathoms, sand, shells, coarse gravel,
is referable to Sladen’s torquata which deserves recognition as a deep water forma of
japonica. (See pl. 72, fig. 4.)

—~ forma TORQUATA Sladen.
Asterias torquata SuapeN, 1889, p. 570, pl. 102, figs. 1-4.

Off Yokohama; Tsuruga Strait, 5 to 47 fathoins.

Genus ASTERIAS Linnaeus, p. 5
1. ASTERIAS AMURENSIS Litken, p. 6.
2 ASTERIAS AMURENSIS ROLLESTONI Bell.
Asterias rollestoni Beuu, Proc. Zool. Soc. 1881.—Dépearier, 1902, p. 33.
Asterias amurensis Suapen 1889, p. 575.
Allasterias forficulosa Vennritx, 1914, p. 194, pl. 83, figs. 3~3e; pl. 84, fig. 1; text figs. 10, 11.

Warmer waters of Japan.

3. ASTERIAS AMURENSIS MIGRATA (Sladen).

Asteracanthion rubens (Linnaeus) var migrafum Staves, Journ. Linn. Soc. vol. 14, 1879,
p. 432,

Allasterias migrata Verniut, 1914, p. 373.

Korean Strait. The two types which I have examined are very immature, R
being 16 mm. in the larger. They are possibly the young of versicolor. The skin is
mottled dark and light brown, as in that species.

4. ASTERIAS FORBES! (Desor).
Aateracanthion forbesi Drson, Proc. Boston Soc. Nat. Hist. vol. 3, 1848, p. 67.
Asterias arenicola Stimpson, Proc. Boston Soc. Nat. Hist., vol. 8, 1861, p. 268.
Asteracanthion berylinws A. Acassiz, Proc. Amer. Acad. Arts Sci., 1863.
Asterias forbesii Vernitt, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 345—C.arx,
Bull. U. 8S. Fish Comm. for 1902, p. 552, pl. 1, figs. 1, 2; pl. 4, figs. 14, 15.
Aateracanthion novae boracensis Pennier, Pédicellaires, 1869, p. 41, pl. 1, fig. 9a,
Maine to Gulf of Mexico; low water to 27 fathoms.
5. ASTERIAS RUBENS Linnaeus.
Aaterias rubens Linnazvus, Syst. Nat. ed. 10, 1758, p. 661; ed. 12, 1766, p. 1099. For

synonymy: Perrier, 1875, p. 47 under rubens and violacea; Stapen, 1889, p. 572;
Bett, Catalogue, 1892, p. 100. Synonyms: Asterias clathrata Pennant, 1777; Amterics

206 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

glacialis PENNANT, 1777 (not Linnaeus); Asterias violacea O. F. MUiER, 1788; Asterias
holsatica Revzrus, 1805; Asterias minuta Retzius, 1805; Asteracanthium distichum
Branpt, 1851; Asterias murrayi Bevu. Various combinations of the above names
under Asteracanthion, Asteracanthium, Stellonia, Uraster.

From the White Sea and Iceland (but not Greenland) down to the Senegal
coast; exceedingly common al] around the British coasts. Accidentally in the Medi-
terranean at Cette. From upper tide level to about 400 meters; exceptionally as
much as 650 meters. (Mortensen, 1927.) -

6. ASTERIAS VERSICOLOR Sladen.
Asterias versicolor SLADEN, 1889, p. 573, pl. 104, figs. 1-4.

The types were taken in depths of 8 to 50 fathoms off Kobe and Awadji Sima,
Japan. I have examined the specimens from Challenger station 233, off Kobe. The
species is obviously related to amurensis, but appears to be perfectly distinct from
both amurensis and rollestoni. It will not be strange however if versicolor is found to
intergrade with rollestoni. The general color is yellowish mottled with brown. Each
abactinal spine is the center of a yellowish zone. The crossed pedicellariae and the
straight pedicellariae are very slender. Sladen’s figures are excellent.

7. ASTERIAS VULGARIS Verrill.

Asterias vulgaris VeRRILL, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 347.—Crark, U.S.
Fish Comm. Bull. for 1902, p. 533, pl. 1, figs. 3, 4; pl. 4, figs. 16, 17.

Asterias stimpsoni VERRILL (part), Proc. Boston Soc. Nat. Hist., vol. 10, 1866.

Asterias rubens Gouup, 1841; Asteracanthion rubens Dusor, 1848; Asteracanthion violaceus
Srimpson, 1853; Asteracanthion rubens Stimpson, 1853; Asteracanthion pallidus A.
Aaassiz, 1863 (nomen nudum); Asterias vulgaris Packarp, 1863 (nomen nudum);
Asterias fabricti PERRIER 1875, p. 56; Asterias pallida Goto, 1898.

In shallow water, this species ranges from Labrador to the eastern part of Long
Island Sound, while in deep water it continues on to the neighborhood of Cape Hat-
teras. Low tide to 358 fathoms, exceptionally to 633 fathoms (lat. 39° 15’ N., long.

y

72° W., Arcturus expedition).
Genus EVASTERIAS Verrill, p. 139

. EVASTERIAS ECHINOSOMA Fisher, p. 152.
2. EVASTERIAS TROSCHELII (Stimpson), p. 139.
—— forma TROSCHELH, p. 141.

—— forma ALVEOLATA Verrill, p. 144.

—— forma ACANTHOSTOMA Verrill, p. 148.

Genus GASTRASTER Perrier

-~

Gastraster Perrier, Expéd. sci. Travailleur et Talisman, 1894, p. 103. Type, Pedicellaster
margaritaceus PeRRIeR, 1882.—FisuEr, 1923, p. 596—MorvTEnseEn, 1927, p. 137.

Diagnosis.—Abactinal plates relatively large, imbricated in three very regular
longiseries; abactinal and superomarginal spinelets small, spaced, delicate; infero-
marginal plates with an oblique series of three or four very flat, bladelike spines which
form a prominent fringe along edge of ray; superomarginals broader than inferomar-
ginals and abactinal plates, strongly imbricated, four lobed, bearing a crescent of
small spinelets; actinal plates small, one corresponding to each inferomarginal;
adambulacrals diplacanthid, the spines without attached pedicellariae; first adam-
bulacrals not in contact behind mouth plates; the latter flare a little at inner ends

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 207

and carry two suborals and a tiny lateral marginal spinelet; a few scattered very
small crossed pedicellariae; and a few very small broadly lanceolate straight pedi-
cellariae on actinal surface; tube feet biserial proximally.

The type and only known species is an immature form, which, when adult, may
lose the primitive character of the skeleton especially of the mouth angle.

The relationships of the genus are obscure. Perrier placed it in the Pedicellas-
teridae while Mortensen believes it is nearer Stichaster. As it stands, it is as isolated
as Stichastrella and Neomorphaster.

GASTRASTER MARGARITACEUS (Perrier).
Pedicellaster margaritaceus Perrier, Rapport sur la Faune sous-marine de A. Milne Edwards,
1882, p. 50.
Gastraster RE Baty Pernier, 1894, p. 103, pl. 9, fig. 3—Mortensen, 1927, p. 137.
I have examined the type specimen, R 17.5 mm.
Off the Azores; Bay of Biscay 938 to 1,225 meters; taken also at 53° 07’ N., 14°

50’ W., 738 to 900 meters (Mortensen),
Genus LEPTASTERIAS Verrill, p. 23
Subgenus LepTrasTeRIAs Verrill, p. 24

1, LEPTASTERIAS ARCTICA (Murdoch), p. 24.
~—— forma ARCTICA, p. 26.
~~ forma BERINGENSIS, p. 27.

2. LEPTASTERIAS AUSTERA (Verrill).

Asterias austera Verritt, Amer. Journ. Sci., vol. 49, 1895, p. 209.

George’s Bank, off Cape Cod, 33 to 35 fathoms (Verrill).

1. LEPTASTERIAS FISHERI Djakonov, p. 42.

4. LEPTASTERIAS GROENLANDICA (Liitken), p. 45.
~~ forma GROENLANDICA, p. &.
~~~ forma CRIBRARIA, p. 47,

5. LEPTASTERIAS HISPIDELLA Verrill.
Leptasterias hispidella Verniui, Amer. Journ. Sci., vol. 49, 1895, p. 210.
Albatross Station 2494, 45° 14’ 30’’ N., 59° 06’ 45’’ W. (off Nova Scotia), 50
fathoms.
“Allied to ZL. littoralis but has much longer and very acute spines, which are
less numerous.”’ (Verrill.)

6. LEPTASTERIAS HYLODES Fisher, p. 35.
7. LEPTASTERIAS HYPERBOREA Danielssen and Koren.

Asterias hyperborea DANIELSSEN AND Koren, Nyt Mag. for Naturvidensk, vol. 27, 1882,
p. 269; Norwegian North Atlantic Exp., 1884, p. 10, pl. 3, figs. 1-7.

Asterias normani DANIELSSEN AND Koren, Nyt Mag. for Naturvidensk, vol. 28, 1883, p. 1,
pls. 1, 2, figs. 1-9.

? Asterias milleri var. floccosa LevinseN, Kara-Havets Echinodermata, 1886.

Vicinity of Spitzbergen; to 35 fathoms (see von Hofsten, 1915, p. 63); Baffin
Bay, 98 fathoms (see p. 34). Exclusively high arctic.

§. LEPTASTERIAS LEPTALEA Verrill, p. 44.
9. LEPTASTERIAS LITTORALIS (Stimpson).
Asteracanthion littoralis Stimpson, Invert. Grand Manan, 1853, p. 14. (Grand Manan,
Bay of Fundy.)
Asterias littoralis Vernitt, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 349.

208 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Leptasterias littoralis VerR1L1, Prelim. Check List of the Marine Invertebrata of the Atlantie
Coast, from Cape Cod to the Gulf of St. Lawrence. New Haven, 1879, p. 14.

& Casco Bay, Me., to Gulf of St. Lawrence; low tide to 23 fathoms. Alhed to
miilleri.
10. LEPTASTERIAS MULLERI (Sars).

Asterias hispida Pennant, British Zoology, vol 4, 1777, pl. 30, fig. 58 (according to Morten-
sen).

Stellonia hispida Forspes, Mem. Wern. Soc., vol. 8, 1833, p. 123.

Uraster hispida Forses, British Starfishes, 1841, p. 95.

Asteracanthion miilleri Sars, Archiv f. Naturges., vol. 10, 1844, p. 169; Fauna litt. Norveg ,
1 Heft, 1846, p. 56, pl. 8, figs. 38-43.

Asterias miilleri Norman, Ann. Mag. Nat. Hist., ser. 3, vol. 15, 1865, p. 127. For literature
see Ludwig, Fauna Artica, vol. 1, 1900, p. 481.

Leptasterias miilleri Verritu, Proc. Boston Soc. Nat. Hist., vol. 10, 1866, p. 350.

North Sea, Irish Sea, southwest Ireland; Shetland Islands; Iceland; Scandinavian
coasts. The arctic records of miilleri (Spitzbergen, Greenland, Siberian Sea) probably
apply to hyperborea; while those of boreal waters from Maine northward, to L. tenera,
or allied forms.

11. LEPTASTERIAS OCHOTENSIS Brandt, p. 57.
12. LEPTASTERIAS OCHOTENSIS SIMILISPINIS (Clark.)

Asterias similispinis H. L. Cuarx, Bull. Mus. Comp. Zo6l., vol. 51, no. 11, 1908, p. 288
(Taraku Island, Hokkaido, Japan).

Boreal waters of Japan.

13. LEPTASTERIAS ORIENTALIS Djakonoy, p. 40.
14. LEPTASTERIAS TENERA (Stimpson).

Asterias tenera Stimpson, Proc. Boston Soe. Nat. Hist., vol. 8, 1862, p. 269.
Asterias compta Stimpson, Proc. Boston Soe. Nat. Hist., vol. 8, 1862, p. 270.
Asteracanthion flaccida AGAssiz, 1863.

Leptasterias compta VERRILL, Proc. Boston Soe. Nat. Hist., vol. 10, 1866, p. 350.
Leptasterias tenera VeRRILL, Amer. Journ. Sci., vol. 7, 1874, p. 504.

—— forma TENERA (Stimpson).

Cape Cod to Newfoundland, 10 to 129 fathoms.

—— forma COMPTA (Stimpson).

Atlantic coast of North America from north latitude 37° 19’ to 45° 29’; 10 to 100
fathoms. Large and abundant in the cold areas south of Rhode Island and Marthas
Vineyard, in 20 to 50 fathoms (Verrill).

Subgenus HEXASTERIAS Fisher, p. 59

15. LEPTASTERIAS AEQUALIS (Stimpson), p. 120.
—— forma AEQUALIS, p. 123.
—— forma NANA Verrill, p. 123.
16. LEPTASTERIAS ALASKENSIS (Verrill), p. 124.
—— forma ALASKENSIS, p. 125.
—— forma SHUMAGINENSIS Fisher, p. 128.
—— forma PRIBILOFENSIS Fisher, p. 129.
17. LEPTASTERIAS ALASKENSIS ASIATICA Fisher, p. 131.
18. LEPTASTERIAS ALASKENSIS MULTIPSINA Fisher, p. 133.
19. LEPTASTERIAS ALEUTICA Fisher, p. 101.
20. LEPTASTERIAS ASTEIRA Fisher, p. 103.
21. LEPTASTERIAS CAMTSCHATICA (Brandt), p. 91.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 209

22. LEPTASTERIAS CAMTSCHATICA DISPAR Verrill, p. 94.
~~ forma NITIDA Fisher, p. 95.
~~ forma NESIOTIS Pisher, p. %.
~~ forma DISP AR, p. 97.
2. LEPTASTERIAS COE Verrill, p. s4.
MM. LEPTASTERIAS CORI TRUCULENTA Fisher, p. 6.
25. LEPTASTERIAS HEXACTIS (Stimpson), p. 107.
—— forma HEXACTIS, p. 109.
~~ forma ASPERA Fisher, p. 110.
~—— forma REGULARIS Verrill, p. 111.
~~ forma SIDEREBA Verrill, p. 112.
—— forma PLENA Verrill, p. 113,
25. LEPTASTERIAS HEXACTIS VANCOUVERI (Perrier), p. 115.
27. LEPTASTERIAS LEPTODOMA Pisher, p. 105.
23. LEPTASTERIAS POLARIS (Miller and Troschel), p. 60.
—— forma POLARIS, p. 60.
——— forma SUBACERVATA Fisher, p. 60.
2. LEPTASTERIAS POLARIS BOREALIS (Perrier), p. 61.
30. LEPTASTERIAS POLARIS ACERVATA (Stimpson), p. 62.
~~ forma APHELONOTA Fisher, p. 63.
—— forma ACERVATA, p. 66.
~~ forma POLYTHELA (Verrill), p. 71.
31. LEPTASTERIAS POLARIS KATHERINAE (Gray), p. 77.
32. LEPTASTERIAS PUSILLA Fisher, p. 118.

Subgenus NESASTERIAS Fisher, p. 135
33. LEPTASTERIAS STOLACANTHA Fisher, p. 135.
Genus PISASTER Miiller and Troschel, p. 162

1. PISASTER BREVISPINUS (Stimpson), p. 180.
~—— forma BREVISPINUS, p. 181.
~~ forma PAUCISPINUS (Stimpson), p. 184.
2 PISASTER GIGANTEUS (Stimpson), p. 172.
3. PISASTER GIGANTEUS CAPITATUS (Stimpson), p. 177.
4. PISASTER OCHRACEUS (Brandt), p. 164.
—— forma CONFERTUS (Stimpson), p. 168.
~~ forma OCHRACEUS, p. 165.
~~ forma NODIFERUS Verrill, p. 169.
5. PISASTER OCHRACEUS SEGNIS Fisher, p. 171.

Genus STENASTERIAS Verrill, p. 138.
1. STENASTERIAS MACROPORA (Verrill), p. 138,
Genus STEPHANASTERIAS Verrill, p. 156

1, STEPHANASTERIAS ALBULA (Stimpson), p. 157.
2. STEPHANASTERIAS GRACILIS (Perricr).

Asterias gracilis Perrier, Bull. Mus. Comp. Zodl., vol. 9, no, 1, 1881, p. 4; Nouv. Arch. du
Mus. d’Hist. Nat., vol. 6, 1884, p. 204.
Stephanasterias gracilis Verrivy, Trans. Conn. Acad., vol. 10, 1899, p. 223; West Indian
Starfishes, 1915, p. 25, pl. 9, figs. 2-2c.
Lesser Antilles, Gulf of Mexico off Florida and off Havana; off east coast of
Florida; 56 to 200 fathoms.

Genus STICHASTRELLA Verrill
Stichastrella Vernitu, Shallow-water Starfishes, 1914, p. 40 —Fisner, 1923, p. 597. Type,
Asterias rosea O. F. Miller.
Diagnosis.—Plates small, with numerous, small, granuliform, close-set spinelets;
general appearance suggesting Henricia; abactinal skeleton close-knit with small
papular areas, rather irregularly distributed dorsolaterally but in definite marginal

210 BULLETIN 76, UNITED STATES NATITONAL MUSEUM

and actinal longiseries; actinal plates in two or three series; carinals regular; supero-
marginals slightly broader than inferomarginals; crossed pedicellariae surrounding
papular areas, not in circlets around spinelets; adambulacral plates with transverse
series of three or four spines proximally, usually three distally, all devoid of attached
pedicellariae.
1. STICHASTRELLA ROSEA (O. F. Miiller).
Asterias rosea O. F. Mistupr, Zool. Dan. Prod. 1776, p. 234.
Stichaster roseus M. Sars, Oversigt af Norges Echinodermer, 1861, p. 86.
Stichaster roseus var. ambiguus Farran, Fisheries Ireland; Sci. Invest., 1912, no 6 (1913),
p. 12, pl. 2, fig. 3.
Stichastrella rosea VERRILL, 1914, p. 40.
Bay of Biscay to Lofoten on the Norwegian coasts; British Isles; 4 to 430 meters
(Mortensen).
—— forma AMBIGUA (Farran).
Off southwest Ireland, 400 to 1,300 meters.

Genus TARSASTROCLES Fisher

Tarsastrocles FisHpr, Ann. Mag. Nat. Hist., ser. 9, vol. 12, 1923, pp. 597, 605. Type,
Hydrasterias verrilli Fisher.

Diagnosis.—See key, p. 5.
1. TARSASTROCLES VERRILLI (Fisher).
Hydrasterias verrilli Fisher, U. 8S. Fish Comm. Bull. for 1908, pt. 3, 1906, p. 1106, pl. 41,
figs. 3, 3a, 3b.

Hawaiian Islands, 284 to 290 fathoms.

Subfamily NOTASTERIINAE Fisher
Genus ICASTERIAS Fisher

Icasterias Fisoer, Ann. Mag. Nat. Hist., ser. 9, vol. 12, p. 601. Type, Asterias panopla
Stuxberg.

Diagnosis.—Abactinal skeleton very weak, irregularly reticulate, sometimes
degenerated into more or less disconnected plates; a definite series of carinal plates
bearing acicular spines; a few dorsolateral isolated spines; marginals smaller than in
Urasterias; superomarginals monacanthid; inferomarginals diplacanthid or sporadi-
cally monacanthid; alongiseries of spineless oblong intermarginal ossicles; adambulacrals
diplacanthid; no actinals; crossed pedicellariae smaller than in Urasterias, with rela-
tively stouter jaws, scattered thickly over the abactinal and lateral surfaces, but not
in wreaths or clusters on the abactinal, marginal, or adambulacral spines; gonads
opening just above superomarginal plates a short distance from base of ray.

1. ICASTERIAS PANOPLA (Stuxberg).

Asterias panopla Stuxsere, Ofversigt af Kungl. Vetensk Akad. Férhandl., 35te Argang,
no. 3, 1878, p. 32.—Danietssen and Koren, 1884, p. 17, pl. 5.—von Horsten, 1915,
p. 68.

Asterias panopla vars. brevimana, gracilis, inermis KALISCHEWSKY, 1907, p. 46.
Arctic; both coasts of Greenland, Spitzbergen, Barents Sea, Kara Sea, east-
ward to longitude 114° 35’ E.; 18 to 560 meters (von Hofsten).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 211
Genus URASTERIAS Verrill

Crasterias Vernitt Amer. Journ. Sci., vol. 28, 1909, p. 67; 1914, pp. 51,187. Type, Aster-
acanthion linckii Miller and Troschel. Fisher, 1923, pp. 249, 599, 600.

Diagnosis. —Large Notasteriinae with a weak abactinal skeleton consisting of
small lobed plates united by slender ossicles into an irregular, loose, open reticulum;
marginal plates conspicuously larger than dorsolateral, the superomarginals often
trilobate proximally with a long descending process; carinal and dorsolateral spines
acicular, isolated, usually without but sometimes with wreaths of crossed pedicellariae;
crossed pedicellariae very large and of an unusual form, with slender serrate jaws
terminating in an unexpanded unguiculate tip; marginal spines prominent, acicular,
one to a plate, the inferomarginals with a very large cluster or cushion and the supero-
marginals with a wreath of crossed pedicellariae; very numerous, large, compressed-
ovoid, straight pedicellariae; gonads opening just above superomarginal plates in
interbrachial angle; actinals absent (or extremely rudimentary); adambulacral plates
predominantly diplacanthid, the spines normally devoid of attached pedicellariae,
although these, on peduncles attached to plates, are numerous on furrow margin.

1, URASTERIAS LINCKT (Miller and Troschel).

Asteracanthion linckii Mtituer and Troscuer, 1842, p. 18.

Asteracanthion stellionura Perrier, 1869, p. 48, pl. 1, fig. 10.

Asterias gunneri DANreELSSEN and Koren, Nyt Mag. Naturvid., vol. 27, p. 268; 1884, p. 7,
pl. 2; pl. 3, figs, 8, 9.

Asterias lincki var. robusta et var. sibirica KAuiscnewsky, 1907, p. 51.

Urasterias linckii Verrity, 1909, p. 67; 1914, p. 52.

Spitzbergen; Greenland; Nova Scotia; Arctic Ocean from Norway to New Siberia

Islands; 5 to 280 meters (von Hofsten, 1915, p. 66).
2 URASTERIAS ENOPLA (Vervill).
Asterias enopla Verritt, Distribution of the Echinoderms of Northeastern America, Amer.
Journ. Sci., vol. 49, 1895, p. 208.
Off Nova Scotia, 53 to 100 fathoms.
Possibly a young linckii.

Subfamily NEOMORPHASTERINAE Fisher
Genus NEOMORPHASTER Sladen

Neomorphaster Sutapen, Challenger Asteroidea, 1889, p. 436. Type, N. eustichus Sladen.—
Fisner, 1923, p. 596.

Calycaster Pennier, Comptes-rend., 1891, p. 1226; Mem. Soc. Zool., vol. 4, 1891, p. 258.
Type, C. monoecus Perrier.

Diagnosis.—Abactinally resembling Zoroaster; primary plates of disk forming a
regular rosette; abactinals and marginals sparsely granulated, imbricated in seven
very regular longiseries (and in regular transeries as well), with large papulae at the
corners, also in longiseries; two or three series of actinal plates each carrying two or
three short, somewhat flattened spinelets; adambulacrals diplacanthid, the spines
without attached pedicellariae; adoral carina composed of three or four pairs of postoral
adambulacrals; tube feet quadriserial proximally, biserial at end of ray; small crossed
pedicellariae scattered on sides and abactinal surface; one or two straight pedicel-
lariae on inner edge of every second adambulacral.

212 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

1. NEOMORPHASTER TALISMANI Perrier.
Stichaster talismani Perrine, Ann. sci. nat., vol. 19, art. 8, 1885, fig. 22.
Neomorphaster euslichus SLADEN, 1889, p. 438, pl. 66, figs, 3, 4; pl. 68, figs. 9, 10.
Calycaster monoecus Perrier, Mem. soe. zool. France, vol. 4, 1891, p. 262; Rés campag. sci.
Prince de Monaco, fase. 11, 1896, p. 28, pl. 2, figs. 2, 2a; pl. 3, figs. 3, 3a.
Bay of Biscay; Azores; off the coast of Morocco; southwest of Ireland; 400 to
2,000 meters, exceptionally to 5,413 meters. (Mortensen, 1927.)

2. NEOMORPHASTER FORCIPATUS Verrill.
Neomorphaster forcipatus Verriuu, Proc. U. 8. Nat. Mus., vol. 17, 1894, p. 269; Amer.
Journ. Sci., vol. 49, 1895, p. 206.

Off Georges Bank and south of Marthas Vineyard, 852 to 990 fathoms.

I have examined No. 11131, U.S.N.M., listed by Verrill as one of his types.
Verrill writes (1894, p. 270) that there are ‘‘attached to the adambulacral spines and
in the ventral interradial spaces many acute, ovate, forcipate [straight] pedicellariae,
often mixed with crossed pedicellariae and scarcely exceeding the latter in siz3.”
I find no pedicellariae whatsoever attached to the adambulacral spines which are two
to each plate. On the furrow face of alternate adambulacral plates are single small
straight pedicellariae or groups of two and three. I find no actinal interradial
straight pedicellariae but crossed ones larger than the lateral and dorsal. This
species appears to be a typical Neomorphaster.

The broad carinals are flanked by a single series of much narrower dorsolaterals;
these are followed by superomarginals as broad as the carinals; then a row of in-
feromarginals about as broad as the dorsolaterals; then a row of actinals, on the
side of ray proximally and forming the ventrolateral border distally. Between
these and the adambulacrals are two short rows of ventral actinals, the outer scarcely
reaching middle of ray, and each plate bearing three or two spines, as do the distal
plates of the lateral actinal series (which distally defines the actinolateral margin of
ray).

LIST OF SPECIES AND VARIETAL NAMES APPLIED TO ASTERIINAE, NOTASTE-
RIINAE, AND NEOMORPHASTERINAE OF THE NORTHERN HEMISPHERE

In the first column is given the species and original describer; in the second
column, the original genus and date; in the third column, the genus to which the
species has been assigned in the foregoing summary.

Valid species and forma names are in roman; synonyms in italics.

acanthostoma Verrill__=____________- Asteria; 1909). 2 ee ee Evasterias.
acervata Stimpson. .--......._.____- ASteTIas 8622-2 =o eee eee Leptasterias.
aequalis Stimpson___.--_._______-__- Asteriag; 186220 fou 45 steer Leptasterias.
Slaskensis \Werrils spe 8 = se Asterias. 9092s 5.2. a Ce eee Leptasterias.
albertensis) Vermll: <2 232 Se Paristerias Olt en ase eo eee Asterias.

Sl DUS StI psON ee ean eee eee Asteracanthion, 1853___._-..-__-- Stephanasterias.
alveclataiVerrlless he wes ore ie ates Hivastenias, 1 OkAwe ose as haere ee Evasterias.
ambiguus WarranUl fue usb. 2UL La Stichaster; 191ISEGS. Sao 22a ie Stichastrella.
amurensis Liitken-__-_--....-/----1. Asterias ul Site: tex hee taehpcl ast Asterias.
anomalanViermls 92) 5 bo ele =o Allasterigs 1900E foi ted ee Asterias.

arctican Murdoch zens seea- = 2 Se Apterias, 88beew se eey oa oe ae Leptasterias.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 213

arcticus Danielssen and Koren... .. ~~. Stichaster, 1884.

arenicola Stimpson... ........------- Asterias, 1862............-- _. Asterias.
MONTE VERS oe seine d oe aanwew cunt ANU, SONU 26. pwn acae ane oe _ Leptasterias.
MEIUMNAVOis sc ougucch ws nsaccnns iy oe ... Pisaster.
LL + sh nS Sa Sem a ae Asteracanthion, 1863............. Asterias.
borealis Permtersc.-. 2.25.2 <55.5. =~. = Avior, 10(0-..-»«2=-+->- _. Leptasterias.
Wackiala Perrier... ..----.---+.- Asterias, 1875................... Evasterias.
brevimana Kalischewsky.......----..- PRUs, 100) beh «tenuesnosans . Teasterias.
brevispina Stimpson..........-.-..-. REMMI NOT esac oo na canoe . Pisaster.
camtschatica Brandt... -........-.-- AMMEN, 1600... 6. ce ae -po a= Leptasterias.
capitata Stimpson... ...............- Asterias, 1862_._................ Pisaster.
SNISEUGRN Ga oy cna tawo owen en 5 Leptasterias, 1914_._....._._.... Leptasterias.
PRAEe ROODENG. <.-.us2-5ccolce. Asterias, 1777... ..- eee Sere. Asterias.
ENTREE swine wes Crete eo Leptasterias, 1914_ _ Leptasterias.
ere, WORPUe ee CL SoS - cot Leptasterias, 1914. ...........--. Leptasterias.
Sununu peon os. oe oo) oo. Asterias, 1866_._....-..-- . Leptasterias.
Oo OR el ES ee ae Leptasterias, 1914... _. Leptasterias.
conferta Stimpson-_-.---.-....------ Asterias, 1862_......._-- _.. Pisaster.
cribraria Stimpeon-.2-..--.-.---.---- Asterias, 1862...........-- _.. Leptasterias.
Me: VERT ree eee OO eCE Evasterias, 1914_......_._....___._. Evasterias.
[OS OD epg nape RS EE Pas at ae Peptasterias, 1918. -2 22-2 -22<-2.- Leptasterias.
quemcwee bimaie. 2.) ose on oe. =e. Asteracanthium, 1851__...._._.._ Asterias.
MORI EERE aa tie en ph nae an SERRE TES ORD eect rere Siete ae ee Leptasterias.
MUOL TORU ane eto e sank cea oe BOTA LWP eee a ooe phantom.
echinosoma Fisher..........-.------- Evasterias, 1926...-.-.------...- Evasterias.
CN VOPR et ee ore ee eae RROOS, LOM es bee cee ee ean Urasterias.
BIOnOre ALOU Gss onan canons nemo oa Asterias, 1835.

exquisita deLoriol. -.--......----.--- RAE TOCA A rose hcalce ott are ares Pisaster.
TUES EMERY art sce eee ee Neomorphaster, 1889... _....._~. Neomorphaster.
tie POMIer ons eons aan aonon = ROR SER ton et wean ae a Asterias.
PTL REIT ott ete ee on Leptasterias, 1929. =. --:-22---:-- Leptasterias.
fissispina Stimpson _...-.....----.---- PEEL 2 time pail apatnnn fea Pisaster.
“flaccida A. Agnssiz_._.....-:....-..-- Asteracanthion, 1863__...._.._--- Leptasterias.
PCCUNE SVM. wag ons caen ea UES SOC tt on eee _. Leptasterias.
RD ORES. < oc neni oases nanae Asteracanthion, 1848............. Asterias.
WCNINSRIE WER Rli vee oc ee aad aa Neomorphaster, 1894__-__.-_._.. Neomorphaster.
yoracmioeG Vexrit..-.-.------------5- Allasterias, 1914._..._._........-- Asterias.
glacialis Pennant__- ~~... ene een tea Asterias, 1777. -.--_-.- eeatmee> Asterias.
gigantea Stimpson. __...........-.-- Asterias, 1862....._.._. eee Pisaster.
MERE BOR etl © neg on oo ad oe BEORIEE, IOGk ene ee nace a Stephanasterias.
gracilis Kalischewsky _.-..--.......-- Ty Riad Rs pf ie poh ee Teasterias.
DOV OUT ohana o¢aanaacascnasens PIANO: LU tie tree eae ee ee phantom species.
groenlandicus Littken_.......-.-.-..- Asteracanthion, 1857- ~~ _._..-..- Leptasterias.
gunneri Danielssen and Koren. -.__-.. BNGMES, £004 - cee wckes ses Urasterias.
hexactis Stimpson... .....-....---.- Baan, 1O0ie Ea eens stone. Leptasterias.
PRR ONTO Sel Sa Go de arte es co oor BEEN, ASE eee eee wane Leptasterias.
DDIGGUS WOOGIE. chpeuscacnecscucsese Leptasterias, 1895... _........_.. Leptasterias.
pT el el Be NEL 1 arama aul dp maps al da autre Asterias.
hyperborea Danielssen and Koren.-- - . Asterias, 1882... ...... ...--.... Leptasterias.
ene SER ee a Cees rest. cao o Leptasterias, 1914............ Evasterias.
De ee eer neen tana ARO, 1OBlonancnen anh caccn «a Leptasterias.
inermis Kalischewsky.........-..- ... Asterins, 1907......_..._.._..... Teastorias.
(en Ave tee nccot t+. 6 SRM ROOia ton. nssacenses>e =~ Leptasterias.
TORLMAE DRRUUs « ou ae nee neo oes 2 SG kode ens ane oe . Pisaster.

214 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

japonica Bell. --~-------------------- PAB TOLISS ul 9 lene ee ere Aphelasterias.
katherinae! Graye-- -=-- noe see = IASterias elo 40 ses ee Leptasterias.
leptalea Verse eee Leptasterias, 1914___------------ Leptasterias.
linckii Miller and Troschel----------- Asteracanthion, 1842___--------- Urasterias.
littoralis Stimpsonessoe- ee =e Asteracanthion, 1853 ------------ Leptasterias.
longimana Kalischewsky - ------------ / Asteriassl OOo ere tae en Leptasterias.
hitkents SumpsonDe ase es == —— = — Agtering SG ea ee Pisaster.
macount Verrier ae ‘Leptasterias, 19452 -> = "a> = ==== Evasterias.
macropors) Verna =se= == =e Astenas, 1009 es= === a= Stenasterias.
margaritaceus Perrier---------------- Pedicellaster, 1882_-------------- Gastraster.
margaritifer Miller and Troschel--_-_-- Asteracanthion, 1842_----------- Pisaster.
merriamt Verrile o-.- 525 == 52 --==5---— Orthasterias 191 4e. === — == ee Leptasterias?.
mexicanum Liitken-_---------------- Asteracanthion, 1859- -- --------- Leptasterias.
migratim Siadenes= sees see Asteracanthion, 1879_.----------- Asterias.
miliaris, Vermlle = eee ee eptasterias; 1914. === = eee Leptasterias.
MUnUla KR ADMICLUBS == Soe eek a= = aa = IABteriaswliiQ0 see == oe =a eae Leptasterias.
minuta Retzius] 2-92 --- esas Asteria: 1805: -<22- 25/2 22-25 See Asterias.
monoecus Perrier—-=--=.-.---=-_-_-_= @aly.caster, 118915 =-—= === ee Neomorphaster.
miUllert Sarge ees Asteracantnigne === == === ers Leptasterias.
ATUIELC LAU) VICI See Asterias. 10142 2= 27-22 Seen Leptasterias.
MUTTGY) BON Ran ee eS See ee Agteriass =2-) Sete ee Asterias.
mana, Vermrlle = ep te ee Teptasterias’ 1914s =*_ Sess n eee Leptasterias.
naulanums Beller at aoe eee eS Asteriasy(pr24)=--—-2- == ae Leptasterias.
NididGaV erm te ea a sas Stichaster, 1 86Gs5——55—=— == nee Stephanasterias.
nodiferus Wermils se eee Pisaster; 1904. == See ee Pisaster.
normani Danielssen and Koren----~--- Aisteriag « V88oo-2 58) 25) aeo eee Leptasterias.
MOTLONETISTS OV OUI ee ee es Adlasterias, 1909s == ene aee ae Asterias.
novaeboracensis Perrier_-------------- Asteracanthion, 1869___---------- Asterias.
oblectaVierile=— = eee eee ae ee Jueptasterias, 19142= esse 2aseees Leptasterias.
Ochotense; Brandt==-=- == =5 oe = Asteracanthium, 1851____-------- Leptasterias.
ochracea) Brandt=- = san. on ee Asterias, 1830-052 -==--2 Se ae Pisaster.
orientalis Djakonov_---..---.------- Leptasterias, 1929_____--_----___ Lepatsterias.
pallidus A Agassiz. 22 -6--4-- <2 cee Asteracanthion, 1863------------ Asterias.
PANnOplA Sts bere S ae ae eee Asteniaswil $7822 =e = eee Teasterias.
PUpULOSarV Cre ae ses ee ee Asterias (Pisaster), 1909___------- Pisaster.
DOLLS DING AV CLT ae eee eee ee By sis berias yO Ame ess ee eee Evasterias.
paucispina Stimpson...-.-....----__- Asterias18625-=2 >= 22 2c wee Pisaster.
PeClin ata BTaNO Ges sae ee ee eee Asteriass S30 eee = eee eee Asterias.
pith Dp Dit DEM a —eee ee o he Asterias, | S8ileeee mc seas eae Leptasterias.
plenatVienn ee een ee eee Leptasterias, 1914______--__--_-- Leptasterias.
polaris Miiller and Troschel__________ Asteracanthion, 1842_________-_-- Leptasterias.
poly thelayVernU === == see eee Seen PAS terias yi Ql ea eee eee Leptasterias.
problema Steenstrup----.-----___-__. Asteracanthion, 1857____-_------ Stephanasterias.
pugetana Verrill_---_-______________. Iheptasteriasy 1OlAs= = ee Leptasterias.
pycoopodiawitishers=---2 5-2. 22-5 eo Aphanasterias, 1923___________-- Aphanasterias.
rathbuni Werner: 2: = sees ee oe Allastering i009 s es = ae = aoe eee Asterias.
TERULATIS AV Gn eee ee ne Leptasterias, 1914______________- Leptasterias.
robusta Kalischewsky ---_._--__-____- Asterias) 19072: eee eee Urasterias.
rollestonisBellie ono Asterias, 188s 3%s== 2) see ae Asterias.
MOSCA eres ee ee ARterias: l7jOc essen ae ee ee Stichastrella.
rubens Linnaeus---_---------____--- Acterins, L758 topo en ee ee Asterias.
rudis Verrill______ s =o == === ------- Hvasterias, 10] 45:20 ee ewe ole Evasterias.
saanichensis deLoriol----__-___.__-_- Asterias 1857 eee ees Evasterias.

satsumana Déderlein________________ Asterias, 1902

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 215

segnie Fisher.< G4 $2 soos wuss dat Pidasten, 1086 soi 65 st canny ~tive Pisaster.
sibirica Kalischewsky ........-.-....- Asteries; 1007 6.d< uns 46 si pseu Urasterias.
ORTON ON TN LA a ecg lt es fl ws Leptasterias, 1914............... Leptasterias.
similispinis Clark... ..........-...-- Aeeting, THOR iss a en code Leptasterias,
O08 BOY. din ncoakh oc cindadeutn Asterias, 1825 (part).........-..- Asterias.
apitzbergensis Danielssen and Koren... Asterias, 1I884_.................. Leptasterias,
stallionura Perrier. .<......-....-..<.. Asteracanthion, 1869... _._..... Urasterias.
SUR MNMIINR WEIR Gi oi arm fee ree ge By Ra in elegant in Asterias.
euonodoed Versi. 2 Bone Oe ee Eveaterian, 19182) 84 ast Evasterias
subnodulosa Verrill.............--.-- Leptasterias, 1914..............- Evasterias.
subregularis Verrill._._........-....- Leptasterias, 1914.._._._..._-.-- Leptasterias.
PAUOST) POTION. o ch wqeans mene Stichaster, 1885................. Neomorphaster.
WeneeR HIDDNON. ooo. 5 oe denne ou ABTA GOL b= ode os aes Uawa ce Leptasterias.
MGUN PURUON = sagen deem onan nnen = RAUEEIREO. cote nd wad ane Aphelasterias.
troschelii Stimpson----........-.---- Amerian, S01 22b 33.2% 25 ete 2 Evasterias.
vanoouveri Perrier. ...............-. Antériag JB76 2s. 3r. sis aes. 2: Leptasterias.
SWURRETILE BIGDOG? ooo os der dab etsbrces ita ass Hydrasterias, 1906.._..........-. Tarsastrocles.
WoImicOlOr PLsGeN Aaterian, 1880 52 oo. do os aw bce Asterias
WINONA VOITULS once cman cacdcana PAAR LRT sooo fiery osc ge Evasterias.
violacea O. F. Miller... -.......-.--- AMUBTIEN NE POG ina Bos ot hos ote an Asterias.

WULmeEee Venti oo se ea he Adterias; T80bae06e. 2022 Sal. od Asterias.

216 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

SUMMARY OF THE GENERA AND SPECIES OF ASTERIINAE AND
NOTASTERIINAE OF THE SOUTHERN HEMISPHERE

The following synopsis is not intended to be a revision in the usual sense, since
I have not had sufficient material for such an undertaking. It is an attempt to
set in order the species of Asteriinae known to occur in the Southern Hemisphere,
particularly in antarctic and subantarctic regions. I have also essayed to define
more sharply a number of genera which are still poorly understood.

Diplasterias Perrier has been resuscitated for Asterias brandti and near rela-
tives. There seems to be no alternative under existing rules of nomenclature but
to recognize this name and to discard Podasterias which was first used by Perrier
for a Cosmasterias, then for Diplasterias; later by Koehler (1912) for Diplasterias
spinosa; still later by Verrill (1914) for Diplasterias (part).

So far as known no genus of southern Asteriinae and Notasteriinae is repre-
sented (at least well authenticated) in the Northern Hemisphere.

The following species have not been assigned a place in the summary.

Cosmasterias radiata Koehler, 1923, p. 36, pl. 2, fig. 5; pl. 3, figs. 1, 2. This
species does not seem to me to be congeneric with Cosmasterias lurida. It was dredged
from Shag Rock Bank between south Georgia and Falkland Islands, 160 meters.

Diplasterias spinosa Perrier, 1891, p. 82. Koehler, 1908, p. 273; 1912, p. 17,
pl. 1, figs. 2, 9; 1917, p. 29. This species was assigned to Podasterias by Koehler
in 1912 as sole species, and in 1917, with hesitation, to Cosmasterias. Koehler
indicates that there are three longiseries of actinal spines, but an examination of
his photograph of the actinal surface in conjunction with Perrier’s description sug-
gests rather that the inner series is referable to the actinal plates, while the two
outer are on the inferomarginal plates—as in Diplasterias. This interpretation and
the absence of definite longiserial arrangement of abactinal spines would rule out
Cosmasterias. The long narrow adoral carina is not found in Diplasterias, ss. The
species probably represents a new generic group, as Doctor Koehler has already
suggested. (1917, p. 29.) ;

The type and only known specimen was taken latitude 47° 29’ S., longitude
66° 45’ W.

Gastraster studeri de Loriol, 1904, p. 34, pl. 4, figs. 3, 3a-3f. Verrill, 1914, p. 360.
The type is probably a very immature Cosmasterias lurida and was taken along
with adult lurida at Port San Antonio, Gulf of San Mathias, Argentina. It is
certainly not congeneric with the type of Gastraster, G. margaritaceus, which I have
examined at the Museum d’Histoire Naturelle.

Asterias borbonica Perrier, 1875, p. 61. The type in the Museum d’Histoire
Naturelle carries a label ‘‘Parasterias borbonica—I Bourbon, 1852.” It is now
in alcohol but was dried at one time. R.17mm.,r5 mm. The specimen probably
represents a young Sporasterias, but no examination of the gonads was made.

Reunion Island, Indian Ocean, latitude 21° S.

Asterias inermis Bell, Proc. Zool. Soc., 1881, p. 512, pl. 47, figs. 2, 2a. The
type, supposed to be from Ecuador, is in reality Leptasterias groenlandica forma
cribraria, probably from Bering Sea. (See Fisher, 1926, p. 198.)

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—-FISHER 217

Asterias nautarum Bell. Listed by Sladen, 1889, p. 824. The type specimen
labeled ‘Ecuador, Hassler Collection,” is Leptasterias arctica (Murdoch) and is
probably from Bering Sea. (See Fisher, 1926, p. 198, and antea, p. 24.)

Asterias philippii Bell, Proc. Zool. Soc., 1881, p. 511, pl. 47, figs. 1, la. The
two types in the British Museum are labeled “South America, Hassler Coll.” These
specimens are Leptasterias polaris acervata (Stimpson), with a well-developed central
spine to each abactinal heap of enlarged granules. They are very nearly identical
with Stimpson’s type of Asterias acervata and undoubtedly came from Bering Sea.

Asterias fragilis Studer, Trans. Roy. Acad. Sci. Berlin, 1884; Farquhar, 1909,
p. 127. Farquhar gives a translation of the original description. The type meas-
ured R 16% mm., r 4 mm.; breadth of ray at base, 8 mm. Studer places it next
to Cosmasterias lurida. It is obviously young. Found east of New Zealand, 35°
21’ S., 175° 40’ E., 597 fathoms.

Asterias (Smilasterias) terwieli Emilie Goldschmidt, Ann. Mag. Nat. Hist.,
ser. 9, vol. 14, 1924, p. 499, text figs. 1 to 3. This species does not belong to the
Asteriidae.

Subfamily ASTERIINAE Verrill (emended)

KEY TO THE GENERA OF ASTERIINAE OF THE SOUTHERN NEMISTHERE

a'. Plates of the abactinal surface of ray have the appearance of forming rather definite longi-
tudinal series even when the dorsolaterals are not in regular series (in this case the dorso-
lateral region is narrow and dominated by the broad carinals and superomarginals);
abactinal plates generally with several spinelets or granules, but if with only one spine,
then this is small and not conspicuous; adambulacral plates never carry attached pedi-
cellariae singly or in clusters, although straight pedicellariae may occur on oral spines
and on furrow margin of adambulacral plates.

b'. Adambulacral plates with two spines.

c’. Actinal plates in one series (or rudiment of second) dominated by the inferomarginals
which form the ventrolateral margin of ray; no large unguiculate straight pedicella-
riae; crossed pedicellariae scattered, not in clusters of wreaths around abactinal spines;
superomarginals conspicuously broader than the inferomarginals.

d'. Carinals and superomarginals of exaggerated width in proportion to length; dorso-
lateral area broad, the plates in three to five series on each side of the carinals; one
madreporic body; not fissiparous; superomarginal plates without beaded area on
surface; (gonoducts open dorsally, those of male with a proximal swelling; inter-
brachial septa heavily calcified. Includes Coelasterias australis Verrill, 11 rays).

Stichaster Miller and Troschel.

@. Carinals and superomarginals broader than other plates but not excessively so; dorso-
lateral area narrow, the plates in a single straight or zigzag series (altogether, three
series of abactinal plates); fissiparous; multiple oe bodies; superomarginal
pistes‘ with beaded surface.hs Jes iec fou. Socdetk . wate. 2 dt Allostichaster Verrill.

@. Young Diplasterias may lead here; see under a*.

@. Actinal plates in two or more prominent series forming the actinal or the actinal and actino-
lateral surfaces of ray (and not dominated or overhung by the inferomarginals).

d', Large unguiculate straight pedicellariae; actinostome sunken, the adoral carina long,
consisting of more than three pairs of contiguous postoral adambulacral plates.

Cosmasterias Sladen.

®. No large unguiculate pedicellariae; actinostome not sunken; adoral carina short, con-
sisting of one or two pairs of contiguous postoral adambulacral plates. Plates

large with coarse, often capitate, globose spines and tubercles; abactinal skeleton

with large, more or less irregular meshes in two dorsolateral series on cither side

of the irregular carinal series; superomarginals with one to several capitate, stout

218 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

spines; inferomarginals with one to three spines; one to three series of actinal plates,
usually two or three; one well defined though sometimes irregular series of dorso-
lateral plates; adambulacrals very regularly diplacanthid__-___-_- Uniophora Gray.

b?. Adambulacral plates monacanthid; dorsolaterals not very numerous, trilobed, in one or two
fairly regular series, more or less concealed by integument; superomarginals very broad;
inferomarginals (each bearing an oblique comb of two to four prominent spines) form a
sharp ventrolateral border to ray; an inconspicuous series of actinal plates; rays short
and thick; gonads) open yentrally--==—- ==>" =2-=_-=— === ee tee Calvasterias Perrier.

@. Abactinal skeleton an irregular net with meshes of various sizes, the plates being sometimes
closely but irregularly imbricated by their lobes; abactinal plates not in rather obvious
longitudinal series (but occasionally in more or less evident transverse series) ; carinal series
usually but not always distinguishable and frequently very irregular; body sometimes coy-
ered by a pulpy skin which obscures the plates and sometimes even the spinelets.

b!. Adambulacral plates with two, rarely three spines, which never carry attached pedicellariae,
although large and small straight pedicellariae may occur on the surface of the plates or
on fleshy peduncles attached to plates near base of spines; clusters of the latter in dried
specimens occasionally appear to spring from the spine-sheath, but in reality do not.

c!. Abactinal plates and spinelets normally developed; both series of marginal plates nor-
mally developed.

d', An actinolateral chevaux-de-frise of inferomarginal spines in transverse series of three or
four; abactinal spinelets numerous, small, close-set; superomarginal spines small,
not forming a well-differentiated, single or double longiseries; pedicellariae scat-
tered, not in circlets at base of spines, nor on sheaths surrounding spines; skin not
thick or puffy.

e’. Actinolateral spines devoid of attached pedicellariae; actinal spines in one series or

absent; gonads opening dorsalliy=.22==_— 23= S ee Smilasterias Sladen.
e. Actinolateral spines carrying pedicellariae; gonads opening ventrally; paedophoric; one
series (ofvactinall spines_/o2 22232 24. Set Se eee Neosmilaster Fisher.

d*, Superomarginal, inferomarginal, and actinal spines forming longiseries with circlets or
bouquets of pedicellariae; abactinal crossed pedicellariae associated with spines, in
cirelets or attached to the base.

e!, Spinelets slender, mobile, more or less sheathed with soft skin carrying crossed pedi-
cellariae; gonads opening ventrally; young brooded by adult; superomarginals
monacanthid or diplacanthid; inferomarginals monacanthid or diplacanthid; one
or two series of monacanthid actinal plates (exceptionally spineless); adambula-
crals:‘repulanliy; diplacanthid==-221-- ash Sas eee Diplasterias Perrier.

e. Spinelets coarse, tuberculate, sometimes very large, capitate or globose; skin not soft
or pulpy; gonads opening dorsally; actinals in one to three series__Uniophora Gray.

c. Either the superomarginal plates, or those of both series, not easily distinguishable; abacti-
nal skeleton well developed; inferomarginal spines not in close-set transverse series.

d'. Only the inferomarginals, of the two series, clearly distinguishable; to them are joined
small lateral transverse ares (containing the superomarginals) separated, consecu-
tively, by broad papular areas or skeletal spaces; abactinal skeleton an irregular
network of plates, without indication of a carinal series; abactinal and inferomar-
ginal spinelets slender, cylindrical, in small groups; two long adambulacral spines;
ambulacral furrows wide, with large 4-ranked tube feet; rays 9 or 10.

Saliasterias Koehler.

d. Marginal plates not superficially distinguishable; a close granulation more or less uni-
form, without serial arrangement, interspersed with scattered pedicellariae, overlies
relatively large three to five lobed plates in an irregular abactinal reticulum; mar-
ginals four lobed; an incomplete series of actinals; size small; gonads opening ven-
trally. (Includes Hemiasterias Verrill, type G. biseriatus Koehler.)

Granaster Perrier.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-—-FISHER 219

¢. Abactinal skeleton weak or else abortive; abactinal integument thick, papillose; no actinal
plates.

d'. Marginal and abactinal plates present; marginal plates well developed; the supero-
marginals with one spine wreathed with pedicellariae and having long descending
processes which form a broad, vertical, intermarginal channel; inferomarginals with
two webbed spines (each with a cluster of pedicellariae on outer face); abactinal
surface soft, rather irregularly and finely papillose; abactinal plates thin (with rudi-
mentary spinelets only) forming an irregular reticulum entirely concealed by the
skin; interbrachial septum strongly calcified; gonads opening ventrally, eggs huge;
good sized, broadly lanceolate, straight pedicellariae on furrow margin; two pairs of
postoral adambulacral plates contiguous; adambulacral plates regularly dipla-

NON IN are Sots a enn apn base eis ab nap a Goro we ad isn ceo Cryptasterias Verrill.
@, Neither marginal nor abactinal plates present, the skeleton reduced to the ambulacral
and adambulacral systems. (See sec. a#.)_..._____________-_- Adelasterias Verrill.

. Adambulacral plates with a single spine, devoid of attached pedicellariae.

c'. Abactinal skeleton more or less well developed, consisting of lobed and oblong plates united
into an open or fairly close meshwork, in which not very regular longiseries can some-
times be distinguished; abactinal spinelets fairly to very numerous; skin typically
not unusually thick pulpy and rugose (exceptions); actinal plates in one series (may
be absent in young).

d'. Abactinal and superomarginal spines surrounded by circlets of crossed pedicellariae;
inferomarginal and actinal spines cylindrical or clavate, the former, two or three to
a plate in an oblique series, with an associated group of crossed pedicellariae;
RDN 2 ees J ee uo es Stal Je essa San Sporasterias Perrier.
@, Pedicellariae rare and scattered, not in circlets around abactinal and superomarginal
spinelets; inferomarginal and actinal spines chisel shaped, devoid of pedicellariae,
closely coordinated into oblique transverse combs forming a chevaux-de-frise along the
actinolateral border of ray; gonads unknown_____..._._____- Eremasterias Fisher.
@. Skeleton hidden under a pulpy, sometimes very rugose pustulated skin; abactinal skeleton
irregularly reticulate, weak, with few spinelets; gonads opening ventrally.
d'. Skin very thick, complex, pustulated, the abactinal spines being surrounded by the

: same cauliflower structure; actinals spineless; rays eight or nine; adambulacrals
Giplstanthid at base of ray 20 0525. . 220 ee Koehleraster Fisher.

@, Skin thick but not having a complex cauliflower structure.
e'. Both series of marginals well developed._.................-_._-- Anasterias Perrier.
e. Superomarginals weakly developed, forming a rather irregular series and bearing spo-
radically isolated spinelets.__...............-.2222__.- Kalyptasterias Koehler.

. Adambulacral plates with a crowded transverse comb of four or five spines (three to six)
which carry attached pedicellariae; actinal area extensive with three to six longiseries of
well-developed 4-lobed, spiniferous plates; mouth angle rather broad, without com-
pressed adoral carina, two or three postoral pairs of adambulacral plates in contact;
PSrEmee (ORINIO® CMBR e ey B Perissasterias Clark.

a’. Abactinal surface devoid of a connected skeleton, soft and flexible, the abactinal plates, if
present, isolated, spaced, and superficially invisible.

b'. Marginal and abactinal skeleton consisting of lateral spaced, transverse tongues of plates
(composed of two reduced marginals plus sometimes one or several abactinals) abutting
on the adambulacral plates; dorsal surface with much reduced, disconnected, scattered
platelets; dermis thick, surrounding the spinelets in mammillated pustules; adambulacrals
monacanthid; gonads opening ventrally. (Anasterias authors, not Perrier.)

Lysasterias Fisher.

&. Neither marginal nor abactinal plates present, the skeleton reduced to the ambulacral and
adambulacral systems; body covered with numerous papillae, containing each a slender
spinelet, disposed without order on all surfaces; adambulacrals diplacanthid.

Adelasterias Verrill.

220 BULLETIN 76, UNITED STATES NATIONAL MUSEUM
Genus ADELASTERIAS Verrill

Adelasterias VeRRiILL, Shallow-water Starfishes, 1914, p. 360. Type, Diplasterias papillosa
Koehler.—FisHEr, 1923, p. 607.

Diagnosis.—Neither marginal nor abactinal plates present, the skeleton reduced
to the ambulacral and adambulacral systems; body covered with numerous papillae,
each containing a slender spinelet, disposed without order on all surfaces; adambu-
lacral plates diplacanthid.

1. ADELASTERIAS PAPILLOSA (Koehler).
Diplasterias papillosa Korner, 1906, p. 21, pl., figs. 2-5; pl. 2, figs. 18, 19; 1912, p. 21.

Graham Land and vicinity, western Antarctica; low tide to 6 meters.
Genus ALLOSTICHASTER Verrill

Allostichaster VeRRILL, Shallow-water Starfishes, 1914, p. 363. Type, Asieracanthion polyplax
Miiller and Troschel.—Cuark, 1916, p. 70.—FisHEr, 1923, p. 575.

Diagnosis.—Body covered with small granuliform spinelets; skeleton close-knit,

the plates in definite longiseries; carinals and superomarginals broader than other
plates; inferomarginals forming ventrolateral edge to ray; dorsolateral area narrow,
the plates in a single straight or zigzag series (altogether three series of abactinal
plates); one series of actinal plates; inferomarginal plates with oblique transverse
series of a few spinelets along ventrolateral margin of ray; cleaned superomarginal
plates with minutely beaded surface; adambulacral plates diplacanthid; pedicellariae
scattered among spinelets; multiple madreporic bodies; fissiparous; rays six to eight.

1. ALLOSTICHASTER POLYPLAX (Miiller and Troschel).

Asteracanthion polyplax Miuuer and Troscuen, Arch. Naturges., 1844, p. 178.
Asterias polyplax Perrier, Réy. Stell., 1875, p. 63.—Ciakx, Echinoderms Western Austra-

lian Mus., 1913, p. 151.
Stichaster polyplax SLADEN, 1889, p. 432.—F arquuar, 1897, p. 196; 1898, p. 313; 1909, p. 129.—

BENHAM, 1909, p. 15.
Tarsaster neozealanica FaRquuAR, Trans. New Zeal. Inst., vol. 27, 1895, p. 207, pl. 12.
Allostichaster polyplax Cuark, Biol. Res. Fishing Exper. Endeavour, vol. 6, 1916, p. 70.—
Korn LER, 1920, p. 85, pl. 18, fig. 7-11; pl. 69, fig. 2—Morrrnsrn, 1925, p. 315, fig. 19a,b.

Tasmania, Australia, New Zealand.

2. ALLOSTICHASTER INSIGNIS (Farquhar).

Stichaster insignis FARQUHAR, Trans. New Zeal. Inst., vol. 27, 1895, p. 205, pJ. 13, fig. 1;
vol. 30, 1898, p. 188; Proc. Linn. Soe. New South Wales, 1898, p. 314.—Brnuam, 1909,
p. 15.

Allostichaster insignis Koruunr, 1920, p. 85, pl. 20, figs. 2-7; pl. 65, fig. 2—Fisuer, 1923,
p. 595.— Mortensen, 1925, p. 316, fig. 19, c, d.

New Zealand; Auckland Islands.
3. ALLOSTICHASTER INAEQUALIS Koehler.
Allostichaster inaequalis KorHLER, 1923, p. 50, pl. 1, figs. 7, 8; pl. 7, figs. 1-3.

Magellanic region, 7 to 100 meters.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——-FISHER 221

4. ALLOSTICHASTER HARTI (Rathbun),

Leptasterias hartii R. Ratunun, Echinoderms of Brazil, Trans. Conn. Acad. Sei., vol. 5,
1879, p. 145.—Vernitt, 1915, p. 23.

The type was taken about 30 miles east of Cape Negro, Brazil, in 62 fathoms.
I have a specimen from the Museum of Comparative Zoélogy labeled “Patagonia,
44 fathoms” which, like the type, has six rays, and measures R 19 mm., r 3.5 mm.
It has one larger and two smaller madreporic bodies.

Another species which probably belongs to Allostichaster is Stephanasterias hebes
Verrill (1915, p. 26, pl. 9, fig. 3), which was taken by the Albatross at station 2766,
off the east coast of South America (lat. 36° 47’ S., long. 56° 23’ W.). The de-
scription isincomplete but the actinal aspect of the ray, as figured, suggests Allostichaster
rather than Stephanasterias. It is possible that the specimen of hartii referred to
above, really belongs in hebdes.

5. ALLOSTICHASTER REGULARIS Clark.
Allostichaster regularis CLarx, Rec. South Austral. Mus., vol. 3, No. 4, 1928, p. 400, fig.115.
South Australia: Spencer and St. Vincent Gulfs.

Genus ANASTERIAS Perrier

Anasterias Perrrer, Réy. Stell., 1875, p. 81; 1891, p.91. Type, Anasterias minuta Perrier.—
Fisner, Smithsonian Mise. Coll., vol. 52, 1908, p. 52; Zool. Anz., vol. 33, 1908, p. 356; Ann.
Mag. Nat. Hist., ser. 9, vol. 10, 1922, p. 592; vol. 18, 1926, p. 197.

Asteroderma Perrier, Comptes-rend., vol. 106, no, 11,1888, p. 763; Mission sci. Cap Horn,
1891, p. 96.

(Not Anasterias Ludwig, 1903; nor Koehler, 1906, 1908, 1912, 1920, 1923; nor Verrill, 1914.
See Lysasterias.)

Diagnosis.—Resembling Sporasterias but the abactinal skeleton typically
reduced to an open, delicate irregular mesh, entirely hidden by thick pulpy skin,
which in the adult, even when dried, may conceal the underlying plates. Both series
of marginal plates well developed; the superomarginals commonly monacanthid (or
spineless), the inferomarginals diplacanthid; one series of actinals, sporadically
spiniferous; adoral carina composed of about three pairs of contiguous postoral
adambulacral plates; interbrachial septum strongly calcified; gonads opening
ventrally —paedophoric.

Remarks.—The above diagnosis is intended to characterize two known species
of a larger group which includes Sporasterias and possibly Kalyptasterias, and which
by right of priority would be called Anasterias. This diagnosis of Anasterias is
therefore of the subgenus Anasterias. The diagnoses of Sporasterias and Kalyptas-
terias are also of subgenera. I have listed these groups separately in order to direct
attention to the peculiar characters of Anasterias, s. s., concerning which there has
been some misapprehension.

The generic name Anasterias has usually been incorrectly applied since Ludwig
first set the fashion in 1903 by using it for a group of small, paedophoric sea stars,
almost devoid of abactinal skeleton but characterized by a soft, pustulated, glandular
integument. These bear a strong superficial resemblance to Anasterias minuta

63160—30——15

222, BULLETIN 76, UNITED STATES NATIONAL MUSEUM

which, on the contrary, has a reticulate abactinal skeleton. Several authors, notably
Koehler and Verrill, have followed Ludwig. The genus erroneously called Anas-
terias was named Lysasterias by me in 1908.

The type specimen of Anasterias minuta was indicated by Perrier in 1891
(p. 93) as that bearing the label ‘‘ Voyage de l’Astrolabe; M. M. Hombron et Jacqui-
not,” and he gives reasons for believing it to have been collected at Port Famine,
Magellan Strait.

In 1923, thanks to the cooperation of Prof. L. Joubin, I examined this specimen
at the Museum d’Histoire Naturelle and published some notes on it. **

The type specimen of A. minuta is in alcohol and is labelled E 792, M. M. Hom-
bron et Jacquinot, 1847. The rays are short. Measurements, according to Perrier,
R 12 mm.,r6mm. The integument entirely conceals the abactinal skeleton. I
subjected the abactinal surface of one ray to treatment with potassium hydrate, which
revealed a fairly regular series of small carinal plates, between which and the regular
inferomarginals (each with one short spinelet) is an irregular, very open, incomplete
net of plates, strongest next to the superomarginals. The net is in part composed of
about 14 irregular transverse tongues of plates on either side, extending inward toward
the carinals and each corresponds roughly to a marginal plate. The inferomarginals
have two spines, which are the largest on the body and are separated from the smaller
single superomarginal by a fairly broad channel with a row of papulae. Actinals
apparently not developed; adambulacral plates strictly monacanthid. The abactinal
surface though soft and somewhat pulpy is not papillose in the sense that Lysasterias
and Koehleraster are. Abactinal spinelets are few and scattered.

It is quite necessary to fix the attention on this type specimen of Anasterias
minuta which establishes the identity of the species and hence of the genus. The
specimens of Anasterias minuta listed by Perrier in 1891 may or may not be A.
minuta and have nothing to do with the status of Anasterias.

It is a fact that Anasterias Perrier has a connected dorsal skeleton, whereas
Anasterias Ludwig has not; A. minuta also lacks the pustulated or papillose abactinal
integument which is described as being characteristic of Anasterias Ludwig (Lysas-
terias Fisher).

Anasterias Perrier is closely related to Sporasterias, as exemplified by S. ant-
arctica and its alleged synonym, S. rugispina (the genotype). In fact Leipoldt,
1895, synonymized Anasterias minuta with S. antarctica and S. rugispina; while
Perrier, himself, writing in 1891,%" says:

Les étoiles de mer que j’ai nommées Anasterias minuta, Asterias rugispina Stimpson; celles que

M. Studer appelle Asterias antarctica Liitken; celles que M. Bell a appellées Asterias spirabilis
sont done des formes extremement voisines, sinon identiques.

Furthermore Koehler (1920, p. 11) writes:

Je dois dire que j’ai eu en communication plusieurs Anasterias minuta du Jardin des Plantes
[but evidently not the type] et j’ai pu constater que toutes étaient parfaitment identiques A de
jeunes Sporasterias antarctica. J’estime donc.]’A. minuta forme jeune d’une Sporasterias ou peut-
étre d’une autre Anasterias, ne doit pas figurer parmi les éspeces du genre Anasterias.

*¢ Notes on the Asteroidea. IV. Ann. Mag. Nat. Hist., ser. 9, vol. 18, 1926, p. 197.
7 Miss. sci. Cap Horn, p. 15.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 223

It seems to me probable that the type specimen of A. minuta is not the young of
antarctica, nor of any species of Lysasterias, because I have examples of an Anasterias
from the Falkland Islands, which are distinet from Sporasterias antarctica, and which
_ I believe to be true A. minuta. (See below.) The open, weak, recticular, abactinal
skeleton, overlaid by a thick pulpy skin is like that of the type of A. minuta.

The species of Anasterias and of Sporasterias are extremely variable and the two
genera appear to me to be confluent. The distinguishing features for Anasterias are
the weaker, more open, abactinal skeleton and the thicker, pulpy skin, entirely ob-
scuring the abactinal plates and also sometimes the spinelets. But these characters
are relative. Some specimens of Sporasterias have a rather weak, open, abactinal
skeleton (especially when young); and a skin obscuring the plates and forming a
thick welt around the spinelets is also rather common. I have some specimens col-
lected by Dr. W. L. Schmitt near Teal Inlet, Falkland Islands, which on the basis
of skeleton and spinelets are Sporasterias, but of dermal characters are Anasterias.
It is probable that the thicker skin is directly correlated with a weak abactinal
skeleton. The formation of local formae is undoubtedly encouraged by the seden-
tary habits of these sea stars and by their habit of rearing a relatively few young
in a temporary suboral brood chamber.

Kalyptasterias conferta Koehler is closely similar to Anasterias minuta. <A well-
hardened alcoholic specimen of the latter resembles the figures of K. conferta in
Koehler’s sea stars of the Swedish Antarctic Expedition (1923, pl. 4, figs. 3 and 4),
The principal anatomical difference is in the weaker, less regular, superomarginals of
Kalyptasterias—a distinction which seems to me to be specific or subgeneric, rather
than generic.

In tabular form the species would be arranged as follows:

Genus Anasterias Perrier, 1875.
1. Subgenus Anasterias—including A. minula, A. pedicellaris, and possibly A.
victoriae Koehler (Lysasterias victoriae).
2. Subgenus Sporasterias Perrier, 1894—including A. antarctica, A. directa, A.
mawsoni, A. rupicola, A. perrieri, A. sphaerulata (see below).
3. Subgenus Kalyptasterias Koehler, 1923—including A. conferta.

If it should eventually be proven that the type specimen of Anasterias minuta
is really a young Sporasterias antarctica, then obviously the same species is the type
of both Anasterias and Sporasterias. The broader genus would continue to be called
Anasterias, the oldest name, which would also be the name of the subgenus called
Sporasterias above. The subgenus Anasterias of the present scheme would be called
Asteroderma, with A. papillosum for type (unless it, too, is a young antarctica).

I have included the two following forms:

1. ANASTERIAS MINUTA Perrier.

Anasterias minuta Pernter, Réy. Stell., 1875, p. 81; Mission sci. Cap Horn, 1891, p. 93 (in
part).

? Asteroderma papillosum Pexrrer, Comptes-rend, vol. 106, no. 11, 1888, p. 765; Mission
sci. Cap Horn, 1891, p. 96.

? Asterias minula var Asteroderma papillosum Pennier, 1891, pl. 10, fig. 3a—3c.

Strait of Magellan, Tierra del Fuego, Falkland Islands.

224 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

The type of this species is in the Museum d’Histoire Naturelle (EK 792 M. M.
Hombron et Jacquinot, 1847, alcohol). See antea. Perrier considered the type
locality to be Port Famine, Magellan Strait.

Two dried specimens from Darwin Harbor, Choiseul Sound, Falkland Islands
(No. 2623 Mus. Comp. Zo6l.) apparently belong to this species. The smaller meas-
ures R 18 mm., r 6 mm.; the larger R 48 mm., r 13 mm. In the smaller example
the abactinal skeleton is clearly visible and consists of a weak, irregular reticulum
resembling the condition in A. pedicellaris as figured by Koehler (1923, pl. 5, figs. 1
and 4) and closely similar to that of the type specimen of minuta. Most of the super-
omarginals carry one spinelet and the inferomarginals two, while scattered along the
intermarginal channel and inside the furrow margin are rather numerous, lanceolate
straight pedicellariae two-thirds the length of the superomarginal spinelets. In the
larger specimen, however, the abactinal integument has thickened and conceals the
skeleton which is quite weak and irregular as in Koehler’s Plate 5, Figure 1, alluded
to above. The proportions are about as in Figure 4. There are a few actinal plates
and spines at the base of the ray and the adoral carina is composed of three pairs
of contiguous adambulacrals, the first pair larger than second, and the second larger
than third. The superomarginal spines have been mostly absorbed; pedicellariae as
in the small example. A third specimen (No. 2624) carries a cluster of young.

Sixteen specimens from Port Stanley, Falkland Islands, collected in 1927 by
Dr. W. L. Schmitt. These are evidently conspecific with the Darwin Harbor examples.
A well-hardened alcoholic example (R48 mm.) resembles the Kalyptasterias conferta
figured by Koehler (1922, pl. 4, figs. 3 and 4). The abactinal plates are slender,
delicate, and form an irregular reticulum, with very large meshes, and are entirely
hidden, until dried, by the soft pulpy integument. Dorsal spinelets few and widely
scattered; only a few abactinal crossed and straight pedicellariae. Superomarginal
plates normal, not massive, each with one blunt, terete, slender spinelet, 1 to 1.5 mm.
long; inferomarginals with two, decidedly stouter and longer spines; actinal plates
with one spine, slightly smaller, the series extending two-thirds length of ray, each
spine forming with the inferomarginal spines a transverse series of three. Numerous,
rather thickly lanceolate, subobtuse straight pedicellariae, decidedly longer than
broad are scattered on the marginal and actinal plates, in the intermarginal channel
and along edge of furrow. No associated cross pedicellariae, except near end of ray
and there only a few. [In Sporasterias antarctica the superomarginals are normally
surrounded by crossed pedicellariae, and the inferomarginal plates carry at least a .
few on the intermarginal side of the spines.]

Another lot of nine from Port Stanley (April 16, 1927) differs in having numerous
small capitate abactinal spinelets, and fairly numerous abactinal (but not marginal)
crossed pedicellariae; straight pedicellariae scattered over abactinal surface and
distributed laterally and actinally without associated crossed pedicellariae; integu-
ment thick, pulpy, in aleohol. When dried a specimen would pass for an aberrant
Sporasterias antarctica, with weak dorsal skeleton. One specimen carries a thick
mass of eggs in the oral concavity.

Fourteen specimens from near Teal Inlet, Falklands, also collected by Doctor
Schmitt, do not belong to this species but are mentioned because they have the thick
dorsal integument and abactinal spinelets of the above lot. When dried they

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 225

resemble Sporasterias antarctica, almost devoid of crossed pedicellariae and with only
a few straight pedicellariae laterally (but with a series of small! ones on furrow margin).
In some but not all specimens a few crossed pedicellariae occur sporadically here and
_ there on the upper side of the inferomarginal plates (where there are none in typical
specimens of minuta). At the present writing I do not know what these are.

I was not able to find a specimen of Perrier’s Asteroderma papillosum at the
Museum d'Histoire Naturelle.

2. ANASTERIAS PEDICELLARIS (Kochler.)

Sporasterias pedicellaris Konuuter, Swed. Ant. Exp., vol. 1, no. 1, 1923, p. 18, pl. 5, figs.
1-6; pl. 6, figs. 1-5, 7-10.

Sporasterias antarclica Koenver, 1920, p. 78, pl. 18, fig. 4.

? Anasterias perrieri Perrier, Mission sci. Cap Horn, 1891, p. 7 (not Studer, 1885). I
have seen a specimen at the Museum d'Histoire Nature'le.

This species is fully described and figured by Koehler. He notes that in some
specimens the spines and characteristic pedicellariae may be very slightly developed,
or are even lacking. He states also that the external appearance, in respect to the
more or less soft consistency of the abactinal wall, resembles that of Lysasterias.

Under this nominal species I have tentatively classified several specimens,

mostly in a bad state of preservation, which were collected by Dr. W. L. Schmitt
at Port Stanley, Falkland Islands. The abactinal spinelets are fairly numerous,
and scattered among them are numerous crossed pedicellariae. In contrast to the
condition in minuta, a few crossed pedicellariae here and there accompany the con-
spicuous lanceolate pedicellariae of the superomarginal and inferomarginal plates.
R 40 mm., r 12 mm.
. Koehler’s material was taken at Tierra del Fuego, 36 meters, and the Falkland
Islands, 7 to 10 meters. Certain specimens collected at low tide, Sparrow Cove,
East Falkland, are probably the same as my minuta, since “les piquants et les pedi-
cellaires sont trés peu développés et ils font méme complétement défaut sur certains
exemplaires.”’

It is likely that pedicellaris and minuta are two ‘‘manifestations’
species.

of a single

Genus CALVASTERIAS Perrier

Calvasterias Pennier, Rév, Stell., 1875, p. 84. Type, C. asterinoides Perrier.—Fisnen, 1922,
p. 597; 1923, p. 597.

Stichorella Koru.en, Austral. Ant. Exp., vol. 8, 1920, p. 89. Type, Stichaster suteri de Lorio!.
Diagnosis —Broad-rayed, monacanthid, paedophorie species, with the skeleton
more of less concealed by smooth integument, and with a rather sharp ventrolateral
margin to ray marked by oblique combs of two to four short inferomarginal spines;
superomarginals very broad (largest plates of all) bearing one or two granuliform
spinelets; dorsolaterals not very numerous, trilobed, in one or two fairly regular
longiseries; a definite series of strongly imbricated 4-lobed carinals; abactinal skeleton
with only a few granules, or with numerous granules in longitudinal series; an in-
conspicuous series of actinal plates, a few of the proximals sporadically spiniferous;
gonads opening ventrally; actinostome sunken, the adoral carina narrow and com-

posed of four or five pairs of contiguous postoral adambulacral plates.

226 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

1. CALVASTERIAS ASTERINOIDES Perrier.
Calvasterias asterinoides PERRInR, 1875, p. 84.
Falkland Islands; probably not Torres Strait, as specified by Perrier.

2. CALVASTERIAS ANTIPODUM Bell.
Calvasterias antipodum Brun, Proc. Zool. Soc., 1882, p. 122, pl. 6, fig. 1.

Mr. G. A. Smith of the British Museum (Natural History) has sent me a de-
scription of the type together with photographs of the skeleton. This species is a
typical Calvasterias with thick skin and very large superomarginal plates. The
carinal ridge carries a narrow zone of coarse tubercles (one or two wide) continued
as a pentagon on the disk. No others are present. The superomarginals are very
broad, with a large beaded area, and are wider than the dorsolateral area. Each
inferomarginal carries two short, obtuse, granular spines which are set at an angle
of 45° to the wall of the ray. On the dorsal side of their bases these spines have
become engulfed in the thick integument.

3. CALVASTERIAS LAEVIGATA (Hutton).

Asterias rupicola var. laevigata Hurron, Trans. New Zeal. Inst., vol. 11, 1879, p. 353.

Stichaster suteri var. laevigatus FarquHaAR, Trans. New Zeal. Inst., vol. 30, 1897, p. 189;
Proe. Linn. Soc. N. 8S. Wales, 1898, p. 313—BrnuHaAm, Subantarctic Islands of New
Zealand. Echinoderms, 1909, p. 302.

Calvasterias laevigata FisuEr, 1923, p. 606—Mortensen, 1925, p. 311, pl. 13, fig. 12; pl. 14,
figs. 3-10.

New Zealand: Auckland Islands; Campbell Island.

4. CALVASTERIAS SUTERI (de Loriol).

Stichaster suteri pp Lorton, Rev. Suisse de Zool., vol. 2, 1894, p. 477, pl. 23.—Farquuar,
Journ. Linn. Soe. Zool., vol. 26, p. 197; Proc. Linn. Soc. N. 8. Wales, 1898, p. 313.—
Brenuam, Subant. Islands of New Zealand. Echinoderms, 1909, p. 302.

Stichaster littoralis FarquuHaRk, Trans. New Zeal. Inst., vol. 27, 1895, p. 206, pl. 13.

Stichorella sutert KorHuER, 1920, pp. 87-89, pl. 22, figs. 1, 2, 4; pl. 62, fig. 3; pl. 63, fig. 1.

Calvasterias suteri FisHER, 1922, p. 597; 1923, p. 606—MortTENSEN, 1925, p. 310.

New Zealand.
5. CALVASTERIAS STOLIDOTA Sladen.
Calvasterias siolidota SuADEN, Challenger Asteroidea, 1889, p. 590, pl. 101, figs. 3, 4; pl. 103,
figs. 11, 12.

Port William, Falkland Islands; Messier Channel, southern Chile.

Mr. G. A. Smith has sent me a description and photographs of the skeleton of
this species which, from Sladen’s figures, resembles a Kalyptasterias. The dorso-
lateral area is broader and the superomarginals much smaller than in typical Calvas-
terias. The abactinal skeleton is well developed with strongly overlapping carinals.
The mostly 3-lobed dorsolaterals form an irregular open reticulum, in which three
or four meshes can be counted between the carinals and superomarginals. The area
is nearly or quite twice as wide as the superomarginal plates, and hence much broader
than in antipodum, in which the superomarginals are wider than the dorsolateral area.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 227

Genus COSMASTERIAS Sladen

' Cosmasterias Suave, subgenus, Challenger Asteroidea, 1889, p. 576. Type, Asteracanthion
sulcifer Perricr= A. luridum Philippi.
Diplasterias Pennien, part, Mission sci. Cap Horn, 1891, p. 77.— Meissner, 1904, p. 6.
Comasterias Penner, Mission sci. Cap Horn, 1891, p. 159.
Podasterias Pennien, Mission sci. Cap Horn, 1891, p. 160.
Quadraster Pennien, Rés. camp. Prince de Monaco, 1896, p. 27. Type, Stichaster felipes
Sladen.

Diagnosis.—Actinal plates in two or more prominent longiseries forming the
actinal or the actinal and actinolateral surfaces of ray (and not dominated or over-
hung by the inferomarginal plates); adambulacral plates diplacanthid, without
attached pedicellariae; large unguiculate straight pedicellariae; plates of the abactinal
area of ray have the appearance of forming definite longiseries, the carinals and supero-
marginals being conspicuously regular, and the dorsolateral area generally rather
narrow with the plates in discernible longiseries; actinostome sunken, the adoral
carina long, consisting of three or more contiguous pairs of adambulacral plates;
gonads opening dorsally; not paedophoric.

Southern Hemisphere.

This genus is in need of thorough revision. It appears to be distantly allied to
Stichaster. In the following provisional list of species I have not included Asterias
(Polyasterias) fernandensis Meissner,® listed by Verrill (1914, p. 360) as Cosmasterias
fernandensis. The genus is probably represented on Juan Fernandez, but Meissner’s
species is almost certainly the young of Astrostole pla‘ei “ from the same locality.

I have omitted also Cosmasterias radiata Koehler (1923, p. 36, pl. 2, fig. 5, pl. 3,
figs. 1 and 2) dredged in 160 meters on Shag Rock Bank, between Falkland Islands
and South Georgia. The specimens, almost certainly immature, have 8 to 10 equal
or subequal rays, and a single madreporic plate. The species is fully described by
Koehler who has placed it provisionally in Cosmasterias.

1. COSMASTERIAS LURIDA (Philippi).

Aateracanthion luridum Puitirrt, Arch. Naturges., vol. 24, 1858, p. 265.

Asteracanthion sulcifer Pennier, Ann. Sci. Nat., 1869, p. 235.

Asterias sulcifera Pennien, Réy. Stell., 1875, p. 58.

Asterias (Cosmasterias) sulcifera Suapen, Challenger Asteroidea, 1889, p. 562.—Lerrotprt,
Zeit. Wiss. Zool., vol. 59, 1895, p. 552 (literature list).

Comasterias sulcifera Pennten, Miss. Sci. Cap. Horn, 1891, p. 160.

Cosmasterias sulcifera Pennien, Expéd. Travailleur et Talisman, 1894, p. 107.

Cosmasterias lurida Lupwia, Expéd. Antarct. Belge, 1903, p. 40.—pr Lorion, 1904, p. 39.—
Kornter, 1912, p. 22, pl. 2, fig. 1-7; pl. 5, fig. 8; 1923, p. 39.

To this species Leipoldt, 1895, unites Asteracanthion clavatum Philippi, A. fuleum
Philippi, A. spectabile Philippi, A. mite Philippi, and Stichaster polygrammus Sladen.
(See below.)

Region of the Straits of Magellan and adjacent coasts on both Atlantic and
Pacific sides; Tierra del Fuego; south Georgia; low tide to 348 fathoms.

” Meisaner, Arch. Naturges., 1904, p. 104, pl. 6, figs. 1, le-1d.
™ Lieberkind, The Natural Hist. of Juan Fernandes. Edited Skottsberg, vol. 3, 1920, p. 384 (record of a young Cosmasterias).
@ Meissner, 1896, p. 103, pl. 6, fig. 2, Asterias (Coscinasterias) platei. See also Lieberkind, 1920, p. 387.

228 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

2. COSMASTERIAS GERMAINI (Philippi).
Asteracanthion Germaini Puiuipri, Arch. Naturges, 1858, p.£265.— Korner, 1912, p. 19.
Asterias (Cosmasterias) tomidata SuapEN, Challenger Asteroidea, 1889, p. 576, pl. 105, figs.
8-10.
Diplasterias Germaini (Philippi), Meissner, Hamburger Magalhaen. Sammelr., 1904, p. 7.
Cosmasterias tomidata VERRILL, 1914, p. 359.
Gulf of Pefias, Chile, 45 fathoms; region of the Straits of Magellan; off Argentina,
latitude 44° 52’ S., 55 fathoms (Mus. Comp. Zoél.).
3. COSMASTERIAS POLYGRAMMA (Sladen).
Stichaster polygrammus SLADEN, 1889, p. 434, pls. 100, 103.
Cosmasterias polygrammus VERRILL, 1914, p. 360.

West of Magellan Strait, 245 fathoms (Sladen).
4. COSMASTERIAS CAPENSIS (Perrier).
Asterias capensis Perrier, Révy. Stell., 1875, p. 73.
Stichaster felipes SLADEN, 1889, p. 433, pl. 101, figs. 1 and 2; pl. 103, figs. 7 and 8.
Quadraster felipes Perrier, Rés. camp. Prince de Monaco, 1896, p. 27.
Cosmasterias felipes FisHpr, Ann. Mag. Nat. Hist., ser. 9, vol. 10, 1928, p. 198.
Cosmasterias capensis FisHeR, Ann. Mag. Nat. Hist., ser. 9, vol. 10, 1923, p. 198.
Cape of Good Hope, to 150 fathoms.
5. COSMASTERIAS DYSCRITA H. L. Clark.
Cosmasterias dyscrita CLARK, Biol. Res. Fishing Exper. carried on by thel F. I. S. Endeavour,
1909-1914, vol. 4, pt. 1, 1916, p. 71, pl. 29, figs. 1, 2.

South of Gabo Island, Victoria, 200 fathoms.
Genus CRYPTASTERIAS Verrill

Diplasterias Konner, part, Expéd. antarct. Francaise, Echinodermes, 1906, p. 19; 1908,
p. 574.

Cryptasterias VERRILL, Shallow-water Starfishes, 1914, p. 362. Type, Diplasterias turqueti
Koehler—Koeru ter, 1920, p. 57; 1923, p. 25. Fisher, 1923, p. 604.

Diagnosis —Related to Diplasterias, but abactinal surface soft, rather irregularly
and finely papillose and the abactinal plates thin (with rudimentary spinelets only),
forming an irregular reticulum entirely concealed by the skin; superomarginal plates
well developed, bearing one spine wreathed with pedicellariae, and having long descend-
ing processes which support a broad, vertical, intermarginal channel; inferomarginals
with two webbed spines (each with a cluster of pedicellariae on outer face) or with
one spine; no actinal plates; adambulacral plates regularly diplacanthid; with two
contiguous pairs of postoral adambulacral plates; interbrachial septum strongly
calcified; gonads opening ventrally, eggs very large; good-sized, broadly lanceolate
straight pedicellariae on furrow margin; rays five to eight.

Koehler, 1923, p. 25, includes two species, the type and C. brachiata. I am not
certain of the status of Diplasterias induta Koehler (1908, p. 575) which is described
as being close to turqueti and hence probably a Cryptasterias. But Koehler (1923,
p. 27) states that other than branchiata, turqueti is the only known species.

1, CRYPTASTERIAS TURQUETI (Koehler).
Diplasterias turqueti KoruiEr, 1906, p. 19, pl 2, fig. 17; pl. 4, fig. 39.
Cryptasterias turqueti VERRILL, 1914, p. 362.—KoruuER, 1920, p. 57, pl. 17, figs. 1-5.

Booth Wandel Island; South Orkney Islands; 9 to 20 fathoms.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 229

2 CRYPTASTERIAS BRACHIATA Koehler,
Cryptasterias brachiata Korner, 1923, p. 25, pl. 1, figs. 9, 10.

Berkeley Sound, Falkland Islands, 25 to 30 meters.
Genus DIPLASTERIAS Perrier

Diplasterias Perrier, Comptes-rendus, vol. 106, no. 11, 1888, p. 765; Mission sci. Cap Horn,
Echinodermes, 1891, p. 77. Type, Diplasterias litkeni Perrier.—Fisuer, Smithsonian
Miscell. Coll., vol. 52, 1908, p. 89. (Error in assignment of sulcifera as type.)

Podasterias Pernier, Expéd. sci. Travailleur et Talisman, 1894, pp. 107, 108; Camp. sei. du
Prince de Monaco, 1896, p. 35. Type, Diplasterias litkeni Perrier (not Podasterias Per-
rier 1891, p. 160).—Fisner, Ann. and Mag. Nat. Hist., ser. 9, vol. 12, 1923, p. 603;
Zool. Anz., vol. 33, 1908, p. 358.

Diagnosis —Adambulacral plates dipleanathid, without attached pedicellariae;
abactinal skeleton irregularly reticulate; the plates with one to several slender spines
surrounded by a collar of pedicellariae and more or less heavily sheathed with mem-
brane which usually obscures skeleton; marginal plates well developed; supero-
marginal spines one or two, conspicuous, clearly differentiated from abactinal and
surrounded by pedicellariae; inferomarginal spines usually two, sporadically three,
bearing pedicellariae and forming definite longitudinal series separated from supero-
marginals by a well-defined intermarginal channel; actinal spines with attached
pedicellariae, in one or two longiseries (actinals exceptionally spineless); two to four
pairs of contiguous postoral adambulacral plates; not fissiparous; gonads opening
ventrally; young brooded by adult; straight pedicellariae lanceolate, typically not
enlarged nor unguiculate. Differing from Anasterias in having a stronger abactinal
skeleton and diplacanthid adambulacral plates.

Remarks.—In the first reference to Diplasterias (1888) Perrier cites D. litkeni
Perrier and D. steineni Studer as species which brood their young. He adds: “Je
classe dans le genre Diplasterias les Asterias qui ont deux rangées au moins de
piquants adambulacraires.”’

In the Cape Horn report, published three years later, Diplasterias is entered as a
new genus, with the same diagnosis. Five species are enumerated and described:
D. sulcifera (Perrier, 1869), D. loveni Perrier, D. liitkeni Perrier, D. spinosa Perrier,
D. steineni (Studer).

Diplasterias was thus formally published in connection with definite species from
which a type may be selected. If the first reference is to be regarded as a nomen
nudem, as I suggested in 1908 (Fisher, 1908a, p. 89) the enumeration of D. litkeni and
D. steineni (both of which are Diplasterias in the narrowest sense) is useful as an indi-
cation of the species which Perrier had in mind for his new, if somewhat indefinite,
genus. While Perrier’s Cape Horn report was in press, Sladen used Perrier’s first
species, D. sulcifera, for the type of Cosmasterias. My choice, in 1908, of D, sulcifera
as the type of Diplasterias was ill advised, due to a strict adherence to the “first
species rule"’ for the determination of a type when none is indicated by the author.
It seems to me that the oldest known species, Asterias brandti Bell, which includes
both Diplasterias liitkeni and loveni (cited by Perrier in 1891) should be chosen, espe-
cially as D. litkeni appears in the first, or Comptes-rendus, citation.

Diplasterias obviously antedates Podasterias, Perrier 1894, even if the generic
diagnosis is vague in the light of modern information. Diplasterias made no deep

63160—30——16

230 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

impression on Perrier’s mind, because he readily abandons it on page 160 of the Cape
Horn paper, in a comment on Sladen’s recently published subdivisions of Asterias.
Here he merges his Diplasterias with Sladen’s Asterias.

On the same page (1891, p. 160) he unfortunately introduces Podasterias for the
Asterias suleifera group, stating that Cosmasterias has priority for this generic divi-
sion. This is the first occurrence of Podasterias in literature. But in 1894," disre-
carding his Podasterias of 1891, he introduced a quite different Podasterias for species
near A. liitkeni. This is precisely synonymous with his Diplasterias of 1891, and is
the Podasterias used by Verrill, Koehler after 1917, and Fisher, 1923. Podasterias of
1891 must obviously be discarded for Cosmasterias, while that of 1894 had no status
when proposed, being a homonym of the first, as well as a synonym of Diplasterias.

Disregarding his Diplasterias of 1891, Perrier in 1894 * employs Diplasterias in
a different sense, namely for Asterias Linnaeus. In this paper the Asteriidae lack a
genus Asterias.

In 1896 * Podasterias is preserved for Diplasterias litkeni, Asterias is reinstated,
and Diplasterias discarded entirely. ;

Ludwig in 1903 “ used Diplasterias in the sense in which I advocate employing
it. Meissner in 1904 * employed the name for Cosmasterias plus Diplasterias.
Koehler in 1906, 1908, and 1912, used Diplasterias to include Cryptasterias Verrill,
1914. In 1917 and subsequent papers Koehler adopted Podasterias with the limits
I have assigned to Diplasterias. In 1914 Verrill * used Podasterias for Diplasterias
but erroneously assigned Asterias brandti Bell, A. alba Bell, A. neglecta Bell, and A.
obtusispinosa Bell to Cosmasterias, the last three as possible synonyms of C. lurida
(Philippi).

Dr. R. Koehler has contributed two species to this genus and valuable notes and
figures of others in his beautifully illustrated reports on Antarctic sea stars ** wherein
the evolution of our knowledge of the group may be traced.

Subgenus DIPLASTERIAS ss.

Diagnosis.—Actinal plates in one or two longiseries; actinal spines in one or two
longiseries, armed with clusters of pedicellariae; adoral carina composed of from one
to three pairs of contiguous adambulacrals.

1. DIPLASTERIAS BRANDTI (Bell).

Asterias brandti Breuu,Proc. Zool. Soc., 1881, p. 91, pl. 9.

Asterias neglecta Bexu, Proc. Zool. Soc.,1881, p. 94. pl. 9.

Asterias alba Brut, Proc. Zool. Soe., 1881, pp. 92, 506.

Asterias obtusispinosa Bru, Proc. Zool. Soc., 1881, pp. 92, 93.

Asterias belli Sruper, Abh. Akad. Berlin, 1884, pp. 12, 18, pl. 1.

Asterias glomerata SuapEN, Challenger Asteroidea, 1889, p. 571, pl. 105.

Diplasterias lovent Perrier, Miss. sci. Cap Horn ,1891, p. 80.

Diplasterias liitkeni Perrier, Miss. sci. Cap Horn, 1891, p. 81.—Lupwie, 1903, p. 41.

4! Expéd. Travailleur et Talisman, pp. 107, 108 (key).

42 Résultats camp. sci. Prince de Monaco, fasc. 11, 1896, pp. 34, 35.

8 Expédition Anarctique Belge, Seesterne, p. 41.

“ Hamburger Magalhaensische Sammelreise, Asteroideen, pp. 5-9.

*s Shallow-water Starfishes, p. 361.

* See Koehler, 1908, p. 572 (Diplasterias); 1912, pp. 12 et seq. (Diplasterias); 1917, pp. 20-29 (Podasterias); 1920, pp. 35-51 (Podas-
terias); 1923, pp. 27-35 (Podasterias).

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 231

Diplasterias brandti Mrissnen, 1904, p. 7—Korurer, 1908, p. 572, pl. 5, figs, 50, 51; 1912,
p. 19, pl. 1, figs. 3, 5, 6.

Podasterias brandli Koruten, 1917, p. 26, pl. 4, figs. 16, 17; pl. 5, figs. 11, 13, 14; 1920, pp.
41, 51, pl. 13, fig. 11; pl. 16, figs. 5, 6; 1923 p. 27, pl. 2, figs. 2, 3.

Koehler points out that )). brandti is a polymorphic species. This is certainly
the case if glomerata, alba, and obtusispinosa, the types of which I have examined,
are to be ranked as one species. Alba and obtusispinosa are formae of the same species,
but they are wholly unlike glomerata. Bell's neglecta, a fine specimen of which is
in the British Museum (1901-5-10-65, Southern Cross expedition) appears to be
a fully grown example of forma glomerata (R 160 mm., r 24 or 25 mm.), neglecta
having priority.

It is likely that a number of well-marked formae will eventually be recognized.
Koehler has already given glomerata varietal rank (1923, p. 29).

South Georgia; Falkland Islands; region of the Straits of Magellan, north to
latitude 44° 14’ S., and longitude 53° 43’ W.

2. DIPLASTERIAS BRUCEI (Koehler).
Stolasterias brucei Koruer, 1908, p. 569, pl. 5, figs. 46, 47.
Coscinaslerias brucei Koenuer, 1911, p. 30, pl. 5, fig. 5.
Coscinasterias vicloriae Korner, 1911, p. 32, pl. 5, figs. 3, 4; 1912, p. 24.
Podasterias brucei Kornter, 1920, p. 42, pl. 11, figs. 5-7; pl. 13, figs. 1-9; pl. 14, figs. 4,
7-11; pl. 15, figs. 4, 5; 1923, p. 35, p. 13, fig. 2—Dépervein, 1928, p. 295.
South Orkneys; Graham Land; Adelie Land to longitude 92°.
3. DIPLASTERIAS FOCHI (Koehler).
Podasterias fochi Koruxer, 1920, p. 35, pl. 14, figs. 1, 2, 12; pl. 15, figs. 1, 2, 3,; pl. 16, figr.
2, 3, 7; pl. 58, fig. 1.
Antarctic from east longitude 92° to 141°; Adelie Land.
4. DIPLASTERIAS KERGUELENSIS (Koehler),
Podasterias kerguelensis Korner, 1917, p. 24, pl. 4, figs. 3, 4, 12, 13, 19.

Kerguelen.

5. DIPLASTERIAS MERIDION ALIS (Perrier).

Asterias meridionalis Perrier, Réy. des Stell., 1875, p. 76.

Asterias mollis Stuper, Abh, Akad. Wis., Berlin, 1884, p. 8.

Asterias studeri Beww, Proc. Zool. Soc., 1881, p. 91.—Stuper, Abh, Akad. Wiss., Berlin,
1884, p. 8, pl. 1, fig. 1.

Podasterias meridionalis Kornu.er, 1917, p. 20, pl. 2, figs. 11, 12; pl. 3, figs. 1, 2, 4, 5, 6, 7;
pl. 6, fig. 8; pl. 7, fig. 9; 1923, p. 31, pl. 2, fig. 1; pl. 13, fig. 1 —Dépencein, 1928, p. 294.

South Georgia; Kerguelen.
Subgenus Batuyastenias, new

Diagnosis —One series of spineless actinal plates. Abactinal skeleton weak,
irregularly reticulate, with small meshes; body overlaid by soft, puffy, wrinkled
skin which involves base of spines; rather wide intermarginal channel; superomarginal
plates monacanthid with long descending process which strongly overlaps the rather
long ascending lobe of the diplacanthid inferomarginals; conspicuous series of inter-
marginal and actinal papulae in addition to abactinal; actinostome small; adoral

De BULLETIN 76, UNITED STATES NATIONAL MUSEUM

carina narrow, comprising four pairs of postoral adambulacral plates. Type, Asterias
vesiculosa Sladen.
6. DIPLASTERIAS VESICULOSA (Sladen).
Asterias vesiculosa SLADEN, Challenger Asteroidea, 1889, p. 568, pl. 99, figs. 1, 2, pl. 1038,
figs. 1, 2.
Arafura Sea (lat. 5° 41’ S., long. 134° 4’ 30’’ E.), 800 fathoms.
I examined the type in the British Museum. This species seems clearly related
to Diplasterias ss., but lacks actinal spines and has a longer, narrower, adoral carina.

Genus EREMASTERIAS, new

Diagnosis.—Resembling Smilasterias Sladen in having a small-meshed, reticu-
late, abactinal skeleton beset with small irregularly disposed spinelets and a chevaux-
de-frise along the actinolateral border of ray consisting of consecutive transverse
combs of three or four, much larger, flattened, spatulate inferomarginal and actinal
spines, devoid of attached pedicellariae. Differing in having monacanthid adam-
bulacral plates and a more pronounced adoral carina consisting of several pairs of
adambulacral plates. Crossed pedicellariae scattered, rare, resembling those of
Sporasterias; a few incipiently unguiculate straight pedicellariae. One series of
spiniferous actinal plates. Type, Pisaster antarcticus Koehler.

Remarks.—The above diagnosis does not agree with Koehler’s excellent descrip-
tion in one important point. The position of the combs of actinolateral, spatulate
spines, distributed on two longiseries of plates, suggests that these plates are respec-
tively inferomarginal and actinal, as in Smilasterias, which Eremasterias strongly
resembles. Doctor Koehler identifies as marginals two longiseries, bearing small
spinelets, above these combs. I think it likely that Koehler’s superomarignal plates
are in reality a regular series of dorsolateral (abactinal) plates, while his inferomargi-
nals are superomarginals. Doctor Koehler states (p. 32) that the number of spines to
each comb does not diminish toward the end of ray. This is not norma] behavior
for actinal spines and points rather to the interpretation I have given above, namely
that the spines are borne on the inferomarginal and one series of actinals as in Smilas-
terias scalprifera. If Doctor Koehler’s interpretation is the correct one, then the
difference between Eremasterias and Smilasterias is even greater than indicated in
the diagnosis; for there is no doubt that the main part of the fan of spines in Smilas-
terias is carried on the inferomarginals. In S. triremis actinal plates are lacking.

This genus is not closely related to Pisaster, which has highly characteristic,
tiny, straight pedicellariac, with bifid faleate jaws, not found eleswhere among
Asteroidea.

1. EREMASTERIAS ANTARCTICA (Koehler).
Pisaster antarclicus KorHier, Ann. l’inst. océanographique, vol. 7, fasc. 8, 1917, p. 30, pl. 9,
figs. 5, 6, 7, 8, 10, 17.
Kerguelen, littoral.
Genus GRANASTER Perrier
Granaster Perrier, Expéd.sci. Travailleur et Talisman, 1894, pp. 129, 133. Type, Stichas-
ter nuiriz Studer.—FisueEr, 1923, p. 604 (including Hemiasterias).

Hemiasterias VeRRiLu, Shallow-water Starfishes, 1914, p. 362, footnote. Type, Granaster
biseriatus Koehler.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-~-FISHER 233

Diagnosis.—Small, paedophoric sea stars with short robust rays, diplacanthid
and triplacanthid adambulacrals, and the whole body covered with a rather uniform
coarse granulation of short, capitate spinelets, without serial arrangement, interspersed
with scattered pedicellariae; skeleton, not externally visible, consists of three to five
lobed plates, directly imbricated by their lobes, leaving skeletal intervals smaller than
plates; marginal plates four lobed, obscured by granules; an incomplete series of acti-
nals; papulae mostly solitary; interbrachial septum incomplete—simply a calcified
pillar passing obliquely upward from the odontophore to the dorsal wall, leaving a
large, free passage, adjacent to margin, from one ray to next; gonads large, opening
ventrally; adoral carina very short, composed of one pair of plates; tube feet in two,
three, or four series; straight pedicellariae ovoid or triangular, often nearly as large
as spinelets; crossed pedicellariae, few, relatively large, scattered among the spinelets.

This genus is very distinct and of uncertain relationship. The general facies
is unlike that of any other asteriid, the dorsal aspect suggesting a little Cycethra or
Perknaster. Only one species is known.

GRANASTER NUTRIX (Studer).

Stichaster nutriz Sruper, Jahrb. wiss. Anst. Hamburg, vol. 2, 1885, p. 1554, pl. 5a.

Granaster nutriz Perrier, 1904, p. 129.—Korner, 1923, p. 53.

Granaster biseriaius Koen er, 1906, p. 11, pl. 1, fig. 6; pl. 4, fig. 42; 1908, p. 565, pl. 5, figs.
48, 49; 1912, p. 29, pl. 3, fig. 2; pl. 6, fig. 1.

Hemiasterias biseriata Vernriui, 1914, p. 362.

South Georgia; South Orkneys; Graham Land; low tide to 250 meters.

I have examined two specimens from Wandel Island, Graham Land, West
Antarctica.

Genus KALYPTASTERIAS Koehler
Kalyptasterias Kosnuer, Swedish Ant. Exped., vol. 1, no. 1, 1923, p. 43. Type, K. conferta
Koehler.

Diagnosis.—Differing from Anasterias Perrier (which it resembles in having
monacanthid adambulacrals and a weak, irregularly reticulate, abactinal skeleton
overlaid by a thick, soft skin) in the weaker development of the superomarginal
plates, which form a rather irregular series and bear, sporadically, a single spinelet.
Whole body covered by a thick soft integument concealing the contour of the plates;
abactinal surface very convex, the two marginal series at the ventral border of ray;
abactinal surface with four or five irregular longiseries of numerous large papulae,
on either side of carinal line, alternating with longitudinal zones of bare skin, slightly
wrinkled; superomarginal plates rather small, irregular, inconstant in form, consti-
tuting a series very irregular and sinuous on some rays and more rectilinear on others;
inferomarginals normally developed, diplacanthid; one series of small, usually spinif-
erous, actinals. Abactinal spinelets small, fairly numerous to scarce; crossed pedi-
cellariae scarce, not in circles around spinelets; straight pedicellariae lanceolate.

This group is very close to true Anasterias. The latter has both series of mar-
ginals well developed and normal, but with increasing age the superomarginals may
lose their spine. See under Anasterias, Remarks.

There seems to be no question that this genus is perfectly distinct from Caleas-
terias. It will not be surprising however if it intergrades perfectly with Anasterias.

234 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

1. KALYPTASTERIAS CONFERTA Koehler.
Kalypltasterias conferta Kowuter, 1923, p. 43, pl. 4, figs. 4-7.
Falkland Islands (Port Louis, 4 to 7 meters).
Genus KOEHLERASTER Fisher

Koehleraster Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 10, Dec. 1922, p. 596. Type,
Anasterias octoradiala Koehler.

Diagnosis.—Differs from Lysasterias in baving an open but perfectly normal,
irregularly reticulate, abactinal skeleton and well developed marginal skeleton;
differs from Anasierias and Sporasterias in possessing a thick, highly pustulated,
“cauliflower” skin. Adambulacral plates monacanthid, but a few proximally
occasionally diplacanthid; a very inconspicuous series of small spineless actinal plates;
abactinal skeleton an irregular but complete reticulum of small, imbricated mostly
simple elliptical or oblong, occasionally faintly lobed plates; carinal series very
irregular; superomarginals monacanthid, inferomarginals mostly diplacanthid, the
spines surrounded by mammillated sheaths which contain (like those of dorsal sur-
face) a few fair-sized crossed pedicellariae without enlarged terminal teeth; numerous
scattered, narrowly spatulate to compressed, broadly lanceolate, often pedunculate
straight pedicellariae; tube feet crowded, large, quadriserial; gonads open ventrally,
although attached on level with marginal plates; adoral carina composed of four
pairs of contiguous adambulacral plates; numerous (eight or nine) rays.

KOEHLERASTER OCTORADIATUS (Koehler).
Anasterias octoradiata Korner, Science Bull. Mus. Brooelve Inst. Arts and Sciences, vol. 2,
no. 4, 1914, p. 64, pl. 14, figs. 1-7. South Georgia. Also 1923, p. 14, pl. 6, fig. 6.

South Georgia, 10 to 15 meters.

I have examined the type which is now in the United States National Museum.
This is eight rayed, while the second specimen (Koehler, 1923) is nine rayed.

The species is evidently allied to true Anasterias. It has a complete irregularly
reticulate abactinal skeleton consisting of very numerous, small, but fairly robust,
oval, elliptical-oblong, and a few irregularly 3-lobed plates (the latter perhaps repre-
senting primary dorsolaterals). There is an irregular carinal series, of which the
plates are no larger than the others. The abactinal plates are joined to the supero-
marginals by transverse bands of plates which are a little more regular than the others,
and between consecutive trabeculae are broad (but short) papular areas, which form
a zone just above the superomarginal plates. These papular areas, or skeletal meshes,
are larger than the other abactinal and the intermarginal meshes.

The marginal plates are fairly robust and of the form usually found in the
Asteriinae—namely, four lobed. The descending lobe of the superomarginal is the
longest, and strongly overlaps the ascending lobe of the inferomarginal. The supero-
marginals are regularly monacanthid, while most of the inferomarginals are dipla-
canthid. There is a very inconspicuous series of small spineless actinal plates.

The gonads are not present in all rays, and are attached to the body wall on a
level with the superomarginals; but the duct turns downward, and I believe that I

have demonstrated its external opening on the ventral surface, on a papilla, close to
the interradial line.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—-FISHER 235

The thick dermis is raised into conspicuous cauliflowerlike pustules which inclose
spinelets and entirely obscure the skeleton.
The genus is named in honor of Dr. Réné Koehler, of Lyons, the leading authority

on Antarctic echinoderms. ’
Genus LYSASTERIAS Fisher

Lysasterias Fisner, Smithsonian Miscell. Coll., vol. 52, May, 1908, p. 88; Zool. Anz., vol. 33,
Aug. 1908, p. 356. Type, Anasterias perrieri Studer, 1885, but not Perrier 1891.—Fisuer,
1922, p. 594; 1923, p. 606; 1926, p. 197.

Anasterias Lupwia, 1903, p. 42.—Koruter, 1906, p. 12; 1908, p. 566; 1912, p. 10; 1920, p. 11;
1923, p. 11.—Verritt, 1914, p. 354.—Not Anasterias Perrier, 1875; not Asteroderma
Perrier, 1875.

Paedasterias Verritt, Shallow-water Starfishes, 1914, p. 355. Type, Anasterias chirophora
Lupwic.—Koeuter, 1920, pp. 16, 30.

Diagnosis.—Marginal and abactinal skeleton much reduced, consisting of lateral,
spaced, transverse series or tongues of plates (composed of two reduced marginals
plus sometimes one or several abactinals) abutting on the adambulacral plates;
dorsal surface with much reduced, disconnected, scattered platelets; dermis thick,
surrounding the spinelets in mammillated pustules; adambulacral plates monacan-
thid; no actinal spines, the plates if present extremely rudimentary; gonads open
ventrally—paedophoric.

This genus has usually been incorrectly called Anasterias (q. v.).

Verrill’s genus Paedasterias was founded pretty largely upon an erroneous trans-
lation. Ludwig (1903, p. 42), in a synopsis of ‘‘ Anasterias’’ says: “‘ Die Spagen haben
(mit Ausnahme der Armspitze) keinen oberen Randstachel; grosse Tatzenpedicellarien
vorhanden * * * An. chirophoran.sp.’’ Verrill diagnoses the genus: ‘* * *
the upper marginal plates being absent except as rudiments distally.” But Ludwig
in a major division of his synopsis states that the upper marginal plates are present
in chirophora. Koehler (1920, p. 30) has adopted Paedasterias, and added a new
species, joffrei.

Koehler finds in his Paedasterias joffret abactinal cross-bars of several pieces, as
in Lysasterias belgicae (Ludwig); but he thinks that the superomarginals are not to be
found in these ares. Rather, they are one or two plates in the interbrachial angle
above the first inferomarginals, and quite independent of the little cross-tiers of
the rays, where the superomarginal would consequently be lacking. In a specimen
of Anasterias tenera which I have examined (No. 1842, Mus. Comp. Zodl.) these
“antiambulacral” arcs consist usually of two, sometimes three, plates; but the outer
usually spiniferous superomarginal is oriented transversely and the inner, or infero-
marginal, more longitudinally just as in chirophora, lactea, and perrieri. Koehler may
be correct in his interpretation of the lateral plates of joffrei and belgicae. But I
think that the outer of the two plates in the antiambulacral ares of chirophora, lactea,
and perrieri is to be interpreted as superomarginal. The type of Paedasterias equally
with that of Lysasterias would possess superomarginals. The presence in belgicae
and studeri of several plates (instead of two) in each lateral are probably represents
simply a less degenerate condition of the abactinal skeleton. In tenera one or two
small plates are sporadically present above the relatively large, unmistakable, trans-
versely oriented, superomarginal. These small plates are homologous with the addi-
tional lateral plates of belgicae. It follows, therefore, that there is nothing character-

236 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

istic about the marginal plates of chirophora, which are identical with those of the
type of Lysasterias. If Paedasterias is maintained as a genus, it must be on the
strength of the large spatulate unguiculate, straight pedicellariae. But in P. joffrei
these are less developed than in chirophora, while the lateral arcs are not those of
chirophora but of belgicae, in Ludwig’s second group. If the pedicellariae are used as
a criterion for separation, then the structure of the skeleton must be disregarded.
1, LYSASTERIAS ADELIAE (Koehler).
Anasterias adeliae Konner, 1920, pp. 16, 26, pl. 1, figs. 1, 2, 5-10, pl. 56, fig. 1.
Adelie Land, 12 fathoms.
2. LYSASTERIAS BELGICAE (Ludwig).
Anasterias belgicae Lupwic, 1908, p. 51, pl. 6, figs. 61-65; pl. 7, figs. 66-67.—KorEHLER,
1920, p. 17.
Latitude 70° 15’ S.; longitude 84° 06’ W.; 560 meters.
3. LYSASTERIAS CHIROPHORA (Ludwig).
Anasterias chirophora Lupwia, 1903, p. 43, pl. 5, figs. 52-54; pl. 6, figs. 55-60; pl. 7, fig. 78.
Paedasterias chirophora VERRILL, 1914, p 371—KornHLER 1920, p. 16.
From latitude 70° 23’ to 71° 19’ S. and longitude 82° to 87° W., 450 meters.
4. LYSASTERIAS CUPULIFERA (Koehler).
Anasterias cupulifera KonHLER, 1908, p. 566, pl. 5, fig. 52, pl. 6, figs. 58, 59; 1920, p. 16.
South Orkneys, 10 fathoms.
5. LYSASTERIAS JOFFREI (Koehler).
Paedasterias joffrei KoruiEr, 1920, p. 30, pl. 1, figs. 3, 4, 9; pl. 2, figs. 7, 8, 9; pl. 56, fig. 2.
Latitude 66° 50’ S.; longitude 142° 06’ E.; 354 fathoms.
6. LYSASTERIAS LACTEA (Ludwig).
Anasterias lactea Lupwia, 1903, p. 50.—Korn.er, 1920, p. 17.
Latitude 71° 09’ S.; longitude 89° 15’ W.; 450 meters.
7. LYSASTERIAS PERRIERI (Studer).

Anasterias perrieri Sruper, Die Seesterne Siid-Georgieus, Jahrb. wiss. Anst. Hamburg, vol.
2, 1885, p. 153.— Koruter, 1920, p. 17.

Anasterias lysasteria VERRILL, 1914,¥p. 354 (unnecessary substitute name; see Fisher, 1922,
p. 593).

South Georgia.
8. LYSASTERIAS STUDERI (Perrier).
Anasterias studeri Perrier, Miss. sci. Cap Horn, 1891, p. 99.—Kornuer, 1920, p. 17.
Near Atlantic entrance to Magellan Strait, 320 meters.
9. LYSASTERIAS TENERA (Koehler).

Anasterias tenera Konuter, 1906, p. 12; pl. 2, figs. 11-16; pl. 3, figs. 27, 28; pl. 4, fig. 41;
1908, p. 569; 1912, p. 10; 1920, p. 11, 17.

South Shetland Islands; Graham Land; South Orkneys, Cumberland Bay,
South Georgia; 1 to 420 meters.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS-~FISHER 237

10, LYSASTERIAS VICTORIAE (Koehler).
Anasterias victoriae Korner, 1920, p. 17, pl. 2, fig. 5; pl. 3, figs. 1-6; pl. 4, figs. 1-4; pl. 5,
figs. 1-10; pl. 6, figs. 1-4; pl. 57, fig. 1.
From latitude 64° to 67°S., and from longtitude 92° to 145° E., 60 to 354 fathoms.
This species is possibly a true Anasterias.

Genus NEOSMILASTER, new

Diagnosis —Resembling Diplasterias in having diplacanthid adambulacral plates,
and gonads which open ventrally, but differing in having abactinal pedicellariae
scattered among the spinelets rather than in circlets on the circumspinal sheaths; in
lacking a definite, well-marked series of enlarged superomarginal spines; in having
the inferomarginal spines in oblique, transverse combs of three or four (as in Smilas-
terias) but with a few attached pedicellariae; one series of actinal spinelets close to
adambulacrals; an intermarginal series of prominent lanceolate pedicellariae;
abactinal skeleton an irregular, close, or medium open net, the small spinelets, with
interspersed papulae, either close-set or spaced, according to the relative compact-
ness of skeleton; superomarginal spinelets similar to abactinal. Type, Asterias
georgiana Studer.

The species of this group resemble Smilasterias, but differ in having ventrally
opening gonads.

I, NEOSMILASTER GEORGIANUS (Studer). °

Asterias georgiana Stuper, Die Seesterne Siid-Georgiens, Jahrb. wiss. Anst. Hamburg, vol.
2, 1885, p. 150.

Diplasterias georgiana Perrier, 1891, p. 7.

Podasterias georgiana Korner, 1917, p. 26; 1920, p. 41.

Ctenasterias georgiana Korner, 1923, p. 40, pl. 3, figs. 3-7, 10.

I have examined at the British Museum one of the original types purchased
from the Hamburg Museum. Koehler has published (1923) good photographs. In
my opinion georgianus is not closely related to Leptasterias (with which Verrill’s
Ctenasterias is synonymous), although there is undoubtedly a superficial resemblance,
which exists also between some species of Sporasterias and Leptasterias.

The species has been taken at South Georgia, low tide to 75 meters; and Shag
Rock Bank, between Falkland Islands and South Georgia, 160 meters.

2. NEOSMILASTER STEINENI (Studer).
Asterias steineni Stupver, Die Seesterne Stid-Georgiens, Jahrb. wiss. Anst. Hamburg, vol. 2
1885, p. 152, pl. 1, fig. 4a, b.
Diplasterias steineni Penner, 1891, p. 84.— Koenver, 1912, p. 20, pl. 1, figs. 4, 7, 10.
Podasterias steineni Koeuter, 1917, p. 26; 1920, p. 41; 1923, p. 30, pl. 3, figs. 8, 9.

Tierra del Fuego; Falkland Islands; south of Cape Horn; 99 meters.

This species is clearly not congeneric with Diplasterias brandti and brucei. The
small abactinal spinelets with scattered pedicellariae not in circlets; the absence of
a conspicuous single or double series of superomarginal spines, either larger or plainly
distinct from the abactinals; the inconspicuous intermarginal area or furrow; the
crowded inferomarginal transverse combs of three or four spinelets—all segregate
this species from typical Diplasterias and ally it to georgianus.

nw
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BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Genus PERISSASTERIAS Clark
Plate 71, Figure 3

Perissasterias H. L. Cuarx, The Echinoderm Fauna of South Africa, Annals of the South
African Museum, vol. 8, pt. 7, May 1923, p. 307. Type, P. polyacantha Clark.—FisuHER,
1926, p. 199.

Diagnosis.—Adambulacral plates short, with numerous (usually four or five but as
few as two or three and as many as six or seven) spines, in transverse series, bearing
small lanceolate straight pedicellariae; actinal area extensive with three to six longi-
series of well-developed, 4-lobed spiniferous plates separated by regular longiseries of
papular areas; abactinal plates cruciform, connected in transverse direction by
oblong secondary ossicles, leaving medium-sized intervals; dorsolateral skeleton on
either side of the evident carinal series, broad, rather irregular; but sometimes poorly
defined longiseries of plates are evident; spines subacicular but strong, surrounded by
a wreath of pedicellariae; carinals with one to six spines; dorsolaterals with usually
one, marginals with two or three, and actinals with one to five spines; actinostome
small, but not sunken, two or three pairs of postoral adambulacrals being in contact
(pl. 71, fig. 3); papulae numerous; pedicellariae small, without outstanding characters;
the straight are small, lanceolate; the crossed, similar to those of Asterias ss.; gonads
open dorsally; size large; rays five to seven.

An isolated genus more nearly related to Asterias than to any group confined to
Southern Hemisphere. ;

1, PERISSASTERIAS POLYACANTHA Clark.
Perissasterias polyacantha CLARK, 1923, p. 307, pl. 18, fig. 3; 1926, p. 29, pl. 6—FisHER
1926, p. 199.
Cape Colony: Off Cape Town, 156 fathoms; off Dassen Island, 170 fathoms.
2. PERISSASTERIAS HEPTACTIS Clark.

Perissasterias heptactis CuarK, 1926, p. 26, pl. 5.

West from Lambert’s Bay, Cape Colony, 210 fathoms.
3. PERISSASTERIAS OBTUSISPINA Clark.

Perissasterias obtusispina Cuark, 1926, p. 28, pl. 7, figs, 1, 2.
Off Dassen Island, Cape Colony, 214 fathoms.
Genus SALIASTERIAS Koehler

Saliasterias Koruter, Austral. Ant. Exped., vol. 8, pt. 1, 1920, p. 52. Type, S. brachiata
Koehler —Fisuer, 1923, p. 603.

Diagnosis —Rays 9 or 10; skeleton irregularly reticulate; plates unequal and
without indication of carinal or dorsolateral longiseries; on sides of rays transverse
ares of plates, of which the lowest is the inferomarginal, correspond to two narrow
adambulacral plates and alternate with conspicuous, quadrangular membranous
spaces (two adambulacrals in length), containing three or four papulae; no actinals;
abactinal spinelets slender, cylindrical, in small groups, surround the membranous
skeletal intervals containing three or four papulae; inferomarginal spines two or
three identical with dorsals; adambulacrals with two long, slender, cylindrical, ter-
minally rounded or truncate spinelets; crossed pedicellariae, upward of 0.4 mm. long

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 239

and without enlarged terminal teeth, are scattered among the spinelets, rather
abundantly on rays; straight pedicellariae, lanceolate, upward of 0.7 mm. long,
less numerous.
1. SALIASTERIAS BRACHIATA Koehler,
Saliasterias brachiata Koruver, 1920, p. 54, pl. 11, figs. 1-4; pl. 12, figs. 1-8; pl. 58, fig. 3.—
Fisuer, 1923, p. 603.
Adelie Land, 15 to 20 fathoms; latitude 66° 50’ S.; longitude 142° 06’ E., 354

fathoms.
Genus SMILASTERIAS Sladen

Smilasterias Suapen (subgenus), Challenger Asteroidea, 1889, p. 562, 578. Type, Asterias
scalprifera Sladen.

Diagnosis —Small, 5-rayed forms with compact, reticulate abactinal and lateral
skeleton beset with small spinelets; inferomarginals on ventrolateral border of ray,
each with a prominent, oblique comb of three or four flattened truncate spines
which radiate apart and are without attached pedicellariae; adambulacral plates with
two or three slender spinelets, devoid of attached pedicellariae; a series of small spinif-
erous actinals in one species (scalprifera); adoral carina comprises first pair of post-
oral adambulacrals which are longer than second pair; crossed pedicellariae scattered,
not in circlets around spinelets; straight pedicellariae lanceolate, scattered on all
surfaces; in addition, sometimes, large spatulate unguiculate pedicellariae with
curved, interlocking tines; tube feet in four series; one madreporic body; gonads
open dorsally at upper margin of superomarginal plates, near interradius.

1. SMILASTERIAS SCALPRIFERA Sladen.
Asterias (Smilasterias) scalprifera Suapen, Challenger Asteroidea, 1889, p. 578, pl. 100,
figs. 4-6; pl. 103, figs. 1 and 2.
Off Marion and Heard Islands, 50 to 75 fathoms.
2. SMILASTERIAS TRIREMIS Sladen.
Asterias (Smilasterias) triremis Sutapen, Challenger Asteroidea, 1889, p. 578, pl. 101, figs.
5 and 6; pl. 102, figs. 5 and 6.
Between Kerguelen and Heard Islands, 150 fathoms.
3. SMILASTERIAS IRREGULARIS Clark.

Smilasterias irregularis Cuarx, Rec. South Austral. Mus., vol. 3, No. 4, 1928, p. 402, fig. 116.

South Australia: Spencer or St. Vincent Gulf.

This species has only two inferomarginal, flat, square-cut spines; no actinal
plates; adambulacrals diplacanthid.

Note.—Asterias (Smilasterias) terwieli Goldschmidt (Ann. Mag. Nat. Hist.,
ser. 9, vol. 14, 1924, p. 499) from off the coast of Chile not only is not a Smilasterias,
but does not even belong to the Forcipulata.

Genus SPORASTERIAS Perrier

Sporasterias Pernien, Expéd sci. Travailleur et Talisman, 1894, p. 107. Type, Asterias
rugispina Stimpson.—Lupwie, 1903.—Koruer, 1917, 1920, 1923.—Venninn, 1914.
« (Not Ludwig, 1905=<=A mpheraster).—Fisuen, 1922, p. 596; 1923. p. 605.
Parastichaster Korucer, Austral. Ant. Exp., 1920, p. 89. Type, P. mawsoni Koehler.

240 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Diagnosis—Small antarctic, or southern, paedophoric species with monacanthid
adambulacral plates and typically a well-developed, irregularly reticulate abactinal
skeleton, overlaid by a thin pulpy skin; carinals in more or less evident longiseries;
dorsolateral plates usually rather numerous, forming an irrgular net, or arranged in
subregular longiseries; superomarginals not relatively very broad; inferomarginal
plates and spines not forming a sharp ventrolateral border to ray; crossed pedicallariae
when numerous, aggregated around abactinal and marginal spinelets; straight pedi-
cellariae usually lanceolate, sometimes robust, ovoid, or subspatulate; gonads open
ventrally; a series of actinal plates, usually spiniferous, but absent in young specimens.

The relation of this group to Anasterias is discussed under that genus.

Parastichaster Koehler appears to represent the maximum development of
skeleton. In addition to the 6-rayed type, Doctor Koehler describes two other
species—directus having five rays and sphaerulatus with six or eight rays. These
are all stout-armed, monacanthid sea stars, having a rather irregular dorsolateral
skeleton, small carinals, and relatively much smaller superomarginals than has
Stichaster, to which Koehler compares the genus. The gonads open ventrally and
the young are carried in a cluster over the mouth. The stubby inferomarginal
spines are arranged in short, oblique, transverse series.

All these features are characteristic of ‘‘Sporasterias spirabilis” (Bell) and
adult ‘““S. rugispina”’ (Stimpson). In fact P. mawsoni suggests a 6-rayed Sporas-
terias spirabilis. The paedophoric habit and monacanthid adambulacrals are out-
standing common characters of Sporasterias and Parastichaster, and the genera seem
to be confluent.

1. SPORASTERIAS ANTARCTICA (Liitken).

Asteracanthion antarcticum LUTkeN, Vidensk. Meddel., 1856, p. 105.

Asterias rugispina Stimpson, Proe. Boston Soc. Nat. Hist., vol. 8, 1860, p. 267.—Lrrrotpr,
Zeit. Wiss. Zool., 1895, p. 563; literature.

Asterias cunninghami Perrine, Rév. Stell., 1875, p. 75.

Asterias spirabilis Bexu, Proce. Zool. Soc., 1881, p. 513, pl. 48, fig. 4.

Asterias verrilli Beuu, Proc. Zool. Soc., 1881, p. 513, pl. 47, figs. 3, 3a.

Asterias hyadesi Perrier, Comptes-rend., 1886, p. 1146.

Sporasterias spirabilis Perrier, Expéd. sei. Travailleur et Talisman, 1894, p. 107.

Asterias (Sporasterias) antarctica MrissNER, Arch. Naturges, 1896, p. 105, pl. 6, figs. 7,
7a-7b; literature.

Sporasterias antarclica Lupwia, Expéd. antaret. Belge, 1903, p. 39; 1905a, p. 70—Koruusr,
1917, p. 10; 1920, p. 78, pl. 18, figs. 1-4; pl. 28, figs. 1-4; 1923, p. 14, pl. 7, fig. 4.

Anasterias antarclica Fisuer, Ann. Mag. Nat. Hist., ser. 9, vol. 18, 1926, p. 197.

To this synonymy Leipoldt 1895, p. 563, adds: Asteracanthion varium Philippi,
A. fulgens Philippi, as well as several species, such as Anasterias minuta, Asterias
perriert Smith, Asterias rupicola Verrill, and Calvasterias stolidota Sladen, which
I consider to be distinct.

Among the above list of synonyms there are undoubtedly representatives of
several well-marked formae, but material is lacking by which a scheme can be
constructed.

Verrill (1914, p. 353), writes of this complex of forms as “the most common and
abundant littoral and shallow-water starfishes of both coasts of Tierra del Fuego
and Magellan Strait.” i

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS~—FISHER 241

2. SPORASTERIAS PERRIERI (Smith).
Aaterias perrieri Suara, Ann. Mag. Nat. Hist., ser. 4, vol. 17, 1876, p. 106.

Kerguelen.

3. SPORASTERIAS RUPICOLA (Verrill).
Asterias rupicola Verrity, Bull. U. 8. Nat. Mus., vol. 3, 1881.

Kerguelen.

4. SPORASTERIAS MAWSONI (Koehler).
Parastichaster mawsoni Korucer, Austral. Ant. Exp., 1911-1914, vol. 8, pt. 1, 1920, p. 91,
pl. 19, figs. 1-8; pl. 20, fig. 1; pl. 21, figs. 1-6; pl. 23, fig. 4; pl. 24, fig. 5; pl. 28, fig. 3;
pl. 63, fig. 2.
Macquarie Island.
5. SPORASTERIAS DIRECTA (Koehler).
Parastichaster directus Korner, 1920, p. 97, pl. 20, figs. 8-11; pl. 21, figs. 8-12; pl. 23,
figs. 1, 2; pl. 62, fig. 2.
Macquarie Island.
6 SPORASTERIAS SPHAERULATA (Koehler).
Parastichaster sphaerulatus Korner, 1920, p. 101, pl. 21, fig. 7; pl. 23, figs. 5-10; pl. 24,
figs. 1-4; pl. 63, fig. 3; pl. 64, figs. 1,2. *
Macquarie Island.

Genus STICHASTER Miiller and Troschel

Stichaster Mttuter and Troscner, Monatsber. Akad. Wiss. Berlin, 1840, p. 102; Arch.
Naturges., Apr. 1840, p.321. Type, Asterias aurantiaca Meyen (not Linnaeus) = Stichas-
ter striatus Miiller and Troschel.

Tonia Gray, Ann. and Mag. Nat. Hist., Nov. 1840, p. 180. Type, Tonia atlantica Gray =
Stichaster striatus Miller and Troschel—Perrier, Expéd. sci. Travailleur et Talisman,
1894, p. 129.

Coelasterias Venniti, Trans. Conn. Acad., vol. 1, 1871 (1867), pt. 2, p. 247. Type, Coel-
asterias australis Verrill. See Fisher, 1922, p. 598.

Diagnosis —Body covered with close-set, coarse, granuliform spinelets; carinal
and superomarginal plates very short but broad; the latter conspicuously broader
than inferomarginals which form a well-marked ventrolateral angle to ray and
sometimes overhang the single series of small actinal plates; dorsolateral area broad,
the plates in three to five series on each side of the carinals; adambulacral plates
short, with two crowded spines, devoid of pedicellariae; crossed pedicellariae scat-
tered, not in clusters or wreaths around abactinal spines; one madreporic body;
interbrachial septa heavily calcified; gonoducts open dorsally, those of male with
proximal swelling; not fissiparous.

1. STICHASTER STRIATUS Miller and Troschel.
Asterias aurantiaca Mrven, Reise um die Erde, vol, 1, 1834, p, 222 (not Linnaeus).
Stichaster striatus MULitEeR and Troscuet, 1840b, p. 321.—Vernity, 1914, p. 362.
Tonia atlantica Gray, 1840, p. 180; Synopsis, 1866, p. 2.
Asteracanthion aurantiacus Mtvuer and Troscuet, Syst. der Asteriden, 1842, p. 21.

Stichaster avrantiacus Vennitt, 1867a, p. 2903.—Crark, Bull. Mus. Comp Zod, vol. 52,
1910, p. 337, pl. 8, fig. 1.

West coast of South America from Callao, Peru, to Taleahuano, Chile.

242 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

2. STICHASTER AUSTRALIS Verrill.

Coelasterias australis Verritu, Trans. Conn. Acad., vol. 1, 1871 (1867), pt. 2, p. 247.—
Hvurron, Catalog Echin. New Zealand, 1872, p. 5.—Koru er, 1920, pp. 54, 91.

Stichaster australis SuapEN, 1889, p. 431.—FarquHar, Trans. New Zeal. Inst., vol. 27,
1895, p. 202; Proc. Linn. Soc. New South Wales, 1898, p. 313.—FisHeEr, 1922, p. 598.—
Mortensen, 1925, p. 313, pl. 14, figs. 1, 2; text figs. 17, 18.

New Zealand.
Genus UNIOPHORA Gray

Uniophora Gray, Ann. and Mag. Nat. Hist., Dec. 1840, p. 288. Type, Uniephora globifera
Gray = Asterias granifera Lamarck.

Diagnosis.—Typically 5-rayed, coarse-spined, heavy-ossicled, diplacanthid Aus-
tralian forms with one to three series of actinal plates; abactinal skeleton with large,
more or less irregular, meshes in two dorsolateral series on either side of the irregular
carinal series; abactinal spines spaced, coarse, tuberculate, often capitate or globose
and prominent; superomarginals with one to several capitate tubercular spines;
inferomarginals with one to three spines; one well-defined, usually irregular, series
of dorsolateral plates; adoral carina short, composed of one or two pairs of contiguous
adambulacral plates, the first pair conspicuously longer than the second; numerous
small, crossed pedicellariae are scattered along the skeletal ridges surrounding papular
areas; straight pedicellariae inconspicuous, broadly lanceolate.

Some of the nine species listed below will ultimately, I think, be reduced to
formae. Five species are from the South Australian coast, namely, gymnota,
multispina, nuda, obesa, and uniserialis.

1. UNIOPHORA GRANIFERA (Lamarck).

Asterias granifera LaMARcK, Anim. sans verteb., vol. 2, 1816, p. 560.

Uniophora globifera Gray, Ann. and Mag. Nat. Hist., Dec. 1840, p. 288.

Uniophora granifera Brwu, Proc. Zool. Soc. London, 1881, p. 497.—FisHmer, Ann. and Mag.
Nat. Hist., ser. 9, vol. 18, 1926, p. 198.—Cxiarx, Rec. South Austral. Mus., vol. 3,
No. 4, 1928, p. 403, fig. 117.

Tasmania.

I have examined Gray’s types in the British Museum, and a specimen in the
Paris Museum mentioned by Perrier (1875, p. 77), now labeled ‘“ Pisaster graniferus—
Mers Australes, MM. Peron et Lesueur, 1803.” I have two examples (one with
six rays) from Port Jackson, New South Wales (U. S. National Museum).

2. UNIOPHORA DYSCRITA Clark.
Uniophora dyscrita Cuark, Journ. Linn. Soc. Zool., vol. 35, 1923, p. 244.

Western Australia: Garden Island, near Freemantle.

3. UNIOPHORA FUNGIFERA (Perrier).

Asterias fungifera Perrimr, Réy. Stell., 1875, p. 73.

Uniophora fungifera Fisuer, Ann. and Mag. Nat. Hist., ser. 9, vol. 18, p. 198.—CLARK,
1928, p. 416.

I have examined the type, in the Museum d’Histoire Naturelle. It is labeled
“Nouvelle Hollande, 1844, M. J. Verreaux.”’
4. UNIOPHORA GYMNOTA Clark.
Uniophora gymnota CuarK, Rec. South Austral. Mus., vol. 3, no. 4, 1928, p. 405, fig. 118.
South Australia: Spencer or St. Vincent Gulf.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 243

5. UNIOPHORA MULTISPINA Clark.
Uniophora mullispina Cuark, Rec. South Austral. Mus., vol. 3, no. 4, 1928, p. 407, figs. 1194,
119d.

South Australia: Near Adelaide.
6. UNIOPHORA NUDA (Perrier).
Asterias nuda Perrier, Rév. Stell., 1875, p. 71.
Uniophora nuda Fisuer, Ann. and Mag. Nat. Hist., ser. 9, vol. 18, 1926, p. 198.—Crarx,
1928, p. 417.

I have examined the type which is from Port Lincoln, South Australia, R. B.
Harvey, Esq. (British Museum); Perrier erroneously says that it is from Torres
Strait.

7. UNIOPHORA OBESA Clark.
Uniophora obesa CiarK, Rec. South Austral. Mus., vol. 3, No. 4, 1928, p. 409, fig. 120.

South Australia: Rocky Point, Eastern Cove, Kangaroo Island.
8. UNIOPHORA SINUSOIDA (Perrier).
Asterias sinusoida Perrier, Réy. Stell., 1875, p. 74.
Uniophora sinusoida Fisner, Ann. and Mag. Nat. Hist., ser. 9, vol. 18, p. 198.—Cvanrk,
Rec. South Austral. Mus., vol. 3, No. 4, 1928, p. 411, figs. 121la, 121b.
I have examined the type, in the British Museum, from Hobart, Tasmania.
This may be a slender-spined form of fungifera. *
9. UNIOPHORA UNISERIALIS Clark.
Uniophora uniserialis CLark, Rec. South Austral. Mus., vol. 3, No. 4, 1928, p. 413, figs.
122a, 122b.

South Australia: St. Vincent Gulf.

Subfamily NOTASTERIINAE Fisher
Genus NOTASTERIAS Koehler

Notasterias Koruer, British Antaretic Exp., vol. 2, pt. 4, 1911, p. 35. Type, Notasterias
armata Koehler.—Fisuer, 1923, p. 258.

Diagnosis —With crossed pedicellariae of a peculiar type, sometimes of very
large size, having hooked, terminally crossed jaws and usually broad base (macro-
cephalous Koehler) in addition to ordinary crossed and straight pedicellariae (includ-
ing a peculiar large pirocephalous variety). Dorsal skeleton consisting of large,
cruciform, spiniferous carinals (alternating sometimes with secondary smaller plates),
to which short, transverse, sometimes sinuous, dorsolateral arcs of a few plates
connect the superomarginals; a sporadically spiniferous plate in these ares represents
a poorly developed series of primary dorsolaterals. Marginals large, cruciform;
one series of spiniferous actinal plates; two dorsolateral series of conspicuous papular
areas on either side of carinal plates; one intermarginal; marginal and abactinal
spines conspicuous, well spaced, not more than one to a plate, in longiseries, and
devoid of a sheath carrying a collar of crossed pedicellariae; adambulacral plates
diplacanthid proximally and monacanthid distally; or diplacanthid throughout,
the spines without attached pedicellariae.

244 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

1. NOTASTERIAS ARMATA Koehler.
Notasterias armata Koruurr, 1911, p. 39, pl. 5, figs. 6-11; pl. 6, figs. 1-8; 1912, p. 25, ple i
fig. 8; 1920, pl. 7, figs. 7-9; pl. 8, figs. 2-8; pl. 9, figs. 1-6; pl. 10, figs. 1-9; pl. 60, fig. 1;
pl. 61, fig. 1.
South Victoria Land, 10 to 18 fathoms; Emperor Alexander Land, 297 meters;
latitude 64° 32’ to 66° 55’ S.; longitude 92° to 145° 21’ E.; 60 to 354 fathoms.
2. NOTASTERIAS HASWELLI Koehler.
Notasterias haswelli Koruupr, Australasian Anarctic Exped., vol. 8, pt. 1, 1920, p. 70,
pl. 1, fig. 11; pl. 6, fig. 4; pl. 7, figs. 1-7; pl. 9, fig. 7; pl. 62, fig. 1.
Latitude 65° 42’ to 66° 50’ S.; longitude 92° 10’ to 142° 6’ E.; 60 to 354 fathoms.

Genus AUTASTERIAS Koehler

Autasterias Korater, British Antarctic Exp., vol. 2, pt. 4, 1911, p. 35. Type, Asterias
pedicellaris Koehler.—FisHEr, 1923, p. 258.
Diagnosis.—In general structure similar to Notasterias but with monacanthid
adambulacral plates.
1. AUTASTERIAS PEDICELLARIS (Koehler).
Asterias pedicellaris Konuier, Trans. Roy. Soc. Edinburgh, vol. 46, pt. 3, 1908, p. 577;
pl. 7, figs. 61-67; pl. 8, figs. 74-78.
Latitude 71° 22’ S.; longitude 16° 34’ W.; 1,410 fathoms.
2. AUTASTERIAS BONGRAINI (Koehler).
Autasterias bongraini Konner, Deux. Expéd. Ant. Frangaise Echinod., 1912, p. 26, pl. 2,
figs. 10, 11; 1920, p. 74, pl. 6, figs. 5-10; pl. 59, fig. 1.
South Shetland Islands, 420 meters; latitude 65° 42’ to 66° 55’ S.; longitude
92° 10’ to 145° 21’ E.; 60 to 318 fathoms.

LIST OF THE SPECIES NAMES APPLIED TO ASTERIINAE AND NOTASTERIINAE
OF THE SOUTHERN HEMISPHERE

In the first column is given the species and original describer; in the second
column, the original genus and date; in the third column, the genus to which the
species has been assigned in the foregoing summary. Valid species names are in
roman; synonyms in italics.

adelisesKochler22 24.) eee Anasterias 1902022 220 Soe sees eee Lysasterias.
Glo Belle Sees ares eee ee Asterias; 88s. 28245 2 eee ee Diplasterias.
antarcticum Liitken==.=..=-.2 2... Asteracanthion, 18562-22252) sees Sporasterias.
antarchica Koehler == ae to Pisasteryl9Na32- 2. oe eee Se Eremasterias.
SnMa ta ORNICR es eee ene eee Motastenrias ss LOiie =e, eee er Notasterias.
OaniCcOnGTa ye See ee eee PONE, (Sa OS ee ee ee ee Stichaster.
asterinoides Perrier_.-2-_-2---__.-.-1 Calvastertas: Sip. 2-46 -ee ee « Calvasterias
AntipsdumiBell se Sas Fe as Fae Calvasterias, 18822202 0150 Sp bees Calvasterias.
aurantiaca Meyen_-.-=-.-----=----.- Asterias;:U8342):)e beers ways Stichaster.
SISOTALA Over oes ee Pe gets @oelasteriass (le 7die= tee eee Stichaster.
DELS LUC er sac eee Se eae Cia ee Asterias. 1884s pe oe 8 ees Diplasterias.
beletcre Dud wits==- 22 seo ee a Anasterias; 1903S--22 5s eee aoe ee Lysasterias.
Dikerzae Oper ese eee ee ee Granaster*US0Ge seo ee. aes Granaster.

bongrain) Koehlors=2 9s] s—e ee = Autasterias 1OTQUNI eg aimee de Autasterias.
Dborbonica eetmiene sense ee eee Asterias Ji875<- <2 2-4 5- eee Sporasterias.
brachiata’ Koehlers--. ose e-- Salissterias) 19202—2— eee Saliasterias.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER

brackiste’ Rother... 206 sence nenscns Cryptasterias,; 1923.....-..~....--..-
OTOANNCH IN DONG Ces ashore cheer rr me RM GTIED, (RRB eh rere sth Abeer
bruce Haile a. ce eee Stolssteriae, 1908. 3. see
COpennie Verrier l. . oer eee OI CRP INI TIE 10S Setar
chirophora Ludwig--.-......-..------ RUMECONAGPROUGs. ce so te retece ee
dasaluet Philippi-_.-...----.------- Asteracanthion, 1870... .~.....~- ee
Cotiferte Mosher. oi wee eee Kalyptasterias, 1923_............-...
cupulifera Ludwig........-..-.--.--- Anepterias 10062 wei he a
cunninghami Perrier_-- ~~ ----.--~---- EGEIUD, At ae crete eee enw
Ci Coamastering, 1016... ..-....---.-.--
TUNGIIMONEIES O. cwasacecnusodvaucne Uniophdr, 1000 |... <2 ees
direseuneoenhlere eee Farastichaster, "19205. 2. .c.. ee Se
JRE TOREOR Ss oo can akacccenccecs WCUIMEUET, (ROO oe Lc deuce de cece eth
ToChWeneridel. 6 csss2-2encaedcene POUMNVENIN) 19SUL hci oe
STG Th Te ue Sopa eet Cite ae pas See ewceeerend
Julgene wamipple. — ee Asteracanthion, 1870. -..............
Yaaserniipnte. 2 ~ eee eee Asteracanthion, 1870_.......-.......-
fungifera Pemer. <..-..-...----- BARGIIRG, CON One hewn eet ceed
Poy 22) rE ARUP ECR ceri enna
Berman! Piitppt_ el Asteracanthion, 1858. ..__.........--
PO es ARTGNAN, DBR O Oe ocr einiinla ewe
OUT NOY Foe ne etc enk UmoploraISsG. = 2 ee
granifera Lamarck - ~~~ .__..-..------ ARIE, NOLO M i tr errr wirnia
EV UNTIONY COON tao ne errs oo ISTRATION CNSR yea ee owe
TREAD ROO S oerSt hrercnt Dep istatine 1870 oc Reese
Hisswellt Benin. 65 ee nee NO Crea DRR TAR LO Sc hoteles
ebessverslet se Stephanasterias, 1915_............_--
beptactis Glavki) 22 Perissasterias; 192652. ~.=~ 2. =e
RUM ID errr rrr A RURT ERS UAL ita ere eee
inaequalis’ Koehler... .......-..-.... Allostichaster, 1923.................-
SriGitGaRDeMIOrGs oo oo nee ce (DIpISCGrIe W008. wai cece
BROVGE DO Fee e ete cokucca PRURTERE TOUNG Gee cn cette ke aoe
irregitiecty Cleves ooo Joo ee PUDUENURMOO AOSD oo eee tek
insignis Werqulier =... wc eden PIB UMNUTOINUONS ink ec cccneekces
ORT RRND i wie eee PROOASOTINS NN O20 | eee eee
[RGUOE GAS St en eee J eC a ee
oh ea apa | 2 RMORTINE RO e ons cane wceneeceuee
uords Warqohiar 5. J... cee Siichastery IS0Gss So ce el
(ORE PORTION en eee DIP NOLIN IRON ere kie nec kecieecuas
oe A Asteracanthion, 1858_..............--
ee ee Diplasterias) 204 So. oi ccc ccc.
On || a SE AYGSCNOR. Oto aati oe
meridionalis Perrier_...............-- BNCAHOS, BGtan ene pene leas otk
RC OMS oe oo woe eo eke Parastichastar, 1020-0. - 25. 2... 2- =.
WN EMER ascent ect el. Wnseleriae, lee ae seen tees
pea tee ence cae Asteracanthion, 1870..............--
WEN INI ee eC eee Asterias, 1884............. te oie
multispina Clark. ____ eee aeons Uniopnores twas ces eee
nautarum Bell (Sladen) ____......_._- fey yi i a ae i
BR gl a a ARDRITEN, tES Lert tee” ce nen wk con
neozealanica Farquhar... _..._.__- ye ah Se oe ee
Cities PMN ate ek oe eek PEA Ait he not enema a
SNR PRU se nc caecnas nec act na ace Hesueey REUD. ee noo. Sa
octoradiata Koehler. ................ PANES A Mie i dak ess
ROI RR cs chests ean PP oh eared avg CACORE een ta

Diplasterias.
Diplasterias.
Cosmasterias.
Lyeasterias.
Cosmasterias.
Kalyptasterias.
Lysasterias.
Sporasterias.
Cosmasterias.
Uniophora.
Sporasterias.
Cosmasterias.
Diplasterias.
See p. 217.
Sporasterias.
Cosmasterias.
Uniophora.
Neosmilaster.
Cosmasterias.
Diplasterias.
Uniophora.
Uniophora.
Uniophora.
Allostichaster.
Notasterias.
Allostichaster.
Perissasterias.
Sporasterias.

Lysasterias.
Lysasterias.
Calvasterias.
Calvasterias.
Diplasterias..
Cosmasterias.
Diplasterias.
Lysasterias.
Diplasterias.
Sporasterias.
Anasterias.
Cosmasterias.
Diplasterias.
Uniophora.
See p. 24.
Diplasterias.
Allostichaster.
Uniophora.
Granaster.
Koehleraster.
Uniophora.

245

246 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

obtusispina Clark------------------- Perissasterias, 1926__---_____=----_-_ Perissasterias.
obtusispinosa Bell_------------------ PAS VOTERS ipl coco tee Diplasterias.
papillosum Perrier_.----------------- Asterodenman Sole ==. == ee === anaes Anasterias ?.
papillosa Koehler. .----------------- Diplasterias; 906" == — = “= === ea Adelasterias.
pedicellaris Koehler- - --------------- ARTEMAS WlO00==s= === == === ae Autasterias.
pedicellaris Koehler----------------- SpovasterigsyeL923= ee ee Anasterias.
ErricniSLO tee ee Asterias dis Geeeee o-oo eee Sporasterias.
perrier studer- 8 s-e- esos n= ee AnasteriassilS85"--- = 52 Lysasterias.
philipprt Beles = ea ae Asterina (SS lee 8) = ae ee See p. 62.
polygrammus Sladen- - -------------- DCHASten I SSOn a= = = ee ee Cosmasterias.
polyplax Miiller and Troschel_-------- Asteracanthion, 1844__--------_-____ Allostichaster.
polyacantha! Clank. 42 2225) -=-=2—52—5 IPETiISsaStGLIAS OLS = ee Perissasterias.
TAdIatas Koehler. 2-- a2-==--55es=2e==— Cosmasterias; e023 = 5ee= === eae See p. 216.
TOPULATIO = Sees Seek BAe ae Allostichaster ml Q2S ===) eae Allostichaster.
rugispina Stimpson..-~-------------- ANSTOTIOS 9a 8 G0 eee ae ee Sporasterias.
TUpDICOIANV CrmUGS seen eee IAS TOnISS SL Os eee ee Sporasterias.
scalprifera Sladen_-.-..------------- DIOUASTETIAs WIS8Oe 2 eo eee Smilasterias.
BINUSOIGa emer s2— = on eee ae ae Asteriss lS (020-2 ae eee Uniophora.
spEectaurilenebilip pine -n- aaa =e Asteracanthionwil@/Os 22 o-===——=—= = =e Cosmastefias.
sphaerulatus Koehler_-_-------------- PanastichastenwLO2022 2-2 = eee Sporasterias.
SprrabipesBelle ss so- See oe Asteriag (188i —— eee Sporasterias.
BpiNnOssPheITioNe es eee a= Diplasterias) $91 -- -- =" 3 5=-= eee See p. 216.
BteInenitonldel: 2 a -— == == Asterias: [8852<% —2 522-5 Sse Neosmilaster.
stolidota;siaden ~~ ===. 2) 22.22 ess G@alvasterias; 1889.-=.---=- == === Calvasterias.
striatus Miiller and Troschel__-------- Stichaster, 918404. = = Stichaster.
StudenpBelleee sen as ee a ee ees Asterias 88h. -22- 22-55-% = See Diplasterias.
atudensRernien*? =. 3. 22-2 ae Anasterias, Wi 89lle= = = pe Lysasterias.
studendeploniola--——- =. so oe Gastraster 19045." = =* eee Cosmasterias.
eulcifengherrierenea= = a ae ae Asteracanthion, 18692. === = Cosmasterias.
suteriudewonolee =" sae aoe nea Stichaster,,1894-—_..- = eee Calvasterias.
TeneraniOehlerset= = 322 ae eee Anasterias19063- =.=. - 2-2 eee Lysasterias.
terwieli Goldschmidt._—-.-_-.-----_-- Smildsterias sl924= 2. =). eee ee See p. 217.
tomidataiSladenes = -=—-3-42-- === === @osmastenas 8895 o— = eee eee Cosmasterias.
triremisipladent===22 2-2 ee Suiilasterias) s889-_-- 25- -es eeee ee Smilasterias.
turquetinioebler==— —- 25222222252 = Diplasterias w906: = -2=- = =e eee Cryptasterias.
uniserialisGlarks-_=-- = = 2 eee Uniophora 9282". 2 ee eee Uniophora.
varimMye Dp Plos =n ee ee aa oe Asteracanthion, 1870_---.----.-=---=- Sporasterias.
verrillap Bellet Se ee ere Ae Asterias, 1882s 223. 2s Sa ee Sporasterias.
vesicwlosarsladen= = = 2--.-=-.s--22- Asterias S89 ien- = 2 2 eee Bathyasterias.
victorsae woehler=— =. 2 = 2--=- == Coscinastenas aL Olas see Diplasterias.

victoriserhoehlon. = 0 222-2 seen5=aess Amasterias 920%. =. eo eee eee Lysasterias.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 247

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© Complete list of tithes relating to ses stars.

250 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Fisner, Wavter K.—Continued.
1916. Notes on the Systematic Position of Certain Genera and Higher Groups of Starfishes.
Proc. Biol. Soc. Washington, vol. 29, pp. 1-6. January 25. ;
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19-23. May.
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August.
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1-5, pls. 1-2. December.
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Forses, Epwarp.
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Goro, SEITARO.
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Univ. Tokyo, vol. 29, art. 1, 808 pp., 19 pls.

ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 251

Gray, Joun Eowarp.
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pls. 4-8.

252 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Korner, René.—Continued.
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ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS——FISHER 253

Mortensen, Tu.—Continued.
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63160—30——17

254 BULLETIN 76, UNITED STATES NATIONAL MUSEUM

Suapen, W. Percy.

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ASTEROIDEA OF NORTH PACIFIC AND ADJACENT WATERS—FISHER 255

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EXPLANATION OF PLATES

PuatTe 1
Fiaurp 1.
2s

Asterias amurensis, cotype; Amur country, eastern Asia; actinal surface, enlarged.
3.

Same specimen; portion of abactinal surface, enlarged.
Same, Petropavlovsk, Kamchatka; portion of abactinal surface, natural size, alcoholic
specimen.

256

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S AMURENSIS

L E PAGE FACING.

BULLETIN 76, PART 3 PLATE 2

| Pua 2
muren: , station 3242, same specimen as Figure 3} portion of actinal surface -
ne, No. 68, Norton Sound, Alaska, %%.

- - Eeete station, ae 4. Specimen upon which description is based.
a 257

Figure 1.
2,

a

. Same, Petropavlovsk, Kamchatka.
. Same, No. 7631, Norton Sound, Alaska.

Pate 3

Asterias amurensis forma anomala, station 3231; actinal surface, reduced.
Asterias amurensis forma amurensis, Petropavloysk, Kamchatka; portion of actinal s
face, alcoholic specimen. ;

Same, station ae actinal surface of Plate 2, Figure 1.

s, Saco iannl
258

PART 3 PLATE 4

76,

BULLETIN

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Piate 4

Ficure 1. Asterias amurensis forma anomala, station 3231; portion of actinal surface, enlarged.
2. Same, station 3231; abactinal surface.
3. Same, station 3231; abactinal surface of specimen with well-defined series of carinal
spines.
4. Asterias amurensis forma amurensis, station 3335, 93 fathoms, alcoholic specimen.
5. Same specimen, abactinal surface.
6. Same, station 3243; medium-sized specimen, with dorsal skeleton cleaned and a portion

removed to show actinostomial ring; 0, odontophore.
259

PuaTEe 5 _ + ae
Specimens to show growth stages; X20 a *G iY
Noe PS eteh a te y
Ficures 1-3. Station 3241. oe ae '
4-6. Petropavlovsk, Kamchatka. ; a
Nore meoure specimens are easily mistaken for Leptasterins

260

BULLETIN 76, PART 3 PLATE 6

2-8, 10, 11. A. AMURENSIS AND FORMAE

1, 9. ASTERIAS AMURENSIS ROLLESTONI

Piate 6

Figure 1, Asterias amurensis rollestoni, Mororan, Hokkaido. Abactinal crossed pedicellaria, 0.21

mm., 100.

2. Asterias amurensis forma amurensis, cotype, Amur. From an abactinal spine, 100.

2a, 2b. From inferomarginal spines, 0.26 mm., 100.

3. Same, from inferomarginal spine of a Petropavlovsk specimen, with R 87 mm.; pedi-
cellaria 0.32 mm., 100.

3a. From inferomarginal spine of largest Petropavlovsk specimen (R 130 mm.), » 100.

4. Same, from inferomarginal spine of Norton Sound specimen, 0.27 mm., 100.

5. Asterias amurensis forma anomala, station 3231; from an abactinal spine, 0.27 mm.,
100,

5a. Station 3231; from an inferomarginal spine, 0.36 mm., 100.

6. Same, Cape Lisburne specimen; from an inferomarginal spine, 0.3 mm., » 100.

7. Asterias amurensis forma amurensis, station 3225, 85 fathoms; from an inferomarginal
spine, 0.34 mm., * 100.

8. Same, station 3335, 90 fathoms; from inferiomarginal spine, 0.36 mm., 100.

9. Asterias amurensis rollestoni, Mororan, Hokkaido. Abactinal straight pedicellariae, 0.5
mm., 50.

9a. From actinal surface, 0.72 mm., 50.

9b. From a furrow spine, 0.45 mm., 100.

10, Asterias amurensis forma amurensis, cotype, Amur; a, an intermarginal straight pedi-
cellaria, 0.75 mm., 50; 6, c, the tips of two jaws, 100.

11. Asterias amurensis forma anomala, station 3231; a segment across actinal surface, on
basal third of ray; S, superomarginals; J, inferomarginals; a, a, a, three adambula-
cral plates. .

261

PLATE 7

Ficure 1. Asterias amurensis forma amurensis, Norton Sound; a straight pedicellaria from a furrow
spine, 0.67 mm., 100.

la. An abactinal pedicellaria, 0.43 mm., > 100.
2. Asterias amurensis forma anomala, station 3231; an actinal pedicellaria, 0.76 mm., 50.
2a. Tip of an actinal pedicellaria, > 100. ;
2b. A pedicellaria from a furrow spine, 100.
2c. Two abactinal spinelets, upper 1.25 mm., lower 1.08 mm., * 15.
2d. Two inferomarginal spines, 2 mm., 15.
3. Asterias amurensis forma amurensis, station 3250, 17.5 fathoms; actinal pedicellaria, < 50.
4. Forma anomala, Cape Lisburne, Arctic Ocean; a, an actinal pedicellaria, 0.9 mm., 50;
b, tip of one jaw; ec, tip of two jaws, 100.

5. Asterias amurensis forma amurensis, cotype, Amur Province. Abactinal spinelet, 1.2

mm., X25.
5a. Adambulacral armature from furrow face, 10.
6. Same, Petropavlovsk, largest specimen, R 130 mm.; armature of three consecutive
adambulacral plates near the base of ray, 5.
Same, station 3242, two consecutive adambulacral plates, 5. This is the specimen
upon which the account is based.
8. Same, specimen from station 3335, 93 fathoms; adambulacral armature and an infero-
marginal spine, J, <5.
9. Same, Petropavloysk, an actinal straight pedicellaria, 0.76 mm., 50; and tip of another,
100.
10. Same, station 3225, 85 fathoms; an actinal straight pedicellaria, 0.63 mm., 50.
10a. A tip, X150.

“I

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 7

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 8

1-3. LEPTASTERIAS GROENLANDICA

4. L. MULLERI. 5. L. ARCTICA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Puate §

Fiaure 1. Leptasterias groenlandica; an abactinal crossed pedicellaria of forma cribraria, station
$252, 0.21 mm., * 200.

la. Station 3537, end of a blade, 200.

1b. Station 3506, forma groenlandica; end of a crossed pedicellaria, 200,

le. Station 3506, forma groenlandica; a very large axillary straight pedicellaria, 1.20 mm., * 50

ld. Station 3352, forma cribraria; an intermarginal straight pedicellaria and an inside view
of a jaw, 0.8 mm., “50.

le. Jaw of the slender lanceolate sort, 50.

2. Leptasterias groenlandica forma cribraria, station 3252, abactinal spinelet, 0.52 mm., «50

2a, Station 3530, abactinal spinelet, 0.55 mm., 50.

2b. Station 3506, two abactinal spines (forma groenlandica) with pedicellaria drawn to scale,
longer spine 0.6 mm., 50.

3. Leptasterias groenlandica, off Cape York, west coast of Greenland; an abactinal crossed
pedicellaria, 0.225 mm., 200. Similarly shaped adambulacral spine pedicellariae
are 0.27 mm. long.

3a. Same, Jugor Strait, off Kara Sea, north of Siberia. An abactinal pedicellaria,
0.2 mm., 200.

4. Leplasterias milleri, northwest of Bergen, 90-200 fathoms, slender rays; abactinal
pedicellaria, Riksmuseum No. 751, » 100.

da. Same, two jaws, Waideguba, 40 fathoms, Riksmuseum, No. 745, 200.

4b. Same, side view of jaw, northwest of Bergen, Riksmuseum, No. 751, 200.

5. Leplasterias arctica forma arctica, cotype, Point Franklin; jaw of abactinal pedicellaria,
200.

5a. Same (0.26 mm.), 200.

5b. Same, 100.

5c. Same, Plover Bay (0.23 mm.), * 100.

5d. Same, 200.

5e. Same, specimen from station 3559, intermediate between forma arctica and forma

beringensis, length 0.27 mm., 200.
5f. L. arctica forma beringensis, type, station 3277, abactinal, 200.
5g. Same, station 3294, 0.27 mm., 100.

5h. Same, station 3294, outside of jaw, side view, 200.
263

PuaTE 9
Leptasterias arctica

Ficure 1. Station 3294, forma beringensis, abactinal skeleton, external view, <5. The lowermost
row of plates (c-c) are the carinals; AD, direction of disk.

Specimen from northern Bering Sea, forma arctica; abactinal skeleton from coelomic
side; c—c, carinal series; X10.

3. Station 3504, forma beringensis; abactinal skeleton, from coelomic side. This specimen

has a heavier skeleton than Figure 2; c-c carinal plates; 10.

4. Station 3536, forma beringensis; specimen with robust rays. Small portion of abactinal
surface showing carinal spines, <5. ; ;

Station 3512, forma beringensis; odontophore from above; J, interradial septum; ambu-
lacral ossicles on either side; < 10.

Station 3536, forma beringensis; adambulacral, actinal, and inferomarginal spines; three
consecutive adambulacral series are shown; AD, adambulacral; AC, actinal; J,
inferomarginal; >< 10.

bo

oa

oS

7. Forma arctica, straight pedicellaria from a suboral spine, cotype, X50.
8. Forma arctica, an unguiculate pedicellaria from actinal interradial region, cotype, 50.

=

). Station 3543, forma beringensis; two views of an actinal interradial or axillary pedicel-
laria, X25.

10, 11. Station 3543, forma beringensis; two unguiculate abactinal pedicellariae, the larger
0.9 mm. long, >< 50.

12. Station 35438, forma beringensis; a straight pedicellaria from furrow face of adambulacral

plate, < 100.

264

BULLETIN 76, PART 3 PLATE 10

5 WA day,
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N

1lb

1-4. LEPTASTERIAS ARCTICA. 5-11. L. HYLODES

FOR EXPLANATION OF PLATE SEE PAGE FACING.

° Piate 10

Figure 1. Leptasterias arctica forma beringensis, station 3514, abactinal RE “200. Ila
Same, Nazan Bay, Atka, 200,

2, 2a. Leptasterias arctica forma arctica, cotype, Point Franklin, 10 pedicellariae from
furrow spines 0.26 mm., LOO,

3. Leplasterias arctica forma beringensis, station 3294, pedicellaria from furrow spine, 0.27
mam., * 100.

83a. Same, jaw, 200.

4. Leptasterias arctica forma beringensis; young from brood “pouch”’ of female, showing
attachment to common central lobulate mass. Numerous strands without young
are shown issuing from the mass.

5. Leplasterias hylodes, station 2847, abactinal pedicellaria from large specimen, length
0.275 mm., 100.

5a. Same, one jaw, 200.

5b. Same, tip of jaw, 200.

6. Same, dwarf form, station 2856, abactinal pedicellaria, > 100.

6a. A jaw, 200.

7. Same, station 2856; jaw of a furrow-spine pedicellaria, 200.

7a. Station 2856; an adambulacral pedicellaria similar to the abactinal, 200.

8. Same, type (station 2851), an abactinal pedicellaria, 100,

9, Same, station 2847; an axillary unguiculate pedicellaria from actinal interradial region,
1.15 mm., X25.

10. Same, type (station 2851); a dorsal spine, 0.98 mm., 50.

11. Same, station 2856; dwarf form, abactinal spinelet, 0.76 mm., 50.

lla, Same specimen, a superomarginal spine, % 50.

115. Same, a furrow spine, 50.

265

PuateE 11 °

Ficure 1. Leptasterias hyperborea, Icefjord, Spitzbergen, abactinal pedicellaria, 100.
la. One jaw, Riksmuseum, No. 124, 200.
1b. Same, Cross Bay, Spitzbergen; jaw of abactinal pedicellaria, Riksmuseum, No. 330,
200.
le. Same, abactinal pedicellaria, Riksmuseum, No. 726, 100.
ld, le. Ieefjord, Spitzbergen, Riksmuseum, No. 124, *50; 1d, marginal 0.75 mm.; le, abac-
tinal, 0.67 mm.
2. Leplasterias orientalis, station 5024, abactinal crossed pedicellaria, 0.24 mm., 200.
2a. Same, station 4810 (see text).
. Closely related race from station 5047, off Bay of Sendai (see text).
. Orientalis, station 5024; abactinal pedicellaria, 0.27 mm., 100.
. Same, Okhotsk Sea, 57° 36’ N., 140° 34 E.; abactinal pedicellaria, 0.31 mm.
. Same, station 5024; straight pedicellaria 0.8 mm., 50.
2g. Same, jaws of two straight pedicellariae from actinal interradial area of example
upon which description is based, X50.
2h. Same, station 5024, carinal spinelet, 0.9 mm., 50.
3. Leptasterias fisheri, Okhotsk Sea; abactinal pedicellaria, 0.32 mm., 200.
3a. Same, one jaw of an actinal axillary pedicellaria, >< 50.
3b. Side view of a similar pedicellaria, 1.17 mm. long, 50.
3c. Same, pedicellaria from an adambulacral spine, 0.7 mm., 50.
3d. A dorsolateral spinelet, 0.9 mm., X20.
3e. An inferomarginal spinelet, 20.
3f. A carinal spinelet, X20.
3g. A superomarginal spinelet, 20.

266

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U. S. NATIONAL MUSEUM BULLETIN 76. PART 3 PLATE !}

ere

1. LEPTASTERIAS HYPERBOREA. 2. L. ORIENTALIS. 3. L. FISHER!

FOR EXPLANATION OF PLATE SEE PAGE FACING.

A UM BULLETIN 76, PART 3 PLATE 12
U, S. NATIONAL. MUSE MI

LEPTASTERIAS CCHOTENSIS

EXPLANATION OF PLATE SEE PAGE FACINGe

PLATE 12
All figures Leplasterias ocholensis

Fieure 1. Abactinal crossed pedicellaria, 0.235 mm., 200. Specimen No. 1354, Mus. Acad.

Sci., U. R. 8S. 8.; coll. Middendorf.

la. Two profile views of jaws.

1b. Same.

lc. Inside of end of jaw, %200. 1a to le, specimen figured on Plate 27.

2. Crossed pedicellaria from an adambulacral spine, profile view.

2a. Inside of end of jaw (compare with lc), 200.

3. Three abactinal spinelets, * 50.

4. Two abactinal spinelets, that on right from carinal series, 50.

5. An axillary straight pedicellaria, 0.7 mm. long, * 100.

6. A small abactinal straight pedicellariae, < 100.

7, 7a. Straight pedicellaria of adambulacral spine; a profile view, and inside and back of
two different jaws, 100.

8, Sa. Two inferomarginal spines, * only 38.

9. Plan of skeleton of middle of ray from actinal plates (AC) nearly to carinals; supero-
marginal plates stippled, secondary superomarginals cross-lined, secondary infero-
marginal solid black, *5. No. 1354, Mus. Acad. Sci. U. R. 8. S.; coll. Middendorf.

10. Mouth and first two pairs of adambulacral plates, & 10.
267

Puate 13
Leptasterias (Nesasterias) slolacantha

Ficure 1. Projection of skeleton of base of ray, the small circles in the intervals indicating papulae;
base of ray on left; *8. CAR-—C, carinal series; D, dorsolateral; S, superomarginal;
I, inferomarginal; A, two series of actinals; AD, adambulacrals; g, point where
gonad is attached.

2, 2a. Abactinal spinelet, showing the sheath and its crown of small pedicellariae, 10.

2b. Marginal spinelets, > 10.

2c. Outer (left) and inner adambulacral spinelets, 10.

3. Adambulacral spinelet, 1.5 mm., 30.

3a. Marginal spinelet, 30.

3b. Station 3319, marginal spinelet, 30.

4. Mouth, angle; g, genital papilla; 10.

5. Abactinal crossed pedicellaria, 0.135 mm., station 3321, 200.

5a. A jaw, 400.

5b, 5c. Two views of adambulacral pedicellariae, station 3321, 400.

6. An axillary straight pedicellaria from near genital papilla (see fig. 4), 0.43 mm., 100.

6a, 6b. Abactinal straight pedicellariae, 0.36 and 0.29 mm., 100.

6c. From adambulacral spinelet, 0.23 mm., 100.
268

ERIAS) STOLACANTHA

er (NESAS
FO EXPLANATION OF PLATE GEE PAGE FACING.
.

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 14

IES

wes
it, bt ;

LF

.

LEPTASTERIAS ARCTICA

FOR EXPLANATION OF PLATE SEE PAGE FACING

PLATE es
io! Leptasterias arctica
‘Forma arctica, cotype, No. 7625.
ame, probably station 3514, 59° 22’ N.. 168° 21” W.
3. Same, locality lost, cleaned to show skeleton.
4. ame, Nikolski, Bering Islands.
Forma beringensis, station 3303.

station 3302.
/ 269

Fieure 1.

Rw

PuaTeE 15
Leptasterias arctica

Forma arctica, station 3559, not typical.

Forma beringensis, intermediate with f. arctica, station 3510.
Forma beringensis, station 3293.

Same, station 3294.

Same (type of forma), station 3277.

270

» PART 3 PLATE 16

76

BULLETIN

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OF PLATE

Ficvure l.

Puate 16
Leplasterias arctica, enlarged
Forma arctica, 60° 22’ N., 168° 45’ W.

2. Forma beringensis, station 3294.

aoe &

. Same, station 3504.

Same, station 3293.

. Same, station 3294.
. Same, station 3510.

271

PuaTe 17
Lepiasterias hylodes,
Fiaure 1. Station 2856.
2, 3, 4. Station 2847.
5. Type, station 2851.
272

< 134.

IDE

HYLE«

STERIAS

LEPTA

U.S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 18

1-3. LEPTASTERIAS HYLODES. 4, 5. L. (NESASTERIAS) STOLACANTHA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

@

. “Puate 18
eptasterias hylodes, station 2851, x2. |
: a ‘stolacantha, station 3321, X24.

273

PLATE 19 '

Ficure 1. Leptasterias hyperborea, Spitzbergen, No. 207, Riksmuseum, Stockholm, 2.
2. Leptasterias orientalis, Okhotsk Sea, 57° 36’ N., 140° 34’ E., 106 meters, mud, 2.
3. Leptasterias orientalis, station 5024, 2.

274

me

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ea

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U. S. NATIONAL MUSEUM BULLETIN 76, PART 3. PLATE 20

\

| we Ne 3 i 7
a ays at
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4 a Pe 4 te , 3
* ,

1,2. LEPTASTERIAS FISHERI 3. L. ORIENTALIS

FOR EXPLANATION OF,PLATE SEE PAGE FACING-

Piate 20

Ficure 1. Leptasterias fisheri, Okhotsk Sea, 50° 03’ N., 144° OS’ E., 110 meters, mud, 14.
2. Leptasterias fisheri, actinal view of Figure 1, 246.
3. Leptasterias orientalis, Okhotsk Sea, 57° 36’ N., 140° 34’ E., 106 meters, mud. Abactinal
view of Plate 19, Figure 2, «144. se
af

PuaTE 21
Leptasterias groenlandica, <2

Ficures 1, 2. Forma cribraria, station 3252.
3. Forma groenlandica, station 3520.
4. Forma cribraria, No. 1349, Arctie Ocean.

276

U. S. NATIONAL MUSEUM

BULLETIN 76, PART 3 PLATE 22

LEPTASTERIAS GROENLANDICA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLATE 22
Leptasterias groenlandica, 135

Ficure 1. Forma groenlandica, station 3506.
. Same, station 3520.

. Forma cribraria, station 3522.
Same, No. 1349, Arctic Ocean.

. Same, station 3252.

ore wb

bo
wl
“I

PuatTE 23

Leptasterias groenlandica forma groenlandica, *

Ficure 1. From north of Asia, Stanford collection.
2. Station 3531.
3, 4, 5. Station 3252.

278

U. S. NATIONAL MUSEt

LEPTASTERIAS GROENLANDICA

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 24

MA ie cata Ned eg. -
Agee fe fet z, Poth s eRe
Ah up t Lm cal,» “999 80 *
Nord? a bite nse ets #4.

‘a CP 4
iy

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1, 2. LEPTASTERIAS GROENLANDICA.{ 3. L. COE TRUCULENTA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PiaTe 24

Figure 1. Leplasterias groenlandica forma groenlandica, station 3506, 1%. Resembles L.

hyperborea.
2. Leptasterias groenlandica forma cribraria, station 3522, 1%.

3. Leptasterias coei truculenta, station 4779, K1%.-
279

PLATE 25

Sv

Fiaure 1. Leptasterias coei truculenta, type, Captains Bay, Unalaska, 0.
2. Same, station 4779.

280

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BULLETIN 76, PART 3 PLATE 26

NATIONAL MUSEUM

U.S

LEPTASTERIAS COEI TRUCULENTA

PuatTe 26

‘2. a!, a, a’, three series of actinal spines; i, inferomarginals;

Leptasterias coei truculenta, type,
s, superomarginals; g, aperture of gonoduet.
251

PLATE 27
Leplasterias ochotensis, abactinal view of specimen from Zoological Museum, Copenhagen; a
little over 3.

282

bo

SEUM

MU

NATIONAL

Uu

23

PLATE

PART 3

BULLETIN 76,

NATIONAL MUSEUM

Us

LEPTASTERIAS CCHOTENSIS

INGs

OF FLATE SEE PAGE FAC

XPLANATION

E

FOR

PLaTE 28
Leplasterias ochotensis

Figure 1. Actinal aspect of Plate 27.
2,3. Two views of Brandt’s var. macrobrachia, Zoological Museum, Copenhagen, 1.3;

Dr. Th. Mortensen.
283

PLATE 29

Fiaure 1. Leptasterias coei truculenta, station 4779; abactinal pedicellariae, 0.3 mm., 100.

la. From adambulacral spine, 0.27 mm., 100.

1b. From adambulacral spine, 200.

lc. Abactinal, 200.

1d. From adambulacral spine of type, 200.

le. Type, abactinal, 200.

2. Leptasterias coei truculenta, two abactinal spinelets of type,

2a. Same, station 4779, abactinal spinelets, 25.

2b. Type, actinal spine, * 25.

3. Leptasterias coei. Type of Orthasterias merriami Verrill, abactinal pedicellariae, 0.8 mm,
100.

3a. Same specimen; from adambulacral spinelet, 200.

4. Leptasterias coei truculenta, station 4779; abactinal straight pedicellariae, 50.

4a, 4b, 4c. Same, station 4779; three views of intermarginal pedicellariae, the largest
1 mm. high, 50.

4d. Same, type; intermarginal, 1.2 mm., 50.

4e. Same, type; from adambulacral spine; > 50.

4f. Same, type; from border of adambulacral furrow, 0.29 mm., 50.

5, 5a. Leptasterias coei. Type of Orthasterias merriami Verrill; three views of lateral straight
pedicellariae, >< 50.

284

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BULLETIN 76, PART 3 PLATE <9

U. S. NATIONAL MUSEUM

1-4. LEPTASTERIAS COE! TRUCULENTA. 5. L. COE!

FOR EXPLANATION OF PLATE SEE PAGE FACING.

U. S. NATIONAL MUSEUM

BULLETIN 76, PART 3 PLATE 30

1. LEPTASTERIAS POLARIS. 2. L. POLARIS BOREALIS. 3-5. L. POLARIS ACERVATA

FOR EXPLANATION OF PLATE SEE PAGE FACING,

Pirate 30
All pedicellariae abactinal unless specified

Figure 1. Leplasterias polaris, Davis Strait, No. 38834, U. S. Nat. Mus.; one jaw of a crossed

pedicellaria from side, 200.

la. Same specimen, length 0.26 mm., 100.

1b. Same specimen, end of jaw, 200.

lc. Same, Disco Island, Greenland, No, 15815, U. S. Nat. Mus., length 0.29 mm., 100.

2. Leptasterias polaris borealis, West Bank, from a very large specimen, length 0.27 mm.

2a. Same, Caribou Island, Labrador, No. 1307, M. C. Z.; specimen with spaced, rather
long spines, 0.26 mm., 100.

2b. Same, Newfoundland Banks, 0.26 mm., 100.

2c. Same, Gaspé, Quebec, M. C. Z., 1308, 0.27 mm.

2d. Same specimen, a jaw, 200.

3. Leptasterias polaris acervata forma acervata, No. 6111, U. S. Nat. Mus., Cape Lisburne,
Alaska, 0.225 mm., * 100.

3a. Same specimen (with small clusters of spines resembling pl. 37, fig. 2); a jaw, 200.

3b. Same, off Indian Point, Bering Sea, No. 6086, U. S. Nat. Mus. (pl. 37, fig. 4) 0.225
mm., 100.

3c. Same, station 3496; variety with very large bivalved straight pedicellariae, 0.32 mm.,
100.

3d. Same specimen, 200.

3e. Same, station 3235 (pl. 37, fig. 2), 0.22 mm.

3f. Same specimen, 200.

3g. Same, No. 39984, U. S. Nat. Mus., off St. George Island, Bering Sea, 0.30 mm., 200.

4. Same, pedicellaria from an adambulacral spine, 0.31 mm., 100. Station 3496,
same specimen as Figures 5a, 5b.

4a. Indian Point, Bering Sea, same specimen as Figure 3b; pedicellaria from an adambu-
lacral spine, 0.23 mm., 200.

5. Leptasterias polaris acervata forma polythela, station 3518; pedicellaria from an adambu-
lacral spine 0.30 mm., 200. .

5a. Same, station 3251; an adambulacral crossed pedicellaria, > 200.

PuatE 31
All pedicellariae are abactinal unless specified

Freure 1. Leptasterias polaris acervata forma polythela, No. 15817, U. S. Nat. Mus., Aretie Ocean

north of Bering Strait, 0.25, > 200.

la. Same, station 3518, specimen with well developed abactinal spines, 0.27 mm.,

1b. Same, station 3251, very large specimen, 0.26 mm., 200.

lc. Station 3439, 0.29 mm., 200.

ld. Same specimen as 1c, end of jaw, > 200.

le. Same, station 4795, off Kamchatka, 0.27 mm., 200.

1f. Same specimen, 100.

2. Leptasterias polaris acervata forma aphelonota, type, station 2850, 0.27 mm.

2a. Same, type; two pedicellariae showing different contours, 100.

2b. Same, station 3214, 100.

2c. Same specimen, 0.27 mm., 200.

3. Same, type; a pedicellaria from an adambulacral spine, 0.34 mm., 100.

3a. Same, type; a rare form of giant crossed pedicellaria from adambulacral spines, 0.45
mim., 100.

3b. Same, Old Harbor, Kodiak; two adambulacral spine pedicellariae, 0.25 mm., 100;
jaw, 200.

4. Same, type; a rare form of straight pedicellaria from adambulacral spine, 0.16 mm., 50.

X< 200.

, X200.

286

BULLETIN 76. PART 3 PLATE 31

U. S. NATIONAL MUSEUM

LEPTASTERIAS POLARIS ACERVATA

FOR EXPLANATION OF PLATE SEE PAGE FACING

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 32

J

1,2. LEPTASTERIAS POLARIS ACERVATA. 3. L. POLARIS BOREALIS

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Puate 32
All straight pedicellariae

Fioure 1. Leplasterias polaris acervata forma aphelonola, type; large-toothed pedicellaria from

intermarginal channel 1.12 mm. long.

la, 1b, 1c. Same, abactinal and lateral pedicellaria and two detached jaws, 50.

1d. Same, from adambulacral spine, 0.63 mm., 50.

le, 1f. Same, Old Harbor, Kodiak, lateral, 0.8 mm., 50.

1g. Same specimen; two jaws of an axillary pedicellaria, of the type common in the type
specimen but rare in this.

1h. Same specimen, jaw of an actinal pedicellaria, X50.

2, 2a. Leptasterias polaris acervata forma acervata, Cape Lisburne, Alaska, No. 6111, U. Ss.
Nat. Mus.; two views of an intermarginal pedicellaria 0.75 mm. long, 50.

2b, 2c. Same specimen, small abactinal pedicellariae.

2d, 2e. Same, station 3496; two views of intermarginal pedicellariae, 1.2 mm. long, * 50.
This and some other specimens from the region of the Pribilof Islands have very
large straight pedicellariae.

3, 3a. Leptasterias polaris borealis, Western Bank, No. 14339, U. S. Nat. Mus., very large
specimen. Side view of the largest intermarginal pedicellaria, 0.58 mm. long, and

two detached jaws.
287

PLATE 33

Leptasterias polaris katherinae, except Figure 7

Ficure 1. Cotype No. 111, abactinal crossed pedicellaria, 0.27 mm., 200.

la. Jaws from (left) a superomarginal and (right) an abactinal pedicellaria, 0.27 mm.,
< 200.

1b. Another type of abactinal pedicellaria, 0.26 mm., 100.

lc, 1d, le. Adambulacral spine pedicellariae and parts; 1d is the inside view of tip of jaw;
le, 0.38 mm. long; all 100.

1f. Side view of carinal plate, showing three abactinal spines, 10.

2, 2a. No. 1181, Museum of Comparative Zoélogy, Gulf of Georgia, two abactinal pedicel-
lariae, the left 0.225 mm., > 100.

3. No. 110, Gray’s 37) (smallest specimen), an abactinal crossed pedicellaria, 0.23 mm.,
100.

3a. Jaw of a pedicellaria from an adambulacral spine, 0.27 mm.

4, da. Type, straight pedicellariae, intermarginal channel, 0.5 and 0.6 mm., 50.

4b. From furrow spine, 0.375 mm., 50.

4c. Valve of same from inside, 100.

4d. Basal piece, 100.

de. Valve of furrow spine pedicellaria, detached, 0.35 mm., >< 100.

5. No. 110, Gray’s 37b, an adambulacral straight pedicellaria similar to those from other
parts of the body, 100.

6. No. 1181, M. C. Z.; an abactinal straight pedicellaria, >< 100.

7. Leptasterias polaris acervata f. aphelonota, type, various spines, 10.

7, 7a. Inferomarginal.

7b. Superomarginal.

7c. Subambulacral.

7d; a, b, c, d. Abactinal.

288

U. S. NATIONAL MUSEUM BULLETIN 76. PART 3 PLATE 33

1-6, LEPTASTERIAS POLARIS KATHERINAE. 7. L. POLARIS ACERVATA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

BULLETIN 76, PART 3 PLATE 34

U' S. NATIONAL MUSEUM

LEPTASTERIAS POLARIS KATHERINAE

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Ficure 1.

PLate 34
Leptasterias polaris katherinae
Cotype, skeleton of ray from the outside, showing actinal, lateral, and part of dorsal
plates, base of ray to left, <4; D, dorsolateral plates; S, superomarginals; J, infero-

marginals; i 7, intermarginal papular areas; A' A*, outer and inner series of actinal
plates; AD, adambulacrals; AM, outer ends of ambulacral ossicles.

. Cotype, portion of dorsal skeleton from coelomic surface, showing mostly the connecting

plates; CAR-C, radial line of plates; base of ray is to the right, 4.

. Type, plates of the mouth angle. The ambulacral plates are shown on the left. On

the right are the mouth plates and the adoral adambulacrals, & 5.

. Cotype, adambulacral armature near middle of ray. The thick spine above is an inner

actinal.
Cotype, a carinal plate, 10.
Type. an inferomarginal spine, 10.
289

PuatE 35
Leptasterias polaris borealis, X %

Figure 1. Caribou Island, Labrador, No. 1807, Mus. Comp. Zodl.
2. Gaspé, Quebec, No. 1308, Mus. Comp. Zo6l.
3. Caribou Island, Labrador, No. 1307, Mus. Comp. Zo6él. This is fairly typical Leptas-
terias polaris borealis and is one of the lot from which Perrier’s type was derived.
290

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U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 36

1. LEPTASTERIAS POLARIS KATHERINAE 2,3. L. POLARIS ACERVATA FORMA APHELONOTA

FOR EXPLANATION OF PLATE SEE PAGE FACING

PLatpS 36

Fiaure 1. Leptasterias polaris katherinae, portion of abactinal surface of type, 2.
2. Leptasterias polaris acervata forma aphelonota, type, <0.66.
3. Same, portion of actinal surface of type, 1.6.

PLATE 37

Leptasterias polaris acervata forma acervala, 14
Ficure 1. Station 3233, Bristol Bay, Bering Sea, shallow.
2. Station 3235, Bristol Bay shallow.
3. Station 3233 Bristol Bay, shallow.
4. Indian Point, Siberia, 17 fathoms.
292

Uu. S. NATIONAL MUSEUM

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U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 38

LEPTASTERIAS POLARIS ACERVATA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Puate 38
Leptasterias polaris acervata, * 1%

Fiavre 1. Forma acervata, station 3233, Bristol Bay, Bering Sea.
north of Bering Strait (Corwin, No. 15818).

2. Forma acervata—polythela,
3. Forma polythela, station 3277, north of end of Alaskan Peninsula.
293

Puate 39
Leptasterias polaris acervata forma polythela, natural size

Figure 1. Station 3251, north coast Unimak Island.
2. Station 3518, Bering Sea, near St. Matthew Island.
3. Station 3439, Bering Sea, west of Pribilof Islands.
294

PLATE 40

BULLETIN 76, PART 3

MUSEUM

NATIONAL

U.S

tS
eS

LEPTASTERIAS POLARIS ACERVATA FORMA POLYTHELA

SEE PAGE FACING.

XPLANATION OF PLATE

FOR &

Prate 40

Leptasterias polaris acervata forma polythela, natural size

Piaure 1. Station 3251, north coast Unimak Island.
2. Station 3440, Bering Sea, west of Pribilof Islands.

205

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(Puave 410
moet eck META polaris katheri 1

Ficure 1. Portion of actinal surface of type, speci
2: _ Cotype, specimen 111, of type series, <1

296

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 42

tanh] Ans i)
Nee?
ALY

Mi

1. LEPTASTERIAS CAMTSCHATICA. 2-4. L. CAMTSCHATICA DISPAR AND FORMAE

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLATE 42

Figure 1. Leptasterias camtschatica; projection of skeleton from actinals to carinals, at base of ray,
which is at left, X10; AC, two series of actinal plates; AD, direction of base of ray;
CAR-C, ecarinal series; DL, dorsolaterals; J, inferomarginals; S, superomarginals.

la. Same, dorsal spinelet, 0.5 mm., 50.

1b. Same, an abactinal crossed pedicellaria, 0.225 mm., » 200.

lc. Same, two lateral straight pedicellariae, 0.36 and 0.46 mm., 50.

ld. Same, Petropavlovsk, mouth plates, 10.

2. L. camtschatica dispar f. nesiotis; two abactinal spinelets, 0.4 and 0.37 mm., 50.

2a. Same, intermarginal pedicellaria of type (0.42 mm.) and a jaw, 50.

3. L. camtschatica dispar f. nitida; abactinal crossed pedicellaria, 0.16 mm., 200 (St.
Paul, Pribilofs).

3a. Same, Pribilof Islands; two abactinal spinelets 0.7 and 0.5 mm., 50.

3b. Same, Unalaska, abactinal spinelet.

3c. Same, two straight pedicellariae, 0.8 and 0.63 mm., 50.

4. L. camtschatica dispar f. dispar, Attu Island, straight pedicellaria, 0.63 mm., *

da. Same, Nazan Bay, Atka; jaw of crossed pedicellaria, 0.27 mm., 200.

4b. Same, St. George, Pribilof Islands, 0.225 mm.

297

PuiatE 43

Ficure 1. Leptasterias aleutica; two abactinal spinelets, 0.38 and 0.4 mm., 100.

la, la’. Same crossed pedicellariae; a, Adakh; a’, Atka (0.2 mm.), 200.

1b, 1c. Same, straight pedicellariae, 0.3 and 0.24 mm., 100.

2. Leptasterias leptodoma; straight pedicellaria, 0.56 mm., 100.

2a. Same; abactinal crossed pedicellaria, 0.24 mm., 200.

2b. Same, Adakh Island; projection of skeleton of base of ray from the carinals CAR to
actinals A, A; AD, direction of base of ray, DL, dorsolaterals, very thin and irregular;
7, inferomarginals; S, superomarginals, 10.

3. Leplasterias asteira; abactinal spinelet, 0.6 mm., 100.

3a, 3b, 3c. Same, views of straight pedicellariae, 50; 3a, 0.34 mm.

3d. Same, abactinal crossed pedicellaria, 200.

3e. Same, projection of marginal (S, J) and actinal plates (A A), base of ray; AD direction
of disk.

298

U. S. NATIONAL MUSEUM BULLETIN 76. PART 3 PLATE 43

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1. LEPTASTERIAS ALEUTICA. 2. L. LEPTODOMA 3. L. AST

U. S. NATIONAL MUSEUM

BULLETIN 76, PART 3 PLATE 44

LEPTASTERIAS HEXACTIS AND FORMAE

Puate 44

Figure 1. Leptasterias hexactis, cotype, No. 1368, 10. Projection of skeleton from carinals
(c, c, c) to actinals (AC); a few adambulacrals indicated on left; AD, direction of
disk; d, d, d, dorsolaterals; 7, inferomarginals; S, superomarginals.

la. Same specimen; abactinal crossed pedicellaria, 0.2 mm., 200; tip of left jaw on
left, 400.

1b. End of another pedicellaria 400. The outer face is on right instead of left as in la.

lc, 1d. Same specimen, three straight pedicellariae, 0.67, 0.58, 0.54 mm. 50.

le. Same specimen, mouth plates and two adambulacrals, 10.

1f. Same specimen; two abactinal spinelets, 0.4 and 0.5 mm., % 100.

2. L. hexactis f. siderea, Kodiak Island; abactinal spinelet, 0.54 mm., 100.

2a. Same; pedicellaria from large intermediate specimen, Homer, Alaska, 0.6 mm., 50.

3. L. hexactis f. plena, Forrester Island, Alaska, abactinal pedicellaria, 0.2 mm., 200;
and outer face of terminal lip of another, «400.

4. L. hexactis f. aspera, Kodiak; abactinal spinelet, 0.43, 100.

da. Same, Cordova; abactinal pedicellaria, 0.25, 200.

4b. Same, Cordova; jaw of a straight pedicellaria, 100.

4c. Same, Cordova; straight pedicellaria, 0.58 mm., 50.

5. L. hexactis f. hexactis, intergrade with vancouvert (Class III), San Juan Islands, Wash.,
x50.

6. L. hexactis f. regularis, San Juan Islands, Wash.; crossed pedicellaria, 200 (left) and

terminal lip, 400.
299

Puate 45

Ficure 1. Leplasterias hexactis vancouver, San Juan Islands, Wash., 5. Projection of skeleton,

base of ray of a large specimen; CAR, carinals; DL, dorsolaterals; S, superomar
ginals; J, inferomarginals; AC 1, 2, two series of actinals; AD, direction of disk.

la, 1b. Same, two abactinal spinelets, 0.5 mm., 100.
lc, ld, le. Same, straight pedicellariae, 0.45, 0.54, 0.58 mm., ><508
2, 2a. Leptasterias hexactis f. aspera, Kodiak, straight pedicellariae, 0.55 and 0.46 mm.,

x 100.
3, 3a, 3b. Leptasterias pusilla, straight pedicellariae, 50; 3, 3b, from actinal interradial

region; 3a, from adambulacral spinelet.

3c. Same, mouth and two pairs of adambulacral plates, 20.

3d. Same, projection of skeleton, base of ray,*<10; CAR-C, carinals; DL, dorsolaterals,
stippled; S, superomarginals; J, inferomarginals; AC 1, 2, actinals, the second
series in solid black; AD, direction of disk, and seven adambulacral plates.

300

BULLETIN 76, PART 3 PLATE 46

U. S. NATIONAL MUSEUM

Sy Kuhae i

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A

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3. L. AEQUALIS

2. L. PUSILLA.

LEPTASTERIAS HEXACTIS VANCOUVERI.

i:

FOR EXPLANATION OF PLATE SEE PAGE FACINGs

PLATE 46

Figure 1. Leplasterias hexactis vancouveri, a small dorsolateral spinelet, 100.

2. Leptasterias pusilla; dorsal spinelets, 0.32 to 0.36 mm., 100.

2a, 2b. Same, dorsolateral spinelets, 100.

2c. Actinal spinelet, < 100.

2d. An abactinal crossed pedicellaria, < 200.

3. Leptasterias aequalis, Monterey Bay; abactinal crossed pedicellaria, 0.21 mm., 200,
and, to right, outer face of terminal lip showing enlarged lateral teeth, «400.

3a. Same, Friday Harbor, Wash., abactinal, 0.24 mm.

3b. Same, Monterey, abactinal, 0.2 mm., 200.

3c. Same, Monterey, from an adambulacral spinelet, 0.24 mm.

3d. Same, from an actinal spinelet, 0.23 mm.

3e. Same, Monterey, 10; projection of skeleton from carinals, CAR to actinals AA;
DL, 1, 2, dorsolaterals; S, superomarginals; 7, inferomarginals; AD, direction of
disk; five adambulacrals shown at bottom.

301

PuaTE 47
Leptasterias aequalis

Figures 1, la, 1b. Monterey Bay, dorsolateral spinelets from typical large specimen, 0.55, 0.7,
and 0.5 mm., L100.
Ic. Abactinal spinelets from a small, slender-spined individual with R 12 mm., 100.
1d. Carinal spinelets from asmall specimen (R 13 mm.) with short granuliform spinelets,
0.19 and 0.27 mm., 100.
le, 1f. Abactinal spinelets (Monterey specimen), 0.36 and 0.45 mm., »< 100.
2. Mouth plates and two adambulacrals, from typical large Monterey specimen.
2a. Another variation.
3. Two straight pedicellariae from ambulacral furrow, 0.48 mm., 100.
3a, 3c. Two pedicellariae from adambulacral spinelets, 0.24 and 0.225 mm., 200 (about
half as large as 3).
3b. One of the narrower straight pedicellariae from oral spinelet, 0.4 mm., 100.
3d. A similar one from the actinal interradial region, 0.45 mm., > 100.

302

PLATE 47

BULLETIN 76, PART 3

U. S. NATIONAL MUSEUM

——

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LEPTASTERIAS AEQUALIS

FOR EXPLANATION OF PLATE SEE PAGE FACING.

BULLETIN 76, PART 3 PLATE 48

U. S. NATIONAL MUSEUM

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3. L. ALASKENSIS F. SHUMAGINENSIS.

2. L. ALASKENSIS.

LEPTASTERIAS ALASKENSIS ASIATICA.

Ais

5. L. ALASKENSIS F, PRIBILOFENSIS

4. L. ALASKENSIS MULTISPINA.

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLaTEe 48

Figures 1, la. Leptasterias alaskensis asiatica, type, 25. Intermarginal straight pedicellariae

for comparison with Figure 2, same magnification, Left figure, 0.72 mm.

1b. Same, Bering Island, 0.58 mm., 50 (twice enlargement of 1).

1c, 1d. Same, Simushir, Kuril Islands, two detached jaws, 50, for comparison with 2a,
same enlargement.

2. Leptasterias alaskensis f. alaskensis, Unalaska. Three views of a large bivalved lateral
pedicellaria, 1 mm., X25.

2a. Detached jaws, 50.

2b. Crossed pedicellaria from adambulacral spinelet, 200.

2c. Abactinal crossed pedicellaria, Unalaska, 200.

2d. Same from St. Paul, Kodiak, 0.24 mm., 200.

2. Three abactinal spinelets, Kodiak specimen, 0.72, 0.8, 0.6 mm., 50.

2f. Projection of skeleton, basal third of ray, showing irregular carinal series (CAR-C),
broad dorsolateral area (D, D), marginals (S, T) low on side of ray, inconspicuous
actinals (AC); AD, direction of disk, 7.

3. Leptasterias alaskensis f. shumaginensis, X25. Lateral bivalved pedicellaria, 0.63 mm.
(compare with 1 and 2).

3a. Two jaws, *50 (compare with 1c, 1d, 2a, same enlargement).

4. Leptasterias alaskensis multispina, Wrangell, Alaska, * 50. Two lateral pedicellariae
(compare with fig. 2; but note different magnifications).

5. Leptasterias alaskensis f. pribilofensis, St. Paul, Pribilof Islands. Abactinal spinelet
4100 (twice magnification of 2e).

5a. Lateral pedicellaria, 25 (compare with fig. 2, same enlargement).
303

FIGURE 1.

9

3.

. Leplasterias camtschatica dispar (Verrill) forma dispar, Unalaska, Aleutian Islands, % 2.

PLATE 49

Leptasterias camtschatica Brandt, Medni, Commander Islands, 1}%.
Leptasterias camtschatica dispar (Verrill) forma nitida Fisher, St. Paul, Pribilof Islands,

x1.
Leptasterias camtschatica dispar (Verrill) forma nesiolis Fisher, St. Paul, Pribilof Islands,
KIS.

Leplasterias camtschatica dispar (Verrill) forma nitida-dispar, St. George Island, Pribilof
Islands, 114.

. Leptasterias camischatica dispar (Verrill) forma dispar, Attu, Aleutian Islands, «1%.

304

NATIONAL

U. S. NATIONAL. MUSEUM BULLETIN 76, PART 3 PLATE 50

2. 3. L.ALEUTICA. 4, 5. LE. ASTEIRA. ‘6. L. LEPTODOMA
7. L. HEXACTIS FORMA HEXACTIS

1. LEPTASTERIAS ‘CAMTSCHATICA DISPAR FORMA NITIDA.

FOR EXPLANATION OF PLATE SEE PAGE FACING

Figure 1.

NOOR oN

Pirate 50

Leptasterias camtschatica dispar forma nitida, Karluk, Kodiak, > 1's.
Leptasterias aleutica, Unalaska, 3.

. Leptasterias aleutica, type, Adakh Island, Aleutian Islands, % 2.

Leptasterias asteira, Sanborn Harbor, Nagai, Shumagin Islands, 3.
Leptasterias asteira, Adakh, Aleutian Islands, 3.
Leptasterias leptodoma, Atka, Aleutian Islands, 245.
Leptasterias heractis forma hexactis, Orcas, San Juan Islands, Wash., 2.
305

Puate 51

With the exception of Figure 4 all specimens are from Stimpson’s type series of Leptasterias hexactis,
No. 1368, U.S. N. M.; slightly less than twice natural size

Fiaure 1. Forma regularis.

Forma hexactis, the type specimen.

Forma hexactis, skeleton of one arm cleaned.

Forma aspera, Alert Bay, British Columbia (not a Stimpson specimen).
9. Forma plena.

306

2.
3.
4.
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MUSE

Al

NATION

BULLETIN 76, PART 3 PLATE 52

NATIONAL MUSEUM

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LEPTASTERIAS HEXACTIS

FOR EXPLANATION OF PLATE SEE PAGE FACING

Figure 1.

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PuaTe 52
Formae of Leptasterias hexactis

Forma siderea, Karluk, Kodiak, * 1%.

. Forma siderea, Karluk, Kodiak, 154.
. Forma regularis, Friday Harbor, San Juan Islands, Wash., 1's.

Forma regularis, Cordova, Prince William Sound, Alaska, » 2.
Forma aspera-reqularis, Karluk, Kodiak, * 1%.
Forma aspera, largely denuded to show skeleton, Kukak Bay, Shelikof Strait, Alsaka,
xX 1%.
Forma plena, Forrester Island, Alaska, 1).
307

PLATE 53

Leptasterias hexactis vancowvert (Perrier) San Juan Islands, Wash. (See text for explanation of

Ficure 1

m OO bo

or

variations)

. Specimen with broad carinals, groups of abactinal spinelets in longiseries, numerous

superomarginal spinelets (Ia !, b!), 1.1.

Specimen with narrower carinals (Ia!, b?), 1%.

Side view of ray of a specimen in Class Ia!, b? to show vertical combs of spinelets, 17s.

Specimen with numerous superomarginal spinelets but groups of abactinal spinelets not
in apparent longiseries (la?).

Specimen belonging to Class IJ, in which the superomarginal spinelets average three;
this specimen is intermediate between a! and a? and suggests Leptasterias camts-
chatica, X 1%4.

Small specimen of Class III (superomarginal spinelets two, sporadically three with
abactinal spinelets in longiseries (a!), 2.

, 8,9. Three specimens belonging in Class III, but with more or less irregularly arranged

Se 124

dorsolateral spinelets (a?), >< 136.
308

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U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 54

1-4. LEPTASTERIAS AEQUALIS 5s Le JPUSIEKA 6. L. HEXACTIS FORMA SIDEREA

FOR EXPLANATION OF PLATE SEE PAGE FACING

PLATE 54

Ficure 1. Lepltasterias aequalis (Stimpson) ; actinal view of alcoholic specimen, with eggs, Monterey

Bay, Calif., <2.

2. Leptasterias aequalis; abactinal view of alcoholic specimen in which the spinelets are not
in evident longiseries, Monterey Bay, 2.

3. Leptasterias aequalis; actinal view of dried specimen similar to Figure 4, vicinity of
Monterey Bay, X14.

4. Leplasterias aequalis; large typical specimen, vicinity of Monterey Bay, 1%.

5. Leplasterias pusilla Fisher; typical example, dried, for comparison with L. aequalis;
Cypress Point near Monterey Bay, 1's.

6. Leptasterias hexactis forma siderea, Middleton Island, Alaska, * 2.

309

Puate 55

Ficures 1, 2. Leptasterias pusilla, Cypress Point, vicinity of Monterey Bay, alcoholic specimen, <3.
3, 4, 5. Leplasterias aequalis forma nana, Monterey Bay, <3.
6. Leptasterias aequalis forma aequalis, Cypress Point, vicinity of Monterey Bay.
Female carrying young about ready to leave. Some are attached to the partly
everted stomach, 2.
7. Leptasterias pusilla, Cypress Point, vicinity of Monterey Bay. Female carrying young

in cluster over mouth, 2.
310

BULLETIN 76, PART 3 PLATE 56

NATIONAL MUSEUM

Ss

U

LEPTASTERIAS ALASKENSIS FORMA ALASKENSIS

SEE PAGE FACING.

FOR EXPLANATION OF PLATE

Piate 55
Leptasterias alaskensis forma alaskensis (Verrill)

Fiavre 1. Typical example from Unalaska, * 14s.
2. Specimen with smaller than typical bivalved pedicellariae, Karluk, Kodiak, 1.
3. Coarse spined example from Agattu, end of the Aleutian Chain, 1%.
4. Specimen from Attu, end of Aleutian Chain, showing clustering of dorsal spinelets,
1%. The form at end of Aleutian Islands is in some respects intermediate with
asiatica.

3ll

Figure 1.

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Bue CE EC

PuaTe 57

Leplasterias alaskensis forma shumaginensis, Humboldt Bay, Shumagin Islands, 175.
Leptasterias alaskensis forma pribilofensis, St. Paul, Pribilof Islands, xK1%.

Leptasterias alaskensis asiatica, type, Nikolski, Bering Island, 1.

Leptasterias alaskensis asiatica, Nikolski, > 1%.

Leptasterias alaskensis asiatica, Simushir, Kuril Islands, >< 14s.

Leplasterias alaskensis multispina, southern Alaska, 173.

Leptasterias alaskensis multispina, type, Wrangell, Alaska.

312

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U. S. NATIONAL MUSEUM BULLETIN 76. PART 3 PLATE 58

EVASTERIAS TROSCHELII

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Piate 58
Evasterias troschelii and formae (all crossed pedicellariae are abactinal, » 200)

Fieure 1. Evasterias troschelii forma troschelii. Pedicellaria from dorsal surface of giant specimen,

Victoria, 0.27 mm. long, 200.

la. Same, abactinal pedicellaria from typical specimen (pl. 60, fig. 1) Friday Harbor, Wash.,
0.28 mm., * 200.

2. Forma alveolata, Unalaska variety (pl. 61, fig. 2) abactinal, > 200.

2a. Same, coarse spined variety (pl. 62, fig. 2) Puget Sound, 0.324 mm., * 200.

2b. Same, small spined variety, Departure Bay, 0.225 mm., 200.

2¢. Same, Carmel Bay, Calif., multispinous variety, 0.24 mm., 200.

3. Forma acanthostoma, typical, Unalaska (pl. 64, fig. 3), 0.30 mm., 200.

3a. Same, Tongass, Alaska, variety (pl. 64, fig. 2), 200.

3b. Same, from large Victoria specimen, 0.225 mm. (PI. 65, fig. 2.)

4. Forma froschelii, ordinary intermarginal straight pedicellaria, giant specimen, Victoria
(pl. 61, fig. 1), 100.

5. Forma alveolata, actinal straight pedicellaria, Carmel Bay, Calif., 50.

6, 6a. Forma troschelii (pl. 60, fig. 2), two abactinal spines, » 25.

7. Forma alveolata, Departure Bay, abactinal spine, > 25.

7a, 7b. Same, Carmel Bay, Calif., abactinal spines, * 25.
8, 8a, 8b. Forma acanthostoma, Unalaska, abactinal spines, 25.
Sc. Same, Victoria, British Columbia.

Puate 59

Friaure l. Evastlerias troschelii forma alveolata; part of actinostomial ring, showing odontophore (OQ),
mouth plates (VW), and first ambulacrals (A), 10.

la. Same, mouth plates and adjacent ambulacrals, >< 20.
2. EHvasterias echinosoma; side view of mouth plates, showing three marginal spines (usually
two), and relative size of two straight pedicellariae, 5.

2a—2c. Same, straight pedicellariae; 2a, from intermarginal channel, 1.1 mm., 50;
2b, type, actinal interradial channel, 1.5 mm.; 2c, type, base of ray, inter-
marginal channel.

2d. Type, abactinal 0.09 to 1 mm.

2e. One jaw of 2d.

2f. An adambulaeral straight pedicellaria, >< 25.

2q. Same, an abactinal crossed pedicellaria, 0.27 to 0.8 mm., 100.

2h. An adambulacral crossed pedicellaria, 0.38 mm.

3. Hvasterias troschelii forma troschelii; large specimen from Victoria, actinal spine, 10.

3a. Inferomarginal spine.

3b. Forma acanthostoma, actinal spine, >< 10.

4. Evasterias echinosoma, actinal spine, >< 10.

4a. Same, abactinal spine, 10.

314

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 59

1,3. EVASTERIAS TROSCHELI!. 2,4. E. ECHINOSOMA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PART 3 PLATE 60

BULLETIN 76,

NATIONAL MUSEUM

S

U

EVASTERIAS TROSCHELII

SEE PAGE FACING.

OF PLATE

FOR EXPLANATION

Piate 60
Evasterias troschelit forma troschelii

Figure 1. Slender rayed variety with very slender secondary spinelets, Friday Harbor, San Juan
Islands, Wash., 1.

2. Variety with unusually stout primary spines. This is similar to Verrill’s ‘ Asterias
victoriana,”’ Friday Harbor, ™ 1.

3. Variety intermediate with forma alveolata, Orcas, San Juan Islands, Wash., 2%.
4. Type specimen, U.S. Nat. Mus.

315

Puate 61

Ficure 1. Evasterias troschelii forma troschelii, Victoria, British Columbia, natural size; a giant
individual. This is an example of Verrill’s variety rudis. The peculiarities are
largely due to age.

2. Evasterias troschelii forma alveolata, Unalaska (No. 3391) natural size. A subforma
showing the very unequal dorsal spines of forma troschelizi and the reticulate arrange-
ment of alveolata.

3. Evasterias troschelii forma alveolata, Naha Bay, Alaska, X 2.

316

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 62

EVASTERIAS TROSCHELII FORMA ALVEOLATA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLate 62
Evasterias troschelii forma alveolata 1%

Figures 1. ‘“‘Acervate’’ variety, Union Bay, British Columbia.

2. Stout rayed variety, from Puget Sound, near Tacoma, in which there is great disparity
in the size of the dorsal spines.

3. The variety predominating at Unalaska.

4. Typical specimen with a few spines enlarged and with a well-developed dorsal recti-
culum, Oreas, San Juan Islands, Wash.

317

PLATE 63

Evasterias troschelii forma alveolata, 11%

Fiaure 1. Typical specimen from Departure Bay, British Columbia.
9

2. The multispinous subforma, Tongass, Alaska.
3. The same variation with coarser spines, Fort Rupert, British Columbia.
318

DBE pty
wat : cs 1 OE Be 45
Bay

4 ioe 2 Vad
mt god

PLATE 64

3

PART

BULLETIN 76

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U

SHELII

TROSC

EVASTERIA

FACING

PAGE

E

ATE E

tL

XPLANATION OF

Puate 64

Ficure 1. Evasterias troschelii forma troschelii, Friday Harbor, San Juan Islands, Wash., natural
size.

2. Evasterias tros

3. Evasterias troschelii forma acanthostoma,

chelii forma acanthostoma, Tongass, Alaska, natural size.
typical example from Unalaska, natural size.

319

Puate 65
Evasterias troschelii forma acanthostoma
Ficure 1. A very slender rayed variety from Oreas, San Juan Islands, Wash., 5%.
2. Variety with heavy rays from Victoria, British Columbia, apparently occurring with the
similar giant form of troschelii, < %.
3. Unalaska, 214.

320

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BULLETIN 76, PART 3 PLATE 66

U. S. NATIONAL MUSEUM

<a

ae

2. FORMA ACANTHOSTOMA

SCHELII FORMA ALVEOLATA

ROS

T

TERIAS

aleeolata, Unalaska, actinal surface, X13.

321

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BULLETIN 76, PART 3 PLATE 68

U. S. NATIONAL MUSEUM

EVASTERIAS ECHINOSOMA

FOR EXPLANATION OF PLATE SEE PAGE FACING-

PuatE 68

: Evasterias echinosoma, natural size
e specimen from Kamchatka, station 4796.

ample from station 3291.

323

PLATE 69
Evasterias echinosoma, actinal surface of type, station 3278, natural size

324

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PART 3 PLATE 70

BULLETIN 76,

NATIONAL MUSEUM

Use

7. APHANASTERIAS PYCNOPODIA

6,

AS ALBULA

STEPHANASTERI

c
2

1

GE FACING

PA

E

PLATE

EXPLANATION OF

FOR

Puiate 70

} Enlarged % 135

Figure 1. Stephanasterias albula, station 3548; large 7-rayed specimen, with large unguiculate
: ve straight pedicellariae; 7, M, madreporic bodies.

2. Same, station 4792, seven rays.

3. Same, station 4777, eight rays.

4. Same, station 4792, eight rays.

5. Same, station 4792, eight rays.

6. Aphanasterias pycnopodia, type.

7. Aphanasterias pycnopodia, type. a

Figure 1.

la.
1b.
les

ld.

3.

PLatE 71

Stephanasterias albula, station 4792; projection of skeleton from carinals, C, to infero-
marginals, 7, proximal third of ray; 1, 2, 3, 4, dorsolateral series of skeletal meshes;
AD, direction of disk; S, superomarginals; 12. y

Same, station 4792; mouth plates and first two pairs of adambulacrals; >< 20.

Same, abactinal pedicellaria, 0.25 mm., >< 200.

Same, station 3548; a straight pedicellaria from mouth plates, two views, 1.25 mm.,
X25. 5

Same, station 4792; abactinal straight pedicellaria, 0.54 mm., > 50.

Same pedicellaria, 25 for comparison with le.

Same, station 3548; abactinal pedicellaria, >< 50.

Same, station 4792; two abactinal spinelets, 0.4 and 0.5 mm., 100.

Leptasterias aequalis, embryos carried by female, drawn from living specimens, much
enlarged.

Two views of a later stage, rays beginning to appear; 3-lobed larval organ below;
March 25, 1927; 20.

. A still later stage (March 30, 1927), 22.

Stage in which the six rays are well defined and larval organ is disappearing (April
4, 1927). :

Perissasterias polyacantha, 170 fathoms, northwest of Dassen Island, Cape Colony, No.
2875, Mus. Comp. Zodl., oral angle, <4.

326

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 72

S PYCNOPODIA. 4. APHELASTERIAS JAPONICA. 5. STEPHANASTERIAS ALBULA

1-3. APHANASTERIA

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLATE 72

Ficure 1. Aphanasterias pycnopodia. Skeleton of ray; C, carinal, S, superomarginal, /, infero-

marginal, AD, adambulacral plates; 10, from about 14 mm. from oral angle;
base of ray to right. The circles indicate position of the compound papulae.

la-lg. Same, spines enlarged, ><50.

la. A superomarginal, 0.9 mm., with a somewhat smaller than average crossed pedi-
cellaria.

1b. A carinal, 0.67 mm.

1c, 1d, le. Dorsolateral spines (0.5, 0.49, 0.81 mm., respectively); le is from a plate
just above the superomarginals.

1f. A rather heavy inferomarginal, 0.125 mm.

1g. An adambulacral, 0.13 mm.

2. Same, mouth plates and adoral carina, X10. Two adambulacrals are shown with spines.

3, 3a. Same, crossed pedicellariae; 3, dorsolateral, 0.32 mm., 100; 3a, a single jaw,
200. Compare with 4 and 5.

3b, 3c, 3d. Same, straight pedicellariae; 3b is from furrow margin 0.8 mm. long, 100;
3c (1 mm.) and 3d (0.63 mm.) unguiculate form from abactinal area, 50.

4. Aphelasterias japonica, abactinal pedicellaria, for comparison (0.17 mm.), * 100.

5. Stephanasterias albula, abactinal pedicellaria, >< 200.
327

PLatEe 73 .
Crossed and fureate pedicellariae >< 200 (except Sa)

Ficurn 1. Pisasler ochraceus forma ochraceus, Sitka, Alaska.
la. Same, Monterey Bay, Calif., both 0.27 mm. long.
2. Pisaster ochraceus forma confertus, Departure Bay, British Columbia.
3. Pisaster ochraceus forma nodiferus, Tongass, Alaska, length 0.31 mm.
4. Pisaster ochraceus segnis, Laguna Beach, Orange County, Calif., length 0.4 mm.
5. Pisaster giganteus, small specimen, Monterey Bay, Calif.
5a. Same, from a giant specimen, length 0.30 mm.
5b. Same, type specimen; dermal straight pedicellaria 0.15 mm. long, 200.
6. Pisaster giganteus capitatus, Laguna Beach, Calif.
6a. Same; jaw of a fureate pedicellaria; Venice, Calif.
7. Pisaster ochraceus forma nodiferus, Monterey Bay; jaw of a fureate pedicellaria.
&. Pisaster ochraceus segnis, type; jaws of a fureate pedicellaria.
8a. A whole pedicellaria, > 106.
328

U. S. NATIONAL MUSEUM

ARRAN

» Si

is or

1-3, 7. PISASTER OCHRACEUS, 4,8. P. CCHRACEUS SEGNIS
CAPITATUS

BULLETIN 76. PART 3 PLATE 73

5. P. GIGANTEUS. 6. P. GIGANTEUS

FOR EXPLANATION OF PLATE SEE PAGE FACING.

63160—30——27

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 74

1,2. PISASTER OCHRACEUS. 3. P. GIGANTEUS. 4. P. BREVISPINUS

FOR EXPLANATION OF PLATE SEE PAGE FACING

Puate 74
Fureate straight pedicellariae <200 unless otherwise specified

Figure 1. Pisaster ochraceus forma confertus, Departure Bay, British Columbia, from abactinal

surface.

la. Same specimen, an intermediate form, >< 100.

1b. Same specimen, back of jaw, X100. Note the height of base in proportion to blade
compared with giganteus (3a) and brevispinus (4b).

2. Pisaster ochraceus forma ochraceus, Monterey Bay, Calif., abactinal, 0.290.31 mm.

2a. Basal piece of above seen from below.

3. Pisaster giganteus, large specimen, Monterey Bay; an abactinal pedicellaria, 0.20.215
mm.

3a, 3b. Two jaws, same specimen. Note height of base in proportion to blade.

3c. From a small Monterey specimen.

4. Pisaster brevispinus forma paucispinus, Barclay Sound, British Columbia, 0.32 mm.
diameter.

4a, 4b. Same, station 4219; giant specimen.

4c. Forma brevispinus, San Juan Islands, Wash., giant specimen, diameter 0.27 mm.

4d. Forma brevispinus, Monterey Bay (station 4501).

4e. Forma paucispinus, Monterey Bay (three jaws). a

Puate 75
Pisaster ochraceus

Ficure 1. Portion of actinal and lateral skeleton from near base of ray of a medium-sized specimen
from Monterey Bay; S, superomarginals; /, inferomarginals; al—a5, actinal plates,
of which three transseries are shown. The secondary plates are shaded darker than
the primaries. The adambulacrals are adjacent to the three lowermost, or inner-
most, actinals, <8.

Mouth plates and adoral carina of a medium-sized specimen from Monterey Bay, 8.

. Actinal skeleton of a small specimen (R 80 mm.) showing three longiseries of actinal
plates and the lower-lying plates of the interbrachial septum. The adambulacral
plates are shown as a dark band, without detail. J, inferomarginals; a'—a’, actinal
plates; int., interradial line; Monterey Bay; 8.

. Abactinal skeleton of a young specimen (R 20 mm.) from Departure Bay, British Co-
lumbia, X10. On the left is the central portion of the disk, with the dorsal and
lateral skeleton of the proximal part of right half of ray, on right. (In this speci-
men there is one series of well-developed actinal plates, with the second just appear-
ing adjacent to adambulacrals; actinal skeleton not shown.) The upper dotted
line marks the carinal series (R—-R’) while the lower indicates the interradial line.
B, primary interradial plates, one carrying the madreporite; C, primary central
plate; DL, dorsolaterals; 7, inferomarginals; S, superomarginals; FR, primary radial
plates.

. Forma confertus, Departure Bay, British Columbia; seven abactinal spinelets, 0.85 to
1.2 mm. in length for comparison with 5a and 5c.

5a. Forma ochraceus; three abactinal spines from a Monterey specimen, > 10.

5b. An inferomarginal spine from same specimen.

5c. Forma nodiferus, Tongass, Alaska; an abactinal spine, 10.

6. Pisaster ochraceus segnis forma nodiferus, Laguna Beach, Orange County, Calif.; an

abactinal spine, 10.

330

bo

ow

rs

or

U. S..NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 76

1-7. PISASTER OCHRACEUS. 8. P. BREVISPINUS

FOR EXPLANATION OF PLATE SEE PAGE FACING.

PLate 76
Pisaster ochraceus

Ficure 1. Portion of actinostomial ring showing two odontophores (O) with proximal end of
ambulacral system between, mouth plates (M) below odontophores, 10.
2. Forma confertus, Puget Sound; an unguiculate straight pedicellaria from among actinal
spines, 1 mm. long, 50.
3. Forma confertus, Departure Bay, British Columbia; a large unguiculate pedicellaria
from the actinal interradial area, 1.5 mm., 50.
4. Forma ochraceus, Sitka, Alaska; inside of a valve of a dorsolateral unguiculate pedicel-
laria, X50.
5, 5a. Forma ochraceus, Monterey Bay, Calif.; two valves of characteristic unguiculate
pedicellariae, 5 being the inside, 5a the outside.
5b. Same specimen, side view, 50.
6. Forma ochraceus, Monterey Bay; the central large pedicellaria from an adambulacral
cluster (1 mm., 50) with one of the small pedicellariae drawn to scale (0.15 mm.).
6a. One of the small pedicellariae (0.22 mm.) of an adambulacral cluster, 200.
7. Actinal pedicellaria from a variation of forma confertus, Puget Sound (Tacoma), 1.3 mm.,
X25.
8. Pisaster brevispinus, station 4219; a festoon of straight pedicellariae from the adoral
carina, X3. These festoons are very numerous and long in this large specimen.

331

PLATE 77

Ficure 1. Pisaster giganteus; section out of actinal area, viewed from aboral end to show actinal
and inferomarginal spines; medium-sized specimen from Monterey, X10. a, a, a,
actinal plates; ad, adambulacral; am, lower end of ambulacral ossicle; x, super-
actinal buttress mentioned in text, and side of one of the superactinal fenestrated
pits.

2, 2a, 2b. Pisaster giganteus, type; three abactinal spines 2.7-3 mm. long, 10.

3. Pisaster giganteus, Monterey Bay; three abactinal spines, 1.6 mm. long, X10.

3a. Same; an abactinal spine from a medium-sized specimen, 25.

3b, 3c. Same; actinal and inferomarginal spines, same specimen as 3 and 3a, X10.

4. Pisaster giganteus capitatus, Laguna Beach, Calif.; two abactinal spines, 2.8 mm., X10.

4a. Same specimen; an inferomarginal spine, X10.

5. Pisaster giganteus, Monterey Bay; three clusters of abactinal papulae drawn from a
rather smaller than medium-sized live specimen, 6. j

6. Pisaster giganteus; pedicellaria from actinal interradial channel, same specimen as 3 and 5,
X25.

6a. Same specimen, from intermarginal channel, 1.2 mm., 25.

6b. Large specimen from Monterey; an intermarginal stone-hammer pedicellaria, 1.2 mm.,
X25.

6c. Same specimen as 6a, inside of a valve or jaw.

6d. Type of P. giganteus; an abactinal pedicellaria, 1.12 mm., X10.

7. Pisaster giganteus; mouth plates and inner part of adoral carina of a medium-sized
Monterey example, showing distorted development of first few adambulacral
plates, 8. This is from a dry specimen and the prominent festoons of straight
pedicellariae are indicated without detail.

7a. Actinostomial spine of another specimen.

332

U. S. NATIONAL MUSEUM

1-3, 5-7. PISASTER GIGANTEUS. 4. P. GIGANTEUS CAPITATUS

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 78

1-4. PISASTER BREVISPINUS. 5. P. GIGANTEUS

FOR EXPLANATION OF PLATE SEE PAGE FACING.

Puiate 78

Ficure 1. Pisaster brevispinus; adambulacral, actinal, and infermomarginal plates and spines of
forma brevispinus, viewed from a cut end of ray, Monterey Bay specimen, X10. AD,
adambulacral spine; A, actinals; 7, inferomarginals.

2. Same; large specimen of forma brevispinus from Lopez Island, San Juan Islands, Wash.
An adambulacral pedunculate pedicellaria, with a few accessories, 25.

2a. Same specimen, two views of a large actinal axillary pedicellaria, 4 mm. long, 12.5.

2b. Same specimen, abactinal pedicellariae, X25.

3. Same; fo ma paucispinus, from Nanaimo, British Columbia, having very numerous
“major’’ pedicellariae; side view of an intermarginal, 1.6 mm. long, *25.

3a, 3b. Others from intermarginal groove.

3c. One of the smaller abactinal ones, 0.96 mm. long, 25.

3d. One of the larger actinal pedicellariae, same specimen, base of ray 2.4 mm., 25.

4. Forma brevispinus, Monterey Bay; an adambulacral pedunculate pedicellaria, 2.4 mm.,
X25.

4a. Same, Monterey, intermarginal, two views, 1.28 mm. long, 25.

4b. Same specimen, actinal, 25.

5. Pisaster giganteus, type; a slightly twisted adambulacral pedicellaria, 25.

5a. Same, Monterey specimen; adambulacral pedicellaria with one of the smaller of the
cluster drawn to scale, 25.

5b. The small pedicellaria of 5a (from an adambulacral bouquet), 200.

333

PLATE 79

Ficure 1. Pisasler ochraceus forma ochraceus, Puget Sound; mouth plates and proximal adambula-

crals from side, 10.

2. Pisaster brevispinus forma brevispinus, San Juan Islands, Wash.; cluster of spines on
an abactinal prominence, > 10.

2a. Same specimen; mouth plates and adoral carina; eight pairs of proximal adambulacral
plates lack spines.

2b. Same specimen, an abactinal crossed pedicellariae (0.24 to 0.27 mm.), 200.

3, 3a. Pisaster brevispinus forma paucispinus, Monterey Bay, abactinal spines, 10.

3b. Same, Barelay Sound, British Columbia, x 10.

3c. Forma brevispinus; abactinal spine, 10, from large specimen, somewhat intermediate,
San Juan Islands, Wash.

4. Pisaster brevispinus forma brevispinus, Monterey Bay; crossed pedicellaria, abactinal
surface, 200.

4a. Forma paucispinus, Monterey Bay; skeleton of young specimen, baseof ray, 10
C, carinal plates; DL, dorsolaterals; /, inferomarginals; S, superomarginals. The
madreporite is shown on right, at top of an interradial line.

5. Pisaster brevispinus forma paucispinus, station 4219; a large crossed pedicellaria from
inferomarginal spines, 200; abactinal pedicellariae similar to 4.

6. Pisasler ochraceus, Monterey Bay; a furcate pedicellaria 0.25 by 0.25 mm., 100.

304

BULLETIN 76, PART 3. PLATE 80

U. S. NATIONAL MUSEUM

PISASTER OCHRACEUS

FOR EXPLANATION OF PLATE SEE PAGE FACING

Pate 80
Pisaster ochraceus forma ochraceus, slightly reduced

Head, Calif.
Bay, Calif. These are characteristic of the open, exposed, rocky coast.

Puate 81

Ficure 1. Pisaster ochraceus forma confertus, Boundary Bay, British Columbia, an extreme exam-
ple, X1.
2. Pisaster ochraceus forma confertus-nodiferus, Departure Bay, British Columbia,
3. Pisaster ochraceus forma ochraceus, Sitka, Alaska, No. 5984, 1. This is regarded as
typical of forma ochraceus. :

336

U. S. NATIONAL MUSEUM

28

63160—30

U S. NATIONAL MUSEUM

BULLETIN 76, PART 3 PLATE 82

PISASTER OCHRACEUS

FOR EXPLANATION OF PLATE SEE PAGE FACING

a i Puate 82
ag
Pisaster ochraceus forma nodiferus, Tongass, Alaska, 1.
ter ochraceus forma conferlus, Departure Bay, British Columbia, 1.
aster ochraceus forma confertus-nodiferus, Departure Bay, British Columbia, 1.
isaster ochraceus forma confertus, Departure Bay, British Columbia, 1.
337

PuatE 83

Ficure 1. Pisaster ochraceus, Monterey Bay, Calif.; cleaned to show abactinal skeleton.
2. Pisaster giganteus, Monterey Bay, Calif.; abactinal skeleton. ;

338

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BULLETIN 76, PART 3 PLATE 84

PISASTER OCHRACEUS SEGNIS

FOR EXPLANATION OF PLATE SEE PAGE FACING

*
Mia °
7 , ”
_
Puate 84
Pisasler ochraceus segnis, type, Lower California, 16 miles south of international bound-
7 (ary, “1.
2. Same, epecimien with two regenerating rays showing structure of forma nodiferus, 1.
Eos : : 339
®
s

Puate 85

Figure 1. Pisaster ochraceus forma ochraceus, Monterey Bay, alcoholic specimen, 1.
2, Pisaster giganteus, Monterey Bay, X1. Specimens from rock pools of the intertidal
zone are generally of this size to 50 per cent larger.
3. Pisaster giganteus capitatus, San Diego, Calif., young specimen, <1}4.
240

U. S. NATIONAL MUSEUM

> a

ee) *

*e,

PISASTER OCHR

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 86

12-16. PISASTER BREVISPINUS

1-9. PISASTER OCHRACEUS. 10. EVASTERIAS TROSCHELII 11. PISASTER GIGANTEUS.

FOR EXPLANATION OF PLATE SEE PAGE FACING-

Piate 86
Young stages of Pisaster and Evasterias, 1

‘Figures 1-9. Pisasler ochraceus, Departure Bay, British Columbia. Forma nodiferus, Figures 1,
: 3, 5, 6, 8 (alcoholic). Forma confertus, Figure 2. Forma conferlus-nodiferus,
Figures 4, 7. Forma ochraceus-nodiferus, Figure 9 (alcoholic).
a 10. Evasterias troschelii, Departure Bay, British Columbia.
a 11. Pisaster giganteus, Monterey Bay, Calif.
12-16. Pisaster brevispinus, Monterey Bay. All young P. brevispinus are forma paucispinus.

Figure 12 is an alcoholic specimen. at

PLATE 87

Pisaster giganteus, Monterey Bay, Calif.

Figure 1. Six-rayed example of the slender-spined forma from shallow water offshore, >< %4.
2. Forma with heavier spines, although not an extreme example, 44. This is a well-fed
specimen from the piles of a wharf whence it had crawled from shallow water.
3. Actinal surface, base of ray,

342

U. S. NATIONAL MUSEUM

BULLETIN 76, PART 3 PLATE 88

NATIONAL MUSEUM

U.S.

PISASTER GIGANTEUS CAPITATUS

FOR EXPLANATION OF PLATE SEE PAGE FACING

Priate 88

Pisaster giganteus capitatus, X'5{5

Fo rma with more widely spaced and heavier spines, Lower California, 16 miles south of
__ international boundary.
2. Forma with slenderer, more numerous spines, Venice, Los Angeles County, Calif.

a

PLaTE 89
Pisaster brevispinus forma brevispinus, natural size

Ficure 1. An extreme of the multispinous variety from Bolinas, Marin County, Calif.
2. A large specimen from Monterey Bay, Calif.

344

U. S. NATIONAL MUSEUM

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 90

PISASTER BREVISPINUS

FOR EXPLANATION OF PLATE SEE PAGE FACING

Piate 90
Pisaster brevispinus
e Forma brevispinus, actinal view of Plate 89, Figure 2, Monterey Bay, X14. A!-A?,
_ three series of actinal plates (spines); J, inferomarginals; S, supero
2. Forma brevispinus, giant specimen from Lopez Island, San Juan Islands, Wash., *4.
3. Forma paucispinus, typical, shallow water, sandy bottom, Monterey Bay, % Ls.
i 345

PuatTe 91

Figure 1. Pisaster brevispinus forma brevispinus. Giant specimen from San Juan Islands, Wash.,
having much shorter spines than Plate 90, Figure 2, and intermediate with forma
paucispinus, X45.

2. Forma paucispinus, Nanaimo, British Columbia. A subforma with very numerous

abactinal straight pedicellariae and few small abactinal spines.
346

U. S. NATIONAL MUSEUM

U. S. NATIONAL MUSEUM BULLETIN 76, PART 3 PLATE 92

PISASTER BREVISPINUS

FOR EXPLANATION OF PLATE SEE PAGE FACINGe

Puate 92
Pisaster brevispinus forma brevispinus, 1%

Monterey Bay, Calif., specimen close to type in general appearance.

tation 4501, Monterey Bay.
Station 4501, typical.

347

PLATE 93
Pisaster brevispinus

Fiaure 1. Forma paucispinus, Monterey Bay, 1.
2. Same, station 4492, Monterey Bay, unusually few spines,
3. Same, Barelay Sound, British Columbia. This is ,Verrill’s ““papulosus,” Xl.
4. Forma brevispinus, San Francisco Bay. Variety_with fewer than typical number of
spines, intermediate with paucispinus, 1}.
5. Forma paucispinus, young, station 4563, <1\M4.

348

U. S. NATIONAL MUSEUM

INDEX

The following index contains the names of the families, genera, species, subspecies, and formae

of Asteroidea which occur in this memoir.
synonyms, or synonymic combinations.

Roman type indicates valid names; italic type indicates
When a species name follows a genus name that is a

synonym of another genus name, both the genus and species names of the combination are italicized,
although the species may be valid. Heavy-faced type indicates the pages on which descriptions
or revisional a may be found. Names occurring in the explanation of plates have not been

indexed.
Page
Adelasterias._-..........-_.-.._ 136, 219, 220
papillosa ss: <2 sees ck 220, 246
Alexandraster mirus_..-.....--.------- 194
POSER LEN see ee ee te 2 es 5, 6, 9, 22
Whomald ce ao cee 6, 19, 204, 212
ON ECIOS Me re en ene 19, 23, 205, 213
OTMON OM = oo een oc ees See 205
MONLONENS TSS eee enna eee 6, 204, 214
RN a ee = See 6, 22, 204, 214
Allostichaster: --.=..-..=--.-- 24, 157, 217, 220
TRU sees ee eee eee 221, 245
Neébeghe ss 4a ee ce 221, 245
macqualige-=.. 54... 2 faa aee 220,245
PRIS eee a ee Se ee 220, 245
oly plax ies <2 22 fos - Sees 220, 246
TEP MIR IIR Sn fee Soe 221, 246
PT ELASLEL ore eee ee eee 135
BURGUIS een Hen aes aa So ee ae 201
INTO PIUR Sse ee na oe eee ea 201
POEUN Se ee 201
Annsteriags00—--2--225- 20 219, 221, 223, 235
MINNA: ans a oes 221, 222, 233, 241, 245
pedicellarie=—_---=-._--= 223, 224, 225, 246
Anasterias adeliae_-_-..-------------- 236, 244
AEC CO res So See ae 241
belgicdes sere we a a 236, 244
CHSTODNON Ga a a a 236, 245
CUINLN EY eae ae oe ae 236, 245
(ACESS A Se ae ae occ an ae ec os 236, 245
ETE is SS ee eee 236, 245
CLOT GOULD Sn or oe 234, 245
REEMA oo wee 236, 246
CHUET A eee oe eh oa nee 236, 246
ROLE Tie te eR pes res at a Se 236, 246
URCUTAU GS en mae eee ce ae 237, 246
Anteliaster coscinactis_-----~-~.-------- 201
GEG PONV ae es oe a = eee 201
maNnNOGEs= =.--=--=-. Be ees eat 201
MB PNANANGEIOR oo nen = 4, 159, 204
MYCNOPOUIA§ o= = —= eGo aan ae 160, 204

Page

pphelasterisys ooUse5 3 cece 4, 160, 204
JAPONICA oo Soe ok wat epee 160, 205, 213
PORQUaUE 25 25>+ 45.7. canuannoee 205, 215
A steracanthtione.-2s 4-5 een eee eee 5, 206
lind So Fo eee 156, 157, 209, 212
antarclicum. — .~ -<<.--none<npaue 204, 244
GUrOninoUe= = oon eee 241
SOT Fs BE olor cet ceed hl a ached 207
DOP TR ee oie oid ope an ae 205, 213
ECAURIACHONCUS oa = 2 8 eee 91
Claman oe eee at 227, 245
UIC G ean Ge eee 208, 213
OP DORE. oe Sen, Se 205, 213
WUlgeis = Se a eh ee 241, 245
LOD eel ce Aa 245
EPIC Tn ila ere on ee 228, 245
groenlandscua. ..- «= 2-2 oboe 45, 213
RRINGE > ene cee pees 77
TER CR ee 6 cre eee 211, 213
Uillnr alias. on ie aoe 207, 213
LaPeer ie 227, 245
MOT OOTY OF re os on ae 162, 164, 214
SRACICOTNU Wt ce sce ee 24,214
STILE LLIN cs wegen asc a 23, 205, 214
WYO Eee eo ape 227, 245
MILLET G2 Aes Senta 23, 208, 214
novaeboracensis......-..---.----- 205, 214
DRUG a a ene as 206, 214
DOLOTE oc Soe e enn oan aan 60, 61, 214
poppe. ==. Seco Hain oednee nme 220, 246
DT OOGRUES Jc errata arc a . 157, 214
VLD OH a gi ore csp ee rey 206
BpOCGD Ooo oan cocee ae 227, 246
Molioniira. 3 6a ce eneneee 211, 214
SUE ec pete eae kee 3 227, 246
OATRUNE So 6 beac neces . 241, 246
(OLAOUUR 6 aio ace nena ace aen 206
Asteracanthium-_--.-....---.----- x 5, 206
camischalicum ...-....-.-.----- Sud 91
Cis CR no nee 0 ve tee ee 206, 213
DOROENNS 2 3 onc ween an ees snes 57,214

350 INDEX

Page Page

Asteriading=22 2225-555 ee Soe ee 2;-200"| A'siertasigraczis == = =e 209, 210, 213
Asteriass< 2-2 22-4 se2s255 = sece ess 3, 5, 205 Granyerd- 2=-- = a= aoe 242, 245
amurensis_-_------ 6, 8, 12, 69, 204, 205, 212 OTOETIONOL CO == eee 45
ANOM al ane ee ee a= ee 13, 15; 19, 204, 212 quincy 211, 213
forbesiz- Ss == eee 6, 205, 213 heraciiss ase ct ese 89, 107, 115, 213
MU Prabal se ee oe ee ae 205, 214 hispidas eee ane ee 208, 213
rollestont==———--e== 6, 7, 12, 22, 23, 205, 214 holsaticaes aes Pe eee 206, 213
TUDENSe s-seb See eee 6, 8, 17, 22, 205, 214 hyadest 235.2 5-8 SS eee 241, 245
VWersicolor..=- 223225 Se See 6, 206, 215 hypervoreas ae sea 24, 45, 207, 213
Vilgariss= 22 cso shea a ea aee a= 6, 8, 206, 215 UNORINLS eee eee ee 45, 210, 213, 216, 245
Asterias acanthostoma___---------- 140, 148, 212 Janihind= =e ee = ee 164, 167, 213
CCOnUC LO 62, 66, 67, 98, 208, 212 FOPONICW ss an sa een eee 205, 213
QOGUaliS ea ne eee Aa e 89, 120, 212 hathemmoees==- === ee ene 77, 78, 164, 203
OUISKONS1S ae ee 90, 124, 212 laeingatige 2232523252 Se ee 226, 245
Glbgta = as ae eee eer eS 230, 231, 244 Wilonauisec=2= ne en ee 207
Qlbula= =e. Se 5 = es ee 157 longumang= soe 2a sae ee 45, 214
QNOMGIG 23s = ease a See 6 ithenviane se ee ee 172, 177, 213
antanctica: Se 2S == = a= =e 241 MOCK O PONG a ---- 90,91, 138, 214
GnChC@i = sa. = ae ee ee oe ee 24, 212 MONGOTUATENQae == a a ee ee ee 167
GQRCNtCOLG ee 205, 213 MLETLOTON CLUS mae a ee re 231, 245
QUEONILOCOM === eee ee ele 241, 244 MUNULG= 2 32 oe eee ee eee 206, 214
OUSLET Oe yen re ee ea 207, 213 MOIS sade a Soe eee eee 231, 245
Gellz See a a ok ee a Se a 230, 244 MULLER ee ne A ee ee 24, 208
bonbonicdi ean. ane ae ee 216, 244 MuUlitCclavise= =e a 90, 91, 203, 214
DORCAS = se ae ee eee 61, 62, 213 MUTTON Vase ea sce See eee ee 206, 214
OTA ChtAtG ss ee 140, 144, 148, 213 NANUMENSIS aoe ea ee oe eae 203
brandon. ssa ee 216, 230, 245 NUON ee 24, 214, 217, 245
DRED TON. Ge an ee 210, 213 neglecta = van te se a= en 230, 245
bnevispindss= == eee 180, 213 MOT MUU es ee 207, 214
COMISCNONCOn ae = ee eae 89, 91, 213 NOTLOTMLENSUS ee 2 ee ee ee 6, 14
COPENSIS = eS ans ee ee 228, 245 TUG ites Se ee ae ee 248, 245
CY OOO 1 a es 177, 213 ODUVUStSDIN0SGa = a a 230, 231, 246
Clathratas +22) 2S .8 = 2 ee 205, 213 OChOteNS (82 =e ao se ee oe 57
COMP =33 eas ee ena R oe 208, 213 OGNNO CEOs =e ee 162, 164, 214
CONS CRL oe eee Se ne 164, 168, 213 MOLIGG sets ae ee ees 206
CHLUNOMI GS a= eee ee eee 45, 213 DANO pla sss ee eee 210, 214
CUNTINGRUMN == = ea ee 241, 245 PODULOSG—= 3 =e Ee eee 180, 214
GOUGlLaST ae ee ee eee 77, 78, 79, 82, 213 DOUCIS PING == ae 180, 184, 187, 214
CU0t0 ao ee ae ee eons 77, 213 MOCUNGL Gs sae aa e ee e 6, 214
CRO DIG ae en ee eS 211, 213 DEGLCEMOnIS ae = eee ee 244, 246
epichlora__--- 90, 91, 139, 140, 144, 147, 213 DETECT eis ae aes ee a 241, 242, 246
CLQUISI OS aa ee 162, 172, 213 DAUD DUG ee a 62, 214, 217, 246
WObrMCits.— == sae = ee eee 206, 213 POLOMS a = = 60, 61
aSCLCULOniS == eee eee ee ae 24 poly plage. oa eee 220
fCNNONGENSIS = a ere ee ee 227 DOlthelL Gao ee 62, 71, 203, 214
SSIS DUNG see ee 164, 167, 213 ROOUSIG =) pee eee 211, 214
floccosa _---- Ee eee See eae 209, 213 TOSCO 22.5 See oe win = See ee ee 209, 214
nag iste ae = oe ee ae 217 TULDENS st Sea See ee 6
SUNG erdsaae- fe ae A eee eee 242, 245 TUS oe = eae ee ne Se ee 140
Qe0rQtanG) 2 Ses eee = eee eee 237, 245 PUGISDING sue ea oe ee ee 240, 246
Gigdtlea= 2s ae ee ee 172, 177, 213 MUUTLCOLD a sees ee 241, 242, 246
glactalise sonst eee 206, 213 SQUNACHENSIS aa aa ea ee 140, 214
glomenataS: <2 - oe es eee a 230, 245 SAISON eee ee eee 214

INDEX 351

Page Page
Asterias CUA RN oo oad 269 || (Brisinpidae.< 22a eee cee ee
MTV Seo ee mon ee 201214) SMR PINRS soe oe es Sele eee nee 200
Similispinte.-----.-----.--.- 59, 208, 214 Bunodasler ritlert__..........-.-.--=--- 189
REUSE en eee eee eho eke:| Calisastentie. 5 -- ccwennmaancnaseaee 162
BSUMNIORG nea momento eae oe) eld exquisila__...-..-.. nit ata howe pate 172
STATAUIIS sono aan <— 222, 241, 246 | Calvasterias._.------.-.-=---~=---=- 218, 225
spitzbergensis BSG sss ease 23, 45, 47, 56, 214 antipodumie oa bane soee =e 2, 226, 244
steineni eS a ee 237, 246 asterindides.. 6.5. ace n bana 226, 244
SIMADSONY sce oe aa oe ae 206, 214 AB VIR Acne we ier ieee 226, 245
iti Bs? eee eee ae 231, 246 BUONO Oth 2 nee ae .. 2, 226, 241, 246
SUN CUT EY Oe ee ee 229 Biter eset wei ae ann .---- 226, 246
Lene 3 one ee Sees SOB O15 Galycusleres . a2 ans a neon Seema se
LEVIES ee ee ese 217 MONOECUS.. aoa accceacla mR ko Oe
fomidata...---.------__--_-.. 228, 246 | ‘Ceramaster arcticus_..._-.-.---.=..- - 193
OTT Ak Cet te I 205, 215 Clarkt = che ocean ane eee ee
CHL ee ee ee ee 239, 246 JADONICUS:2.—222e=-cacesecoasedee ee Ae
froscheltt» .-=-2-=--=-=-=-=-=- 139, 141, 215 leplocerams=— 0-6 ae ee ee 192
RONCOUVCT Gs = aa s a 89, 115, 215 DAtASONICUAS ceo ae ae 192
DENT oe aan ne 241, 246 | Cladaster validus------.-.------------ 193
DERCHIOSO sess eee ee 5301046; | (Cosliatentas:se— ee tee a ea ee
O1cloniang.=.-----—==— 140, 141, 143, 203, 215 australss. <.6.--25.ccdeeast wags ey eee
ROL Ced ee sa ea 206, 215 |: Comastertas___...--_---=--.---------- 226
RUE AG= os eee ee 2, 201, 203 SU Gs oa ee ee 226
ARLEMINAC Soa == een oe 2, 3, 202, 204, 216, 217 | Coronaster_-----------~----~---------- 135
ASLEriNU Nini Ob=- === = 2=—--=-- = 193'|{@oxcinasteriase==- —- = =-— == see ee 16
JAG aii Pr eee ee eee 221 | Coscinasterias brucet___.--------------- 231
papillosum._---------------=--- 223, 246 WiCLOTOB Ss oka eee ee eee 231
Astrocles actinodetus-_----------------- 900 | Coscinasteriinae---..--.-------------- 201
Astrometis californicus - - - ---~---- 190, 202, 204 | Cosmasterias------------------- 216, 217, 227
eye 0 te oe ee 202, 203, 204 CApensIs.. ~~ 2. << aca e aoe eeee 228, 245
Astropecten armatus- ----------------- 189 Gysorith sooeee eee eee eee
[ped hae ee 190 felipes.....--------~»-----====2=5 228
Galil ON CUS sees oe a 190 Jernandensis__--.----------------- 22
BRINGS ae fe ee ee 189 germaini__-...-....--.------«--- 228, 245
DENS LEO Niles eee = ae eee 190 Wide 2 oecpeccacs shee Deamon pees
OVNatissIMUBS-—--5—2—=~5—-----—--~=-- 190 polygramma. --.--------- _... 228, 246
RIG AIIOINSS (ot ek aoa aa Sse a—he 189 TACIRUAS seer eee es eakaae _. 216, 227, 246
VERE [eee ee a ee ee 190 suletferd.- =.= <<-<-----«-.=5===- 227, 246
Astropectinidac.-_-------------------- 188 (Ofna Gla ena enn . 228, 246
Astrostole platei-_--------------------- 227 | Craterobrisinga synaptoma- --- Sune
iitanteriaveeee a wesen see ae ed Cribrella spiculifera—------- Sakata eae
bongraini_ ~—_---------------- “___ 244 | Crossaster papposus_-.-~.------ ..- 198
pedicellaris--------------------- 244, 246 | Cryptasterias--.------------- .. 219, 228
Bathyasterias ------------------------ 231 brachiava. -...--.-<-.<~ 229, 245
vesiculosa...------------------- 232, 246 turqueti._........----- ... 228, 246
Benthopecten acanthonotus------------ 191 | Cryptopeltaster lepidonotus- - Be - 193
GIRVIPelseese nae ose ae a= 191 | Ctenaslerias -..----------------------- 23, 51
MUtADUhe aos cee aes e ee ee 191 GAOT OT ann dnaee came sass _... 45, 204
Benthopectinidae - - -- ----------------- 191 georgiana ...----- ae = --- 27
Blakiaster conicus.-..----------------- 189} Ctenodiscus crispatus.-.- _ 88
Pe ei a ae eee ane ceana~, 180, 190 Cyllaster clarki-..-- isks a 196
Brisingella exilis- --------------------- 200 polyacantha - - - - - : 196
pannychia-----.----------------- 900 | Dendrogaster-_-.- 5 ee 53

pusilla. ge ----------------------- 200 astericola - Liked a reetiole 53

352 INDEX
Page Page
Dermasterias. imbricata___-_....-----__ Osi (Gastraster= 22) ==-- 5 4=— 5, 206
Diplasterias____.- 6, 216, 218, 226, 228, 229, 230 mMarcaritaccus= a —= esas === ae 207, 214
brandtin 236 2) ee ee 230, 237, 245 SUUCChIR eo an ae eee eee 216, 246
Druceincts= re ees ee 231,231,245) | Gephyreaster swifti--=-__=--_--=="5- == 3 192
Cpichlon@sas = ee ee ee _ TALON Gy hGSten mes ete eee eee 189
fochi® = 22223-0252 ee ee eee Zale 24oi | Gonlasier dace ==.55. 5 eae oe ee 192
Georgiana 22 soos oe ee Se ye2o | Goniasterinaess= == == =e ee eee 192
Germain 22 os os oe eee ee 228 | Goniopectinidae _ - _ __ ee ee 188
TNO eee 228.245) Granastenees= 222) eee ee 218, 232
kerpuclensisve == ese =e eee 231 Diserialuse ae e— eee eee 233
Loven t See ae ern a ee 2, 29, 245 DUPLiNS he were oe ee 233
litkentoes 2 ee ee Pep PA PAY ae Moyeniq oa), sachs calls} ey ee ee L 148
MeLiGClONaliS= ee ee 201;)245) |) Heliaster kubiniji- =2= 22 -S925 2-2 =e se 202
OPULOSU = ae = = Bee a ee eee 22246) lMemiastentas= a= see eee 232
SPULOSC ee ees a 216, 229, 246 brseniatuse. akan aes ee 232, 233
Stewment 2a sa5s tenes eee es 22925) MAennicianaleutlCaesa= 2am a= eee ee 196
SULCIfeT Ose ease ee asa Bae SA ae 229 anmectens= 262 =—72 22 ane eee 196
UT ULL te ae ee ee ea coe 228, 246 anclica tt oes UE ae ees 194
Vesiculosa2- = 252 s+) Sn eee eS 2 232, 246 aSPClate ee eee ee 196
Diplopteraster multipes__...---_---+-_- 199 asthenactis=.=cs-\2-2 soe =e 196
Dipsacasteranopluses-— ==. a 189 bonedtista sae eee Pee WN 194
IDOTCSIS se ween ate ee es eee 189 Claris) 6) 3S ea ee ae eee 196
OxIMIU S's se see Li Sees ee eee 189 dyserita_____- Set aba Stee ee 195
Le tm O DAU S eee = ee eee 189 CSCHUIGIL Yee eee 194
Distolasterias chelifera_.......--=-222 = 204 WNOCTMAMS = fap eee ees 195
NIPONSs sais so See eee aeeeoe 201 lacviusculass= 2222 eee Sees eee 195
Dytaster demonstrans------------+-- 190 longispinad === == eee ae 196
gilbertis. <2 235 5525% Ss = eee 190 ivilamens & ones San Se 195
Hehinastenrobustus====---=—=—--==——=— 194 MULLIS PING te a ee 195
SQNUUS* = = Les asses Seca e eee 194 POlysc snibh ass 196
(enUispINUS= 322 eo = 9 see ee eee 194 sanpuinolenta-2-o ase ee 194
chin astemd ae ss" =e eae eee 194 Spalulufend = seam e oa e ee ee 195
Hremasteniass 22a o eee eee eee 219, 232 spiculifera Ro RS Sa SSS 195
anbarcticale 2 =i ceed es pees 232, 244 tumida- - ---~-------------------- ros
Bremicaster pacificus..----._--_-:=_-_ 188 Heterozonias alternatusese- === == -Seeee 198
“ liexasterias= 1-2. = ss a<seeseaee= =e 59, 203, 208
(LENE DTATINS See ee ee ee 188 - :
: Hippasteriavarmataa==s—==—os = ae = 193
HV ASUCTIAS= aS ae = are eee ele 3, 43, 139 BA eee
Californi Cale seta Se 193
ACANUNOStOM Ye see = Sa ae ee 94, heath 193
139, 140, 144, 148, 204, 206, 212 nes Bene ie. Se Oe eae ee
kurilensis® -<=3. = ec wo pee eee 193
alveolata=ae=— aes aes ae eee 91, s
lelopelitat.) "= = 22 e eoe eeeeeee 193
94, 139, 140, 144, 204, 206, 212 ale :
Spinosad= 5255255 322 ee eae 193
CONS es =e eee ee 140, 204, 213 . ; ‘
: Ey drasterias’=--5= === ees ee 163
echinosomae. += s2=-—--= 139, 152, 206, 213 “ er An. 9
Sette DONT ULL ee a 210, 215
DOWISDINGH = = a 140, 144, 148, 204, 214 Hem cnaster kochleri 199
UL See eee TOUT 144 204 2S eae peters en ES ls ene Cn 199
subnodosa_________- 140, 141, 144, 204, 214 Ny as eee alae A aE MEAS Gente LT ae
ee Quadnispinosusses2e == === == 199
UO C1 1 ese a ee 94, Icasterias 4. 210
9 9 9 29 9 Pe a he EOP ee eae ot Oke Bee ¥
125, 189, 141, 208, 204, 206, 215 Acie eae ead pine
Freyella insignis - ------------------ --- 200! Kalyptasterias._.__.-------- 219, 221, 223, 233
microplax_— ~~~ --~---------------- 200 conferta_____- PSA 223, 224, 234, 245
Freyellaster feeundus: = === .---- 42 -2-+- 200 | Koehleraster_______.______- 219, 222, 234, 245
Borcipulata: = 325-5 ee eee 2, 200 Octoradistatss=2- 2-2 eee 234

INDEX 353

Page

Page

Korethrasteridae_..................... 198 Leptasterias nana. 90,119,120,123,203,208,214
Iheptasteniass.2-f 9205-2 3, 23, 207 nesiotis._____ ... 94, 96, 209
ACCU VAUR re sic ee a 30, 59, 60, 61, 62, nitida___ 94, 95, 103, 108, 113, 124, 203, 200
66, 71, 94, 100, 125, 203, 209, 212 oblecta._................. 45, 56, 204, 214

CCL SNIR eae ae eee eee 59, 89, ochotensis._____ . 1, 24, 57, 208, 214
90, 91, 101, 103, 109, 113, 114, 115, 117, orientalis. ___ ... 1, 24, 40, 44, 208, 214
118, 119, 120, 123, 188, 203, 208, 212 plena_____- ... 90, 107, 108,
CNS an) es en ee 59, 112, 113, 117, 124, 203, 209, 214
j . 70, 90, 91, 97, 103, 105, 107, 108, 112, polaris_............ 59, 60, 61, 62, 209, 214
124, 125, 130, 131, 133, 203, 208, 212 polythela__.._..._ 62, 70, 71, 203, 209, 214
aleuticnZo. .4=--2-e lo detisls 59, 101, 208 pribilofensis._......_..._- ... 129, 208
aphelonota-_-----....-.- 60, 62, 63, 70, 209 pugetana..______ 96, 107, 111, 113, 203, 214
BNChiCNHes Atos 2-22 = be 23, 26, pusilla__________ 59, 108, 118, 123, 203, 209
35, 42, 50, 51, 52, 56, 135, 203, 207, 212 regularis._.._.__.._.._ 90, 91, 107, 108, 109,

BRA Cae eee 59, 98, 128, 131, 208 110, 111, 113, 116, 203, 209, 214
BapenSs se 5 ee 91, 107, 109, 110, 207 shumaginensis__ __ 127, 128, 130, 135, 208
BUG AP ee ee ces SS 44, 59, 103, 208 RIQGYOR ee Ac oad SOs Gia ee 90,
MURGELAS (ee Kee eee 23, 207, 213 107, 109, 112, 14, 1 24, 203, 209, 214
benim Pensige = ses ae eee 26, 207 BIMWISHIMIG> 22 ae eee ese .. 59, 208, 214
borealis. 2-_=~..- 59, 60, 61, 62, 82, 209, 213 StOlAGHN tha a. See oe 135, 209
camtschatica_____ 59, 65, 89, 90, 91, 94, 103, plbAcervata. 2c = 60, 209
105, 109, 117, 124, 125, 203, 208, 213 subnodulosa__ 90, 91, 112, 125, 140, 203, 214
CONE ee ee ee 90, 203, 213 subregularis _-.___---_-_- 90, 111, 203, 215
(Co ee a 59, 60, 84, 203, 204, 208, 213 Llyn a 2 24, 44, 84, 135, 208, 215
COM PACU saeco ao 90, 120, 123, 203, 213 ITNCUENia a ae ee ee 59, 85, 86, 209
ON Ua see are oe 44, 135, 208, 213 vancouveri_. 89, 103, 109, 115, 203, 209, 215
CD TROT TU 90, 120, 123, 203, 213 | Leptychaster anomalus_-_-_--_._..----- 188
GhIDratiae snes cease 47, 56, 204, 207, 213 Srchious. 22.2 ae oe ee eee 188
MUR pire e ae Wo oe ee os 59, 70, 90, 94, 97, INENMUSi see keene s eee : 188
101, 103, 108, 125, 128, 204, 209, 213 MALES PING AS Sa cae eee ee 188
BPRGHLON GE cee eee on 90, 91, 124, 203 pacifieus_______- dis andere ore ae 18s
PBN etiee a wee 1, 24, 42, 207, 213 PYOpinquus._= oa a eee
Broemandica—..- =... --.-<8_-0=+ 24, | Lethasterias chelifera__........__-- . 202, 203
42, 44, 45, 48, 49, 135, 204, 207, 213 NENVUeNsIS.-. ea ee 202, 204

UNTIL Se Ee es Soe peere 24, 221, 245 | Linckia columbiae_- - ~~. --.---.---~-~- ; 193
MEXECHNE Soman ose bce 59, 89, 91, | Lophaster furcilliger___. .. - a iaheeate 198
103, 107, 109, 115, 116, 119, 203, 208 VEX OUOD a ee oie ecto a eer 198

TUS POLO CUL ED PAs ete a ictcsred w wt ccerssi coins 207, 213 | Luidia asthenosoma- -.--- ase rea 191
vlog esas sneer a oe 24, 35, 41, 44, 207 foliolnte sno. ann eee ee 190
RSS KIC ea re el en 24, Od Wikie Rc cee s a oem e ee 191
34, 35, 40, 41, 44, 135, 207, 213 | Luidiaster dawsoni- - - - ~~. ~ - - 191

DURE UL ARES psa an Sas i wl 140):208; 213 | Duidiidse-s2- <2 22.5... <s 190
TESOL Gh Cake ene tee re eek 56, 213 | Lysasterias.._...... 136, 219, 2 2: 229, 223, 235, 236
katherinae __..........- 23, 30, 59, 60, 61, “adeliged 2. 6 oe cmenwates i 236, 244
70, 77, 85, 164, 203, 204, 209, 213 belgicae ____- pene nid 236, 244
leptalea__............ 24, 44, 203, 207, 213 chirophora- . - ~~~. ~- : 236, 245
Jeptodomass. = 282 aoee -Sonen ae 59, 105, 209 cupulifera —........- 236, 245
littoralis______________.- 24, 135, 207, 213 joffrei____. ne 236, 245
macount ___- ~~~ 112, 140, 141, 144, 203, 214 lactea . - - .- Ree : 236, 245
TTC SUCH NE eee ee eee oo a 24, 214 perrieri__ ; 236, 246
WRU Re Pr eee 90, 208, 214 studeri- - - - ame 236, 246
miilleri__ —__ 24, 34, 35, 40, 56, 135, 208, 214 tenera..--. aa apatite . 236, 246

te

multispina- --.------ 59, 127, 130, 133, 208 VIOCONAGe.2k eee 23, 237, 246

304 INDEX
Page Page
Lysastrosoma- ------- ashoo neta eeee 1135) | -Parastropectena == = = ee a 189
MESMIOLAS: 222 fee eee ee 202 IMeNINGS: = coe eos ope aee eee 188
Marthasterias sertulifera_-------------- 203) | Patiniaminiatae. = es 193
Mediaster‘aequalis= = .52-=----5-=. = OD) "Paxillosas=2* oof ae) ee cents s a ee 188
tenellusses- 6 ae a ee eee 192) Pectinastertagassizy=—o— = =e == a= 191
Miyondla x 220-1 ae ea ee 191 evopluse#lne 226 ~S-. See SSL 191
Myonotus intermedius---..------------ 191 | Pedicellaster magister_ + _--+_-------=-- 201
Myxoderma ectenes-_------------ Soe 200 MONG OMLACCUS= sae eee 207, 214
platyacanthume #2=2 25 one ee 201 mepalabis=s22- ok Sse ee ee 201
ThomaleuiM ss" sees ee 201 Bedicellasterinaeeys= a= === see = 201
sacculatuims === --— ese eee 200 |, Peribolaster biserialis_----.-.__-_-----= 198
INIGIUGS CT ee ee er ee 1'56)| Peridontaster crassus=_-=—---=-==-==3==— 191
Nearchaster aciculosus_-.-------.------ JOM tPerissasteriase. === eee eee ea 219, 238
pedicellariss 23: == == 5— =. eae 191 heptactislse- 2s) 32) oes ee eee 238, 245
Variabilisecs ome eee eae ee a= 191 ObtUsispinaee'= 2 ee ee eee 238, 246
INeomorphaster==-— = - =) == ee ee 5; 201 DOW ACN Yee eee ae 238, 246
CUSLLCRUS SE oa te ee ee Pie cllsu| meh aneroz Oa a= a= eee ee 188
TORCIPALUSSE eae eee eee ae Aro Si| PEISaS tetas ae ane: LE aoe ts 5, 9, 162, 209
taligm sniete= ae ae eee 212 ONAN CliCUS a2 ee ee eee 232, 244
Neomorphasterinae- - ------------- 3; 211; 212 qustralis== 2-222 164, 172, 176, 194, 203, 213
Neosmulastens sae = eee 218, 237 brevispinus-___- 164, 180, 181, 203, 209, 213
PCOMQIONUSe Se eee ee ae ee eee 237, 245 capitetisiees =) = === ee 164, 171,
StCINCNi ee! = = Sa 5-252 = Se ee 237, 246 173, 174, 175, 177, 203, 209, 213
INesastenase-s oe Sse ee eee eee 135, 208 GOMPeRGUS ee ee oe OA oO)
IN OUaSteTIAS ters sa eae oe ee 243 140, 164, 165, 168, 170, 203, 209, 213
ALIN GR ee ee ee an SO 244 SUSSTSIDUNUS aaa 164, 165, 203
naswelli =. 2 a5 shen = ae eee 244, 245 gipanteuses ess ee ee 139,
Notasteriinae_-___-------- 31, 210, 212, 216, 243 163, 164, 172, 177, 180, 203, 209, 213
Notomyotasee- == 2) ee ae ee eae 191 GRY ie See eee 77, 79, 164, 213
Odontasternicrassusss = ae 191 hithenve ee 164, 172, 176, 203, 213
Odontasteridae=s = 2555-2 es = ee 191 NOdIferuss. = — == ee 94,
Orthastenias=--5= 5-2-5 = eae 18 139, 140, 164, 169, 170, 203, 209, 214
LOR CUT a alee ee ee a eee 204 OChTaceuse2 aes ee 94, 139,
californicae Craw en oe heen 204 140, 164, 165, 170, 174, 175, 203, 209, 214
COUWMULON Cae ee eee ee See 204 CID UL OS UG a et ae 163,
AGwSON er ae ee 2 eee 204 164, 181, 184, 187, 203, 214
OnCUDULGIO= mee ee a ee 204 paucispinus____ 110, 164, 184, 203, 209, 214
Oren eee ee ee ee 204 BOPnISh= oA See 170, 171, 203, 209
GON0LCNON a a = ee 2045 eROdastentaseeere ae ee 6, 216, 226, 229, 230
Kkochlonios= =e eee ee eee 02204 ORONO apa oe ee er 231
lentolena= Pee Bes ee ee ke 204 briuceis © ee Le os oe ee ee eee 231
leptostyilsi= 45 ee ae ee eee 204 FOCKUES Asse ee ee ee 231, 245
merriami.....------ 88, 84, 85, 86, 204, 214 GEON OU CNGee 237
mOntereyeNsiSaa- = seo eee a aoa 204 Ken QUClLen SUS = =e a 231
Paedastentastss=- 4 ene ea Sa 235 MEnTALONGSS ee ee 231
ChYjO DUNS <2 ae oe a 236 steinent=_=- = Pek a). eRe 237
MOUT eve = aon eee ao 235,236, 2451| Poraniopsis flexilis= => ==-—--=—3-=====—— 194
Parastenias: 22 ee ee eee eee 5, 6 inflataee cree. ee oe ee eer 194
CIDCKIENS1S =e 6, 20, 22, 204, 212 MUO ee ee a ee 194
ROnOstiChastene see ees ae ee _. 239) Porcellanasteridac-_-_----- =o to So ae Bee 188
dimectss.- 22a eee __..... 241, 245 | Pseudarchaster alascensis__—------=-=-- 192
MO WS0ONt sO so eee 241, 245 GissOnls eee Oe ae eo eee 192
SURCEHULALUS == ee _ 241, 246 DALCieee eee ee eee 192

INDEX 355

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Pseudarchaster pusillus.------_-.-____. 192 | Sporasterias rupicola. —.—. --.--~ 241, 246
Pseudarchasterinae_.---_-_--._.--_____ 192 sphacrulata_ —_ . 241, 246
Pteraster arcuatus_—_........._...___.- 199 spirabilis_ 241
ROsCINODE DI swes= =a nena ea ae eee 198 | Stellonia ....___- .. 5, 206
OUOGIUAR see eee 191 hispida__. piss
eves ee at Poa eet ened ae 199 | Stenasterias._.____. ..-- 3, 138, 209
FONQHMNG ono pant ane eciete noone 198 macropora_.....- 90, 91, 138, 204, 200, 214
MIATSID DUS see na eee see 198 macropora ; Sit aRint es aol 62

TT AUS ee eae 8 ee eg 198 | Stephanasterias ‘ _...... 4, 156, 159, 209
PUR TNUGS oe a So oe mob wm 199 albula_......-.--....-- 157, 204, 200, 212
ODSCURUB See ae os ae ewe 199 RAGA ae a Ree kere . 156, 209, 213
DCLG re ee ets oe Aaa eh 199 hebes__ -- 221, 245
WULWUUS S ee eee s eee moo ee LOD VStiObAstéla sor en onan ene een . 217, 241
UNC 26 = a ee 199 albulus_—- ~~. 156, 157
HORSE AOUS Se= en oe soem a 199 QHD GUUS ao naan ee een aoe 210, 212
UMrOnGdOnS = seas — oe an w= 198 OY CREE nee ana ee ete eee 213
Plerasterides aporus_-—-.-------------- 198 UP ONNOCUG ste anon aerate ee 241, 244
DE SSNPS ic et a re 135, 136 BUISDOAUIS eee ee ae eer 241, 244
helianthoides.—.-..--..---------- 202, 204 Ny] UNOS ice a ete Be es ee 228, 245
IPUGnOpOGUNRes fee = ane mo o-oo 202 RTS QUIS doe nie ee a 220, 245
UTGRRUSher sete sea ee seo lr 226 Levi giles fot n> See a 226
AU ueh eee ee ee ae ee 228 NALOY GU oe oo ree ea ee 226, 245
IRSOIStENURG lea ao So tiki 191 Wee ee ee 157, 214
Rath bunaster. =. .2-=--=2-5. 22-56 135, 136 Wulpii ses aba cates aoa ae 232, 245
CAILOMGUSee= 2 ae oe 201 DOWGrGNUNNS = aoa — oe ao 228, 246
ALIAS COLS Ss] eee eee 218, 238 Poly ples an See eee 220
pre chiataoee = 25 SS == i ae 239, 244 POSES Se et ee ee oe ees 210
Sclerasterias heteropaes---------------- 201 BUTIRGUA oon one eee ere 241, 246
Nmlastenias 26 sae sa a 218, 232, 237, 239 Siar Err see eae a a eer eee 226, 246
RGRe GUAR aes ae a 239, 245 lalisnanisn~< sane sooes oe ee 212, 215
RCalpniterdeee see = ass s— a 239-2461) Stichastrellfs..-—= sone === S omen ae ee 5, 209
LOL Clee eee cn AR Sao 217, 239, 246 am Piguae sete fee ee 210, 212
Tye pan aes ee ee 239, 246 TOS@A2 eee a ee 210, 214
OLAR UBL A CUl Che eee ee NOT SStrclor ella ae oc wo ae ea 225
ied thee sere 6 ee ee 198 SH Br iio cee ee 226
CONS ENA S ee ee en a eee 107 | Stolasterias' brucei. --.-.---..--2..-. 231, 245
GsinO rem eerie ee re oe 197), || Stylastenitis. -.-=---=<22---<-Ssacennee Is
BnGE Ud eee eons anaes ees 196 fOYTeYi as nae ees Sern ch wae een ee 201, 204
CSIP UE See ee eae eee 197 | "Tarsaster. -.--<-- - 2-62 sce -e ene eH ene 24
(Uumneeee ere eee senceens |LOG BIBAKONAIS Sts a ae ee ae 201

by OUMrISsUS= == 2==— = ea 198 neozealanica.._.....------- _.. 220, 245

MUD DOS USE ei. == nae a= 198 |. Tarsastrocles___..--------- .... 5, 135, 210
OSRIUSUURE ea a Sen en 197 verrilli____- ane i . 210,215
AIM PSO = sae a ae ee a a= 197 | Thrissacanthias penicillatus. - - 190
NOSALEUIO AG wee ee ae a 196s) COVE ae ts re ...----- 24], 244
Sporasterias_--..------------ 219, 221, 223, 239 allantica_ -. 241
antarctica _----- 222, 223, 224, 225, 240, 244 | Trophodiscus almus-- 189
CLINGC Lees =p te ee Skee 241, 245 uber... --- ; iso
THAWEO Le aaa a a ae we 241, 245 | Uniophora- - . 218, 242
MCHC Osan a= == ==—- > ===-= 225, 246 dyserita. —- 242, 245
ELMAN a eee anos n-ne on=- 261s 246 fungifera : : . 242, 245
PUpIs PNG. =a. -—- -=-=~-~-------== 222 globifera__...-=------- 242, 245

63160—30——30

356 INDEX

Page Page
Wniophora‘gramiferac==22--5-2--6- se 242.245) | AUrastenasaen. sooee eee ee. eee eee 3, 211
FVMNOUs = Sees eee so ee 242, 245 enoplass- 2-22) a. eee ee 211, 213
multispindeas sees eee ee 243, 245 line} 2.58 oe ee eee 211, 213
MUU See eto Sa eee Be eee LOMO AOU e Oba eee oa - = ee ae Se ee 191
ODeSHt Sten) ea ee eee 24324151 Zoroaster actinocles=—.==2=——==s=s+4==se= 200
SINVUSOLdA eee ae ae eee 243, 246 evermannine --o. eee ee eee 200
TNISChIalISes see se ee ee ee 2A 20 MOTO AK S= ae = Se ea ee es 200
Unastensaa = = ana se Sok ee eee 5, 206 Ophiurus: 264=- = =322 6552545550 200
RSPIOW? — nse ate J eas een oe Seo Z208)| OLOsstenidses === = —=— eae =e ae 200

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