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ALLAN (HAN COCK FOUNDATION |
PUBLICATIONS
OF
THE UNIVERSITY OF SOUTHERN CALIFORNIA
First SERIES
ALLAN HANCOCK PACIFIC EXPEDITIONS
VOLUME 8
1940-1948
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1948
ALGAN HANCOCK FOUNDATION
PURE ECAT IONS
ALLAN HANCOCK PACIFIC EXPEDITIONS
VoLuUME 8
1940-1948
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1948
CONTENTS
ok PAGES
. A New Genus of Brittle Stars, Amphicontus by Alex Hill (Plate 1)... 1- 6
. New Ophiurans of the Allan Hancock Pacific Expeditions by Fred C.
ZAesemhennen CEL Aces 9) eases swe orca cee cee censor ee ees rasenencesacceceraneeere 7- 60
. The Holothurioidea Collected by the Velero III during the years 1932
to 1948 Part I, Dendrochirota by Elisabeth Deichmann (Plates 10-30) 61-196
. New Eastern Pacific Sea Stars by Fred C. Ziesenhenne (Plates 31-34) 197-224
5. A Report on the Echini of the Warmer Eastern Pacific, based on the
collections of the Velero III by Hubert Lyman Clark (Plates 35-71)
ACE weed DY Peg ess A as a a a es 225-352
REPORTS ON THE COLLECTIONS OBTAINED BY ALLAN HANCOCK PACIFIC EXPEDITIONS OF
VELERO III OFF THE COAST OF MEXICO, CENTRAL AMERICA, SOUTH AMERICA, AND GALA-
PAGOS ISLANDS IN 1932, IN 1933, IN 1934, IN 1935, IN 1936, IN 1937, AND IN 1938.
A NEW GENUS OF BRITTLE STARS,
AMPHICONTUS
(PLATE 1)
By ALEX HILL
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PUBLICATIONS
ALLAN HANCOCK PACIFIC EXPEDITIONS
VoLuME 8, NUMBER 1
IssuED JANUARY 10, 1940
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
Los ANGELES, CALIFORNIA
A NEW GENUS OF BRITTLE STARS, 4MPHICONTUS
(PLATE 1)
ALEX HILL
Research Associate, Allan Hancock Foundation
Among the several vials of minute brittle stars from the Hancock
Collections given me for identification, one contained a small specimen
from Velero Station 384 collected in 5 fms., three fourths of a mile off
shore, on the east side of Independencia Bay, Peru. The conclusion was
finally reached that the species was new and that it did not belong to
any genus in the collection. However, the shape of the disk, the outline
of the plates, and the small size indicated the Amphiuridae. A sketch and
a preliminary description were forwarded to Dr. Hubert Lyman Clark
of the Museum of Comparative Zodlogy, Cambridge, Mass.
While waiting to hear from Dr. Clark, the author found another lot
of brittle stars, from Velero Station 330-35. Careful examination of these
showed that they were essentially like the specimen from Peru, and one
was then forwarded to Dr. Clark. Station 330-35 is on the south side of
Albemarle Island, Tagus Cove, Galapagos, in 12 fms. This stop had
been made by Velero III, December 12, 1935, on her way to Peru.
Further study confirmed earlier conclusions, and Dr. Clark suggested
that the new genus be named Amphicontus (Amphi—in accordance with
the related genera—plus xoytos—a pole, in reference to the polelike
spines on the disk).
The generic diagnosis is as follows:
AMPHICONTUS, new genus
Amphiuridae with a thick disk covered by a large central plate sur-
rounded by five primary plates of equal size, accompanied by a number
of small scales. Radial shields large, somewhat irregularly oval, smooth,
in close contact with each other. Interradial areas each covered by 8-10
(or more) small plates and scales, three of which carry relatively big,
blunt, polelike spines; one of these lies close to the primary radial plates,
while the other two are on the interradial margin. Side armplates large,
bearing stout, blunt, not flattened spines of nearly equal length. Dorsal
and ventral armplates large, more or less pentagonal, notably similar
GIC4>
ee OS 3 sp
2 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
to each other. Ventral interradial areas covered with scales. Genital slits
very small. No tentacle scales or pores. Three pairs of oral papillae; two
outer pairs more or less vertical to the jaw.
‘This new genus resembles some of the small species of Amphiura in
the contour of the disk, in the superficial appearance of the disk-plates,
and in the shortness of the arms. It differs, however, in having 3 pairs
of oral papillae as in Amphiodia and in the presence of 15-20 polelike
spines on the disk. It also lacks tentacle pores and scales; presumably
the tentacles are themselves wanting.
Amphicontus minutus Hill, new species
Description.—Disk diameter, 2 mm.; arm length, 6 mm. Disk rather
thick and heavy, covered with swollen plates. The central plate slightly
pentagonal with rounded angles, surrounded by five rounded plates of
equal size separated from one another by minute interstitial plates or
scales. Interradial plates, 8-10 or more, the most proximal bearing a
strong, blunt, vertical, polelike spine which marks the proximal, median
point of the interradial space ; the other plates, more or less symmetrically
arranged ; of the two outermost each has a blunt, polelike spine, making
10 polelike spines along the complete margin of the disk. Two young
paratypes have a small spine adjoining the longer spine on its radial side,
making 10 long, inner spines and 10 short, outer spines in the interradial
areas of the disk-margin. No other important differences have been noted
in the study of the 6 paratypes.
Radial shields prominent, less than half the disk radius in length,
longer than wide, irregularly oval, in close contact along the straight
radial margins; interradial margins convex but irregular; distal margins
convex.
Dorsal armplates somewhat pentagonal with rounded corners, broader
than long, with a concave proximal and a convex distal surface to the
plate.
Ventral interradii covered with small, overlapping scales. Oral pa-
pillae, three pairs: innermost, thick, blunt, rounded triangular, not
closely in contact with each other; the middle and distal pairs are erect,
flattened, bluntly pointed, the width at the base about half the height,
the distal pair wider. Two blunt and rounded teeth on each jaw. Oral
shields pentagonal or diamond shaped, approximately as wide as long,
with proximal angle most acute, distal angles rounded. Adoral shields
No. 1 HILL: A NEW GENUS OF BRITTLE STARS, 4 mphicontus 5
longer than broad, meeting fully within, but well separated radially by
the first under armplate, which is rounded pentagonal, approximately as
broad as long with a nearly straight distal margin; subsequent plates
pentagonal, broader than long, with a proximal angle; lateral angles
rounded, distal margin scarcely concave. No tentacle scales or pores. Lat-
eral armplates with prominent spine-ridges, bearing 3 heavy, subequal,
blunt spines; spines longer than arm segment.
Color-—Cream white, with the following exceptions: some of the
primary plates of the disk and some of the dorsal armplates have splotches
of brown or nearly black.
Types——Holotype, AHF no. 3; 6 paratypes, one in the Museum of
Comparative Zodlogy, Cambridge, Mass., No. 5861.
Type locality—South side of Tagus Cove, Albemarle Island, Gala-
pagos, in 12 fms.
Remarks.—At the type locality, on the south side of Tagus Cove,
Albemarle Island, Galapagos, the records indicate that the bottom is
composed of coarse, black sand. Records also indicate that the dredge
contained one other echinoderm, Paulia horrida (Gray), and two species
of crabs, Stenorhynchus debilis (Smith) and Portunus (Achelous) angus-
tus Rathbun.
At Station 384-35, three fourths of a mile off the east shore of Inde-
pendencia Bay, Peru, the bottom was coarse white sand with an abun-
dance of sea weeds and red algae. This haul was rich in spider crabs,
Pyromaia tuberculata (Lockington) and Inachoides microrhynchus
Milne Edwards and Lucas. In addition to the crabs a number of worms
were in the same haul.
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ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
EXPLANATION OF PLATE 1
Fics.
1. Amphicontus minutus Hill, new species, upper surface of most
of disk and part of one arm.
2. Lower surface of part of disk and base of one arm.
3. Two arm segments seen from side.
All drawings by Anker Petersen, staff artist,
Allan Hancock Pacific Expeditions.
No. l
HILL: A NEW GENUS OF BRITTLE STARS, A mphicontus
REPORTS ON THE COLLECTIONS OBTAINED BY ALLAN HANCOCK PACIFIC EXPEDITIONS OF
VELERO III OFF THE COAST OF MEXICO, CENTRAL AMERICA, SOUTH AMERICA,
AND GALAPAGOS ISLANDS IN 1932, IN 1933, IN 1934, IN 1935,
IN 1936, IN 1937, IN 1938, AND IN 1939.
NEW OPHIURANS OF THE ALLAN HANCOCK
PACIFIC EXPEDITIONS
(PLaTEs 2-9)
By FRED C. ZIESENHENNE
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PUBLICATIONS
ALLAN Hancock PAcIFIC ExPEDITIONS
VOLUME 8, NUMBER 2
IssuED APRIL 12, 1940
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
Los ANGELES, CALIFORNIA
CONTENTS
INTRODUCTORY REMARKS
Famity HEMIEURYALIDAE
Genus Sigsbeia Lyman
Sigsbeia laevis, new species
Famity OPHIACANTHIDAE
Genus Ophiacantha Miller and Troschel
Ophiacantha phragma, new species
Genus Ophiomyces Lyman
Ophiomyces multispinus, new species
Famity AMPHIURIDAE
Genus Ophiophragmus Lyman
Ophiophragmus disacanthus, new species .
Ophiophragmus lonchophorus, new species
Ophiophragmus ophiactoides, new species .
Ophiophragmus papillatus, new species
Ophiophragmus stellatus, new species .
Genus Amphichondrius Nielsen
Amphichondrius laevis, new species
Genus Amphiodia Verrill
Amphiodia sculptilis, new species .
Genus Ophiocnida Lyman
Ophiocnida californica, new species
Genus Amphioplus Verrill
Amphioplus philohelminthius, new species
Famity OPHIOCHITONIDAE
Genus Ophionereis Liitken
Ophionereis dictyota, new species .
Ophionereis perplexa, new species .
Genus Ophiodesmus, new genus :
Ophiodesmus amphilogus, new species.
Famity OPHIOLEPIDIDAE
Genus Ophiomisidium Koehler
Ophiomisidium leurum, new species
Genus A mphiophiura Matsumoto
Amphiophiura irregularis, new species
Amphiophiura paucisquama, new species .
Genus Ophiolepis Miiller and Troschel
Ophiolepis plateia, new species
11
13
15
16
18
19
21
22
24
25
27
29
30
33
33
35
36
39
41
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NEW OPHIURANS OF THE ALLAN HANCOCK
PACIFIC EXPEDITIONS
(PLATES 2-9)
Frep C. ZIESENHENNE
Research Associate
Allan Hancock Foundation
In the study thus far of the ophiurans, which have been collected on
the eight Velero III cruises to the Eastern Tropical Pacific, nineteen ad-
ditional new species have been noted, twelve of which were collected on
the 1938 cruise when the new dredging apparatus of the Velero III was
used.
It was the writer’s good fortune to work with Dr. H. L. Clark,
Museum of Comparative Zodlogy, during the ten weeks of the 1938
cruise, and he wishes to express his indebtedness to Dr. Clark for his
invaluable advice and opinions and also for certain field and color notes
used in the preparation of this paper.
Thanks are also due to Mr. Anker Petersen, staff artist of the Allan
Hancock Foundation, for the preparation of the illustrations.
Hemieuryalidae
Genus SIGSBEIA Lyman
Sigsbeia laevis, new species
Plate 2, Figs. 1-3
Description —Disk diameter, 7.5 mm.; arms, about 38 mm. long.
Disk small, swollen, and elevated; disk plates swollen, of various sizes,
irregularly arranged with no central plate or rosette. Interbrachial scales
arranged in 3 rows, a median row of 3 to 5 large, irregular plates sep-
arated from the radial shields by a series of small plates. Radial shields
triangular, longer than wide, interbrachial margin longest, angles within
widely separated by 2 or more large and several smaller disk plates.
Between the distal margins of the radial shields and the arm base are
7 disk plates arranged in a triangle, transversely to the arm. The disk of
the specimen seems to have been crushed at one time, as the radial shields
at one arm base have fused into an equilateral triangle with concave sides.
First upper arm plate oval, twice as wide as long. Succeeding plates
[9]
he
10 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
swollen, pentagonal, about as wide as long, with proximal margins
slightly concave, and lateral and distal margins convex; plates barely in
contact. Side arm plates narrow, bearing 3 blunt arm spines, the upper
one rectangular, very large, not twice as broad as long, resembling a
supplementary upper arm plate. Under and median spines of equal size,
short, flat, rounded, and longer than broad. Side arm plates meeting above
but not below.
Oral shields, except the madreporite, rectangular, almost 3 times as
broad as long, distal margin only slightly convex. Adoral plates rectangu-
lar, meeting fully within, separated distally by a smaller triangular plate.
Jaws prominent, almost twice as long as wide. Three or 4 oral papillae
on each side and a triangular-shaped papilla on the apex of the jaw. Outer
papillae longest, broader than long; median peg shaped, blunt, forming a
straight line with the margins of the outer and apical papillae. Five
bluntly pointed teeth.
Interbrachial spaces below covered with elongated, irregular disk
plates, 2 or 3 elongated plates bordering the base of the arms. Two small,
oval genital slits in each interradius in contact with the oral shield. First
under arm plate trapezoidal, about as wide as long, with a broad, slightly
convex distal margin. Succeeding plates rectangular with concave proxi-
mal and side margins and a slightly broader distal margin. The distal
portion of the plate overlaps the proximal margin of the succeeding plate.
One flat, rounded tentacle scale, about 1% the length of the under arm
plate, attached to the side arm plate.
Color.—Dried specimen, disk white centrally with a brownish tinge
on the margins. An irregular oval brown ring at the base of each arm
and one radiating out from the center of the disk to the interbrachial
area. Each of the larger disk plates bears a brown marking. The upper
arms bear a brown median stripe the width of the upper arm plate,
including the portions of the side arm plates that come within that width.
Bordering the brown median band is a white band, lateral to which is a
narrow brown line that disappears distally. The remainder of the side
arm plates and the median and lower arm spines are pink. The broad
upper arm spine is half white and half brown. The arms are uniformly
white at the tips, and the under arm plates are white; tentacle scales
white with a tinge of pink now and then. In life the disk was pale, its
sides really coral red and the lower surface quite red; no distinct marks
on the disk; a deep yellow stripe (longitudinal) on upper surface of the
arm.
No. 2 ZIESENHENNE: NEW OPHIURANS 11
Types.—Holotype, AHF no. 9; 7 paratypes, one in M.C.Z.
Type locality Station 863-38, clinging to red coral, 30-50 fms., off
Bahia Honda, Panama, March 1, 1938. Five additional paratypes from
Station 948-39, 30-35 fms., off Bahia Honda, Panama, March 28, 1939.
Remarks.—This species is closely related to S. lineata L. & M. of
Cocos Island and the Galapagos Islands but differs in the flat disk scales,
less swollen arm plates, and the very different color pattern. From S.
murrhina Lyman of the West Indies, it differs markedly in the same
characters and in its much smaller size and more slender form. Its dis-
tinctive characteristics besides the color pattern are: irregular disk scales
with no rosette, knobs, or noticeable convexities ; uppermost arm spine, at
least on basal segments, flat, longest at the upper margin, not quite so
large as the other two spines together, and not twice so wide as long. ‘The
name Jaevis has reference to the smooth disk scales.
Ophiacanthidae
Genus OPHIACANTHA Miller and Troschel
Ophiacantha phragma, new species
Plate 2, Figs. 4-6
Description —Disk diameter, 8 mm.; arms, about 38 mm. long. Disk
pentagonal, rather thick, covered with closely compacted, short, rounded
granules. Irregularly placed are some long, stout, pointed disk spines, y,
mm. in length. Radial shields concealed. Disk granulations extend out on
the first few upper arm plates.
First upper arm plate small, partially concealed by granules. Suc-
ceeding plates longer than broad, somewhat hexagonal, proximal margins
straight, distal lateral and proximal lateral margins nearly straight, distal
margins slightly concave; distally the plates become triangular, longer
than broad, with a convex distal margin. Upper plates in contact basally,
separated by the side arm plates distally. Side arm plates broad, in contact
distally. Arm-spine ridge prominent, bearing 7 long, slender, tapering
arm spines, the uppermost longest, more than 2 arm segments in length,
gradually reducing in length to the lowermost, which is shortest.
Oral shields triangular, acute inner angle and rounded outer angles
with conspicuous granules on the distal margin, like those of the disk.
Adoral plates broad and narrow, slightly enlarged distally, meeting fully
within. Jaws long and narrow, bearing medially a pair of granules about
the size of those on the disk. Five to 7 oral papillae on a side. Outermost
12 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
largest, broad, slightly longer than broad with a rounded cutting edge.
Next papillae broad, flat, twice as long as broad. Inner 3 to 5 narrow,
acutely pointed, apical pair slightly largest.
Interbrachial spaces below covered with a dense granulation and
bearing spines centrally but not along the large genital slits. First under
arm plate diamond shaped, twice as broad as long. Succeeding plates tri-
angular, about as long as broad, with a broadly rounded distal margin.
Basally the plates are in contact; distally they are separated by the side
arm plates. Two and sometimes 3 erect, flat, oval tentacle scales on the
first arm segment. Two broad, oval, flat tentacle scales on the next 5 to 8
arm segments, inner scale slightly the longest. Only one flat, oval,
appressed tentacle scale on the remaining arm segments.
Color.—Dried specimen, disk granulations brown, disk spines pale
cream color. Arms light purple with a dark band every 2 to 5 segments.
Under arms and mouth parts are creamy white. A smaller dried specimen
had brown and white markings on the disk with white disk spines. Arms
show longitudinal purplish lines in addition to the bands.
Types.——Holotype, AHF no. 15; 34 paratypes.
Type locality—Station 529-36, 165 fms., off San Francisquito Bay,
Lower California, Mexico, March 1, 1936. There are 34 paratypes as
follows: 2 from Station 431-35, 45 fms., north of Octavia Bay, Colom-
bia, January 27, 1935; one from Station 534-36, 125 fms., off San Fran-
cisquito Bay, Lower California, Mexico, March 2, 1936; one from Sta-
tion 780-38, 47 fms., Chatham Bay, Cocos Island, Costa Rica, January
14, 1938; 2, one in the Museum of Comparative Zodlogy, from Station
795-38, 55 fms., Sulivan Bay, James Island, Galapagos, January 21,
1938; 5 from Station 948-39, 30-35 fms., outside Medidor Island, Bahia
Honda, Panama, March 28, 1939; 10 from Station 874-38, 45 fms., N.E.
of Anacapa Island, California, August 1, 1938; one from Station 909-39,
60-90 fms., 2 miles off Emerald Bay, Catalina Island, California, Janu-
ary 29, 1939; 3 from Station 750-37, 60 fms., Outer Gorda Banks,
Lower California, Mexico, April 4, 1937; 4 from Station 875-38, 50
fms., N.E. of Anacapa Island, California, August 1, 1938; 2 from Sta-
tion 900-38, 40 fms., Long’s Point, Catalina Island, California, Novem-
ber 18, 1938; 2 from Station 213-34, 7-10 fms., La Plata Island, Ecua-
dor, February 10, 1934; and one from Station 201-34, 25-35 fms.,
Gardner Bay, Hood Island, Galapagos, January 31, 1934.
Remarks.—These specimens have a habit of rupturing the upper sur-
face of the disk upon being brought to the surface. The paratypes from
No. 2 ZIESENHENNE: NEW OPHIURANS 13
Station 750-37 had intact disks, and on the disk margins were several
compact groups of 3, 4, or 5 large conical, blunt-tipped spines, heavier
and twice as long as the disk spines. The distinctive character is the wide
oral shields with thin marginal granules. This species is closely related to
O. bairdi Lyman but differs in having a triangular oral shield, 2 or 3
tentacle scales on the 8 to 10 basal arm segments, and a distinct series of
enlarged granules on the distal margin of the oral shields. It is because of
this palisade or fence on the oral shields that the name phragma was
chosen.
Genus OPHIOMYCES Lyman
Ophiomyces multispinus, new species
Plate 3, Figs. 1-4
Description.—Disk diameter, 5 mm.; disk height, 4.5 mm.; arms, 23
mm. long. Disk conical, almost as high as wide; interradius produced to
form a pronounced ridge, while the area directly above the arm base is
concave. Disk scales numerous, circular, and overlapping; many of them
are beset with short, delicate, finely tapering, pointed spines. At the apex
of the disk are 5 heavy, long, gradually tapering spines, about 1 mm. in
length and longitudinally ribbed. No radial shields can be seen, but the
spine-bearing scales are fewer in the region of the arm base.
Arms short, heavy, and rapidly tapering. First 3 upper arm plates
overlapping, small, oval, broader than long, with convex distal margins.
Fourth plate triangular, broader than long with the peak within. Fifth
plate triangular, broader than long, proximal margins concave, distal
margin convex. The Ist to 5th upper arm plates separate the side arm
plates, from which point the side arm plates are in contact above. Suc-
ceeding plates broader than long, proximal margin slightly convex, distal
margin deeply convex. Plates reduce in size distally to the arm extremi-
ties, where they are barely visible. Side arm plates broad with a promi-
nent spine ridge, separated from each other by a naked membrane. ‘They
meet above and below except the 3 basal segments above, where the upper
arm plates separate them. Arm spines, 9, covering the upper side of the
arm. Uppermost fine, delicate, and shortest ; gradually they become longer
below, to the 3rd upper spine, which is the longest, more than 114 arm
segments in length. Undermost 4 spines heavy, curved, longitudinally
striated, undermost 2 slightly shorter; spines become fewer and relatively
shorter distally.
14 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
Oral shields and adoral plates completely concealed by the broad and
numerous oral papillae. By removing the papillae the oral shield is found
to be triangular, acutely pointed within, with slightly concave margins,
distal margin truncated and raised to a median, rounded ridge that bears
one short, heavy, rounded, tapering, blunt spine. Adoral plates large,
roughly triangular, larger than the oral shields, barely meeting within; a
proximal swollen ridge bears the 2 exceedingly wide, fan-shaped, distal
oral papillae. Jaw plates long and rounded, meeting only proximally and
bearing 2 rows of oral papillae, outer and inner rows; outermost papilla
on the jaw plate is the 3rd and last fanlike mouth papilla. Twelve oval
oral papillae on each side, arranged in 2 distinct rows, one on the margin
of the jaw and the other within the margin of the jaw plate. The mar-
ginal row of papillae from the outer to the apical is as follows: outermost
very broad, flat, fan shaped, longer than broad, largest of the row; next
3 fan shaped, flat, longer than wide, diminishing in size to the apex of the
jaw; next 3 short, rounded, tapering to a blunt point. Apical papillae
short, rounded, and bluntly pointed. Inner row of oral papillae 6; the
distalmost exceedingly broad and fan shaped, only slightly longer than
wide; next 3 flat, spatulate, 3 times as long as broad. Proximal 2 narrow,
rounded at the tips, short, only slightly longer than the corresponding
outer row of papillae. Four flat, broad, rounded teeth, widely spaced.
No tooth papillae.
Interbrachial spaces below densely covered with scales, some bearing
spines. ‘I'wo long, narrow, genital slits bordered by long, wide, genital
plates, parallel to each other, obliquely set at arm base. Under arm plates
hourglass shaped, longer than wide, lateral margins concaved by the large
tentacle pores; produced median ridge bearing tentacle scales. Tentacle
scales, 3 pairs on Ist 3 arm segments. One long, narrow tentacle scale on
the side arm plate and 2 median flat, oval, fan-shaped tentacle scales
attached to the under arm plate. Next 4 arm segments with 4 pairs of
tentacle scales, attached to the under arm plate, outer 2 oval and fan
shaped, inner 2 long, paddle shaped, narrow, and pointed. Three pairs of
tentacle scales on succeeding 7 arm segments, inner 2 pairs long and nar-
row, outer one oval and acutely pointed ; succeeding 12 segments bear 2
tentacle scales, an oval outer and a long thin inner scale; near the ex-
tremities only one narrow, pointed, tentacle scale.
Color.—In life, disk grayish blue with a dull yellow stripe in each
interradius; arms pale orange, lighter at base and orally. Dried speci-
mens are nearly white, but the disk has an evident bluish tinge.
NO. 2 ZIESENHENNE: NEW OPHIURANS 15
Types—Holotype, AHF no. 5; 5 paratypes, one in M.C.Z.
Type locality —Station 810-38, 73 fms., off Barrington Island, Gala-
pagos, January 26, 1938. Six diskless specimens from Station 810-38 in
the Museum of Comparative Zodlogy and 2 diskless specimens from
Station 786-38, 392 fms., N.E. of Indefatigable Island, Galapagos,
January 18, 1938.
Remarks.—This species is readily distinguished from the other known
members of the genus by the numerous tentacle scales, particularly the
long paddle-shaped ones. The numerous short and fine spines of the disk
and the numerous slender arm spines are also important characters. This
species has therefore been named multispinus with reference to the
numerous spines.
Amphiuridae
Genus OPHIOPHRAGMUS Lyman
Ophiophragmus disacanthus, new species
Plate 9, Figs. 1-3
Description—Disk diameter, 4 mm.; arms, 33 mm. long. Disk pen-
tagonal, swollen, covered by numerous, overlapping disk scales; a large
central plate and 10 slightly smaller radiating plates. There are from 7 to
9 irregular series of scales in each interradius. The median series overlaps
the adjoining series, and they in turn overlap the adjacent series. ‘The side
and upper disk scales meet to form a distinct disk margin. In each inter-
radius are 2 long, stout, flattened, marginal papillae. The length of the
papillae is 3 to 4 times its diameter. In comparison, the papillae exceed the
largest arm spine in length, but they are not so broad or so heavy. Radial
shields are about twice as long as wide, length less than half the disk
radius, in contact without, separated within by one disk scale. Interradial
edges covered by overlapping disk scales.
Upper arm plates quadrangular, wider than long, side margins
broadly convex, proximal and distal margins slightly convex. Arm plates
fully in contact with each other. Side arm plates very narrow, separated
from each other by a membrane, not in contact above and barely in con-
tact below. Spine ridge large and prominent, bearing 3 short, heavy,
blunt-tipped spatulate arm spines. The spines are less than an arm seg-
ment in length; median spine narrow and short; upper and lower spines
heavy and broad, length 2 to 3 times the diameter of the spine.
16 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Oral shields elongated, almost twice as long as broad. An acute angle
within and a lobe without, outer side margins slightly concave. Adoral
plates triangular, broader than long, widest without. All margins slightly
concave; not quite meeting within. Jaws very small, triangular, broader
than long. Three oral papillae on a side, outermost triangular, about as
long as wide; median bluntly rounded, slightly longer than wide; inner
rectangular, blocklike, longer than wide, widely separated. Four teeth,
ventralmost short and triangular, next 3 broad and rectangular, increas-
ing in length to the uppermost, which is the longest.
Interbrachial spaces below covered with large overlapping disk scales,
as large as those on the upper surface. Genital slits inconspicuous, con-
cealed by the arm spines. First under arm plate small, rectangular, wider
than long, margins straight. Succeeding plates pentagonal, about as wide
as long, angle within, proximal lateral and lateral margins almost
straight, distal margins concave. Plates barely in contact or separated by
the side arm plates. Two small oval tentacle scales placed at right angles
to each other, innermost slightly the longer.
Color.—Dried from alcohol, the holotype was a dark olive buff on
both upper and under surfaces without any special markings.
Types——Holotype, AHF no. 21; 4 paratypes.
Type locality—Station 177-34, 5-20 fms., Sulivan Bay, James
Island, Galapagos, January 23, 1934. Four paratypes from Station 337-
35, 2-5 fms., Sulivan Bay, James Island, Galapagos, December 12, 1934.
Remarks.—This species seems to be confined to the Galapagos
Islands. It can readily be distinguished from all known species of the
genus by the 10 large marginal papillae, longer than the arm spines. The
papillae are well spaced, 2 in each interradius. In the 5 specimens exam-
ined, not more than 2 papillae were found in any interradius. In dredging,
several papillae were crushed and broken, but prominent scars remained
to mark their location. The name disacanthus has reference to the 2 mar-
ginal disk papillae in each interradius.
Ophiophragmus lonchophorus, new species
Plate 4, Figs. 4-6
Description —Disk diameter, 3 mm.; arms, 23 mm. long. Disk circu-
lar and swollen, covered with large, polygonal, overlapping disk scales.
There are several conspicuous disk scales but no central disk plate or
rosette arrangement. In the interbrachial areas are 5 rows of disk scales;
NO. 2 ZIESENHENNE: NEW OPHIURANS 17
the median row overlaps the adjoining rows. There are from 11 to 18
stout, cylindrical marginal papillae in each interradius. Radial shields
longer than wide, length equal to half the disk radius, in contact except
inner ends, where a small scale separates them.
Upper arm plates transversely oval, proximal margin broadly convex,
distal margin straight overlapping the margin of the succeeding plate.
Side arm plates not meeting above or below. Spine ridge bears 3 some-
what flattened, bluntly pointed arm spines, about an arm segment in
length; middle arm spine slightly the longest.
Oral shields diamond shaped, about as long as wide, with rounded
angles. Adoral plates swollen, broader than long, meeting within. Jaws
small and triangular, a deep depression between them. Three oral papillae
on aside, outermost largest, somewhat triangular, about as broad as long;
median small, flat, peglike, longer than broad; apical blocklike, heavy,
slightly longer than broad. Four broad teeth, uppermost longest.
Interbrachial spaces below covered with minute overlapping scales.
The scales are so fine and crowded that they can only be taken for gran-
ules under low magnification; only under high magnification can the
scales be distinctly seen. Genital slits long, extending from the oral shield
to the disk margin; the edge bears a row of overlapping scales, larger
than the under disk scales. First under arm plates small, triangular, angle
within, distal margin straight. Second and succeeding plates pentagonal,
about as broad as long basally, and becoming longer than broad distally.
Lateral, proximal lateral, and distal margins slightly concave. ‘The basal
7 or 8 plates have a notable raised median ridge. Tentacle scales 2, a
small, narrow, oval outer scale attached to the side arm plate and a rela-
tively huge spearhead-shaped inner scale, almost twice as long as broad,
attached to the under arm plate. The smaller scale is not present on the
distal third of the arm. The large scale is not present on the last 15 arm
segments, nor is it conspicuous on newly regenerated segments.
Color.—Dried from alcohol, disk and arms light gray, with irregular
brownish drab arm bands on distal third of arms. Marginal papillae and
arm spines pale gray. Under disk surface light gray, mouth parts and
under arm lighter. The paratype, dried from alcohol, is dark brownish
drab; under disk surface, marginal papillae, and arm spines gray. The
median ridge of the basal under arm plates is also pale gray.
Types—Holotype, AHF no. 19; one paratype.
Type locality—Holotype and paratype from Station 964-39, 2-8
fms., Tenacatita Bay, Mexico, May 8, 1939.
18 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Remarks.—This species can be readily distinguished from other
known members of the genus by the unique inner tentacle scale. It is
strange that after dredging in Tenacatita Bay on seven different occa-
sions this notable new species should now be taken for the first time. The
name lonchophorus has reference to the unique spear-shaped inner ten-
tacle scale.
Ophiophragmus ophiactoides, new species
Plate 3, Figs. 5-7
Description.—Disk diameter, 7 mm.; arms, about 38 mm. long. Disk
covered with small scales; only a few specimens had a large central plate;
scales enlarged about the interradial margins and radial shields. Inter-
radial fence is composed of small, blunt, rounded marginal papillae, vary-
ing from 5 to 23 in number in each interradius; usually there are 15 to 20.
Papillae near the radial shields are the largest, those midway of the arms
smallest and finest. Radial shields oval, longer than wide; % the diameter
of the disk, separated proximally by a single (sometimes a double) series
of long, narrow scales, in contact distally or in some specimens separated
even there by a single narrow scale.
First upper plate triangular, broader than long, angle within, distal
margin convex. Succeeding plates oval shaped, twice as broad as long,
with convex lateral margins, proximal margins convex, distal margins
almost straight, wider than proximal margin. On the distal margin is a
median notch that varies from a slight indentation to a large notch, which
gives the appearance of a concave margin. Side arm plates small, scarcely
meeting below. Three arm spines, longer than an arm segment, oval and
blunt, flattened at the tips; lower spine rather long; median slightly the
longest ; upper spine shortest and flattest.
Oral shields triangular, longer than broad. Adoral plates twice as
broad as long, widening distally and extending to the genital slits, scarcely
meeting within. Three oral papillae on a side, outermost largest, broad
and oval; median narrow and rounded; apical heavy and blunt. Teeth
prominent, flat and rounded, in some specimens somewhat bilobed.
Interbrachial spaces below covered with fine scales. Genital slits equal
to 6 arm segments in length. First under arm plate small, as broad as long,
others hexagonal, broader than long, proximal margin straight, proximal
lateral, lateral, and distal margins slightly concave. Tentacle scales 2, at
right angles to each other; the one on the side arm plate the largest.
NO. 2 ZIESENHENNE: NEW OPHIURANS 19
Color.—Dried specimen, disk dull yellow, with a grayish hue, only
the distal ends of the radial shields whitish. Arms and arm spines light
brown with irregular darker markings on the dorsal arm plates and
spines. Under side grayish brown with some markings of darker brown
on the under arm plates. Color in life green and brown, much like
Ophiactis savignyi (M. & T.) or young Ophiocoma alexanderi Lyman
and not at all like an Ophiophragmus.
Types——Holotype, AHF no. 10; 169 paratypes, 51 in M.C.Z.
Type locality.—Station 403-35, shore, Manta, Ecuador, January 20,
1935. An additional 159 paratypes as follows: 50 from Station 474-35,
shore, Salinas Bay, Costa Rica, February 10, 1935; 2 from Station 207-
34, shore, south of Point St. Elena, Ecuador, February 8, 1934; one from
Station 415-35, 45 fms., south of Port Utria, Colombia, January 24,
1935; 5 from Station 229-34, shore, Cabita Bay, Colombia, February 13,
1934; 97, 51 in the Museum of Comparative Zodlogy, from Station 848-
38, shore, Cape San Francisco, Ecuador, February 23, 1938, and 4 from
Station 400-35, shore, Manta, Ecuador, January 19, 1935.
Remarks.—The fine scaling of the disk without primary plates, the
slender marginal papillae, the narrow and long radial shields, the rela-
tively short arms with rather slender arm spines, and the triangular oral
shields make a combination of characters that readily distinguish this
species from other known members of the genus. The coarse disk scaling
of O. marginatus Liitken and O. chilensis Lyman separates those forms at
once, while the oral shields, arm plates, arm spines, and the marginal
papillae set it apart from O. paucispinus Nielsen and O. tabogensis
Nielsen. This species lives in algae, in sponges or under rocks, in tide
pools, more or less gregariously. Adults are likely to be more or less
buried in the sand, often several together. Owing to the marked super-
ficial resemblance to Ophiactis savignyit (M.& T.) in color, form, and
habits, the specific name ophiactoides has been chosen.
Ophiophragmus papillatus, new species
Plate 4, Figs. 1-3
Description —Disk diameter, 3.6 mm.; arms, 18 mm. long. Disk
thick and rounded, arms short and heavy. Disk covered with circular
overlapping scales; those in contact with the radial shields slightly larger.
The large, distinct, circular central plate is surrounded by 5 oblong radial
plates almost twice as broad as long. Primary plates separated by the
20 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
scales, 10 of which bear stout, cylindrical, bluntly rounded papillae twice
as long as wide, 2 in each interradius. The marginal fence consists of 8 to
10 broad, flat papillae almost as wide as long, heavy and blunt. Radial
shields small, almost twice as long as broad, less than half the disk radius
in length. Their pointed ends proximally are separated by 2 or 3 disk
scales; distally they are in contact.
Upper arm plates broadly oval with rounded lateral margins, fully
in contact with the distal margins overlapping the proximal margins of
the succeeding plates. Side arm plates narrow, separated from each other
by a naked membrane; arm-spine ridge prominent, bearing 3 short, heavy,
flattened, bluntly pointed arm spines, about twice as long as broad, equal
to the length of an arm segment.
Oral shields diamond shaped, about as long as wide; distal margins
slightly concave. Adoral plates broader than long, margins almost paral-
lel, meeting fully within. Jaw plates long and narrow, meeting proxi-
mally to form a V. Three oval papillae on each side, distal 2 broad, block-
like, and rounded; distalmost only slightly larger; apical papillae block-
like, longer than broad, widely separated. No tooth papillae; teeth wide
with rounded cutting edges.
Interbrachial spaces below covered with overlapping scales, more or
less uniform in appearance. Two long genital slits in each interradius.
First under arm plates rectangular, wider than long, with a notched prox-
imal margin. Second plate squarish with a concave distal margin; suc-
ceeding plates about as wide as long, with convex lateral margins; distal
margins slightly wider than proximal, fully in contact with each other,
the outer one slightly larger.
Color.—Dried specimen, disk almost white, a purplish tinge on the
primary plates. Upper arm plates irregularly marked, some plates white,
others almost black with variations of brown plates with dark markings,
white plates with brown and black markings and white plates with par-
allel lines; there is no definite banding arrangement. Arm spines tinged
with purple. Under side entirely white with a few dark bands on the
distal under arm plates.
Type.—Holotype, AHF no. 11.
Type locality—Station 260-34, shore, Tangola Island, Tangola-
Tangola Bay, Mexico, March 1, 1934.
Remarks.—This species is easily distinguished from all known mem-
bers of the genus by the 10 cylindrical papillae on the disk. Because of this
characteristic feature the name fapillatus has been selected.
No. 2 ZIESENHENNE: NEW OPHIURANS 21
Ophiophragmus stellatus, new species
Plate 6, Figs. 6-9
Description Disk diameter, 5.5 mm.; arms, about 65 mm. long.
Disk flat and pentagonal, covered by numerous overlapping scales in an
irregular rosette. Conspicuously placed are the large circular central and
radial plates and one or two large oval plates in each interradius. ‘The
plates are surrounded by numerous irregular disk scales. About 10 rows
of scales in each interradius, median row of scales the largest. Disk mar-
gin bears 15 to 18 long, well-spaced, cylindrical papillae, about 4 times
their diameter in length. Radial shields large, nearly half the disk radius,
longer than broad, acutely pointed within, separated within by 3 or 4
ovate scales.
Upper arm plates oval, twice as wide as long, proximal and distal
margins broadly convex, lateral margins acutely convex, in contact with
each other. Side arm plates narrow, separated by a naked membrane, not
in contact above or below. Spine ridge prominent, bearing 3 heavy, flat-
tened arm spines, about an arm segment in length, uppermost shortest,
undermost heaviest and longest.
Oral shields diamond shaped, slightly longer than wide, outer margins
concave. Madreporite swollen and considerably larger than other oral
shields. Adoral plates triangular, outer margin widest, not meeting
within. Jaws small and triangular. Three oral papillae on a side, outer-
most triangular, broad and flat; median spiniform, more than twice as
long as broad, longer than other papillae, resembling the fence papillae
more than an oral papilla; inner papillae blocklike, longer than broad,
separated from each other. Four broad, blunt teeth, uppermost longest.
Interbrachial spaces below covered with small, uniform, overlapping
scales. Genital slits large and wide. Genital ridge covered with a series
of large, overlapping scales.
First under arm plate small, proximal margin concave, other margins
straight. Second and succeeding under arm plates pentagonal, wider than
long, proximal lateral and distal margins concave, lateral margins
straight, plates in contact with each other. I'wo small, oval tentacle
scales placed at right angles to each other, inner scales slightly the longer.
Color—Dried from alcohol, upper surface pale gray, disk plates
slightly darker. Under surface light cream with some pale gray markings
on under arm plates.
Types.—Holotype, AHF no. 17; 138 paratypes, 71 in M.C.Z.
22 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Type locality—Station 823-38, 30-40 fms., San Juan Bay, Peru,
February 7, 1938. Two paratypes from Station 833-38, 10 fms., Inde-
pendencia Bay, Peru, February 10, 1938.
Remarks.—In the smaller specimens of 2 mm. diameter the rosette
was regular and there were 4 rows of scales in each interradius. Only one
small scale separated the radial shields proximally. The marginal fence
contained only 7 papillae, and the upper arm plates were somewhat tri-
angular. The median oral papillae are spiniform and project above the
other papillae as in the type. The madreporite is swollen and considerably
larger than the oral shields in all specimens. This species can be distin-
guished from Ophiophragmus chilensis Lyman and other members of the
genus by the large radial shields, rosette disk, median oral spiniform
papilla, and the unique enlarged and swollen madreporite. The name
stellatus has reference to the star formed by the large primary plates on
the disk.
Genus AMPHICHONDRIUS Nielsen
Amphichondrius laevis, new species
Plate 4, Figs. 7-9
Description.—Disk diameter, 3.5 mm.; arms, 16 mm. long. Disk pen-
tagonal and swollen, covered with large overlapping disk plates that form
a rosette. Surrounding the large central disk plate are 5 small triangular
plates, which with equally small distal triangular plates separate the 3
large radial plates from one another and from the central plate. There are
a few disk scales surrounding the large plates, while the remainder of the
disk is covered with moderate-sized, overlapping disk scales. Five or 6
rows of scales in each interradius. On the disk margin, midway between
the arms, is a large, conspicuous, elliptical plate, almost twice as wide as
long. The disk granulations of the interbrachial spaces below can be seen
distal to the marginal plates. Radial shields small, longer than broad,
length less than half the disk radius; separated within by one or two
disk scales, outer ends in contact.
Upper arm plates triangular with broadly rounded angles, proximal
margin strongly convex. Basally the arm plates are broader than long;
distally they become longer than broad. The plates are separated from
one another by the side arm plates, which are finely granulated and meet
above and below. Spine ridge large and heavy, bearing 3 stout, erect,
subequal arm spines less than an arm segment in length, upper spine
stoutest and slightly longest.
NO. 2 ZIESENHENNE: NEW OPHIURANS 23
Oral shields small, diamond shaped, about as long as wide, with an
angle within and without. Adoral plates broader than long, meeting with-
in, separated at the outer ends by the first under arm plate. Jaws small,
triangular, depressed medianly. Three oral papillae on a side, outermost
largest, rectangular, broader than long, slightly elevated above the jaw;
median as long as wide, elevated above the jaw; apical slightly longer
than wide, separated from each other and set deeper in the mouth than
other papillae. Four teeth, uppermost longest, lowest shortest. All plates
of the oral frame and mouth more or less swollen and rounded on the
angles.
Interbrachial spaces below covered with a dense coat of minute
granules; those along the genital slits and disk margin slightly larger.
Genital slits long, extending from the oral shields to the disk margin.
First under arm plate small, pentagonal, in contact laterally with the
adoral plates. Second and succeeding plates pentagonal, slightly wider
than long, angle within, proximal lateral margins concave, lateral mar-
gins almost straight, distal margin straight, but becoming notched distal-
ly, basally all angles rounded. Two rather large tentacle scales, one on
the side arm plate, oval, and one on the under arm plate longer than wide;
the two are in close contact.
Color.—Dried from alcohol, disk light gray with a brownish drab
mark centrally on the radial shields and primary plates. Arms light gray,
arm spines white. Interbrachial spaces below light tan, mouth parts and
under arms dull white.
Types—Holotype, AHF no. 18; 35 paratypes.
Type locality —Station 964-39, 2-8 fms., Tenacatita Bay, Mexico,
May 8, 1939. An additional 13 paratypes as follows: one from Station
209-34, 8-10 fms., La Libertad, Ecuador, February 9, 1934; 2 from
Station 259-34, 12-20 fms., Tangola-Tangola Bay, Mexico, February 28,
1934; 4 from Station 281-34, 35-40 fms., Santa Maria Bay, Lower Cali-
fornia, Mexico, March 7, 1934; one from Station 492-36, 35 fms., San
Lorenzo Channel, Espiritu Santo Island, Lower California, Mexico,
February 19, 1936; one from Station 541-36, 60 fms., Puerto Refugio,
Angel de la Guardia Island, Gulf of California; 3 from Station 767-38,
40-50 fms., off Mexico, Lat. 15° 55’ N., Long. 97° 41’ W., January 9,
1938; and one from Station 770-38, 7 fms., off San Jose Light, Guate-
mala, January 11, 1938.
Remarks.—This species differs from Amphichondrius granulosus
Nielsen in having a disk rosette, upper and under arm plates separated by
24 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the side arm plates, radial shields in contact distally, oral shields diamond
shaped, granulations on the genital ridge instead of scales with parallel
furrows, and triangular-oval upper arm plates little wider than long. ‘The
name Jaevis has reference to the very smooth disk.
Genus AMPHIODIA Verrill
Amphiodia sculptilis, new species
Plate 5, Figs. 1-3
Description ——Disk diameter, 5.5 mm.; arms, 26 mm. long. Disk
round and thick, covered with numerous plates. A small pentagonal cen-
tral plate and 5 narrow radiating plates form a star on the disk. Between
the narrow plates are 5 larger radial plates about the size of the central
disk plate. In each interbrachial area are 7 series of long plates, the middle
series broadest and overlapping the other series. Numerous small, overlap-
ping scales radiate out from the center of the disk. Radial shields rounded
triangular, longer than broad, outer and distal margins convex. A small
triangular scale separates the shields within, and in some cases smaller
elongated plates may separate them. The radial shields in all specimens
are separated within by at least one triangular scale; without they are in
contact. Sides of disk covered with overlapping scales. Lateral to the geni-
tal plate and distal to the radial shield is a series of 3 or 4 minute, blunt,
cylindrical spines.
Upper arm plates rectangular, almost twice as broad as long, fully in
contact with each other, margins straight, distal angles slightly rounded.
Side arm plates small, arm spine ridge prominent, bearing 3 short, broad,
bluntly rounded arm spines, uppermost slightly largest.
Oral shields very small and diamond shaped, an obtuse angle within
and truncated without. Adoral plates triangular, broader than long,
widest distally, proximal margins concave, the combination of all the
adoral plates and oral shields gives the under side a star-shaped appear-
ance; within each raylike depression the mouth papillae are located. Jaws
small and narrow, bearing 3 oral papillae on a side, outermost largest,
flat and triangular; median oval, slightly smaller than others; proximal
oblong, twice as long as wide, blocklike, and separated from each other.
Five broad teeth, uppermost longest.
Interbrachial spaces below covered with uniform overlapping scales
that are continuous with the interbrachial scales. Genital slits narrow,
bordered by a series of uniform overlapping scales. First under arm plate
small, triangular, broader than long. Second and succeeding plates larger,
NO. 2 ZIESENHENNE: NEW OPHIURANS 25
rectangular, proximal margin convex, distal margin distinctly concave.
On the basal under arm plates are 2 longitudinal grooves that divide the
plate into a small, oblong center ridge and 2 lateral broader ridges that
appear somewhat like supplementary plates, but the plate is not divided.
Beyond the disk margin the grooves disappear and the plates are smooth,
with parallel lateral margins and convex proximal and concave distal
margins. They are fully in contact. One large tentacle scale, longer than
wide, widest proximally, almost as long as under arm plate, attached to
the lateral margins of the under arm plate.
Color.—Dried specimens were pale variegated gray above and dull
white below; arms purple gray with irregular white bands.
Types.—Holotype, AHF no. 20; 20 paratypes.
Type locality——Station 764-38, 15-20 fms., off White Friar Rocks,
Guerrero, Mexico, January 8, 1938. Twenty paratypes as follows: one
from Station 121-33, shore, Tenacatita Bay, Mexico, March 16, 1933;
2 from Station 212-34, 45-55 fms., La Plata Island, Ecuador, February
10, 1934; 3 from Station 259-34, 18 fms., Tangola-Tangola Bay, Mexi-
co, February 28, 1934; one from Station 265-34, 5-10 fms., Petatlan
Bay, Mexico, March 3, 1934; 2, one in Museum of Comparative Zo-
dlogy, from Station 765-38, 5-10 fms., Chacahua Bay, Mexico, January
9, 1938; 3 from Station 927-39, 4-12 fms., Chacahua Bay, Mexico,
March 22, 1939; and 8 from Station 964-39, 2-8 fms., Tenacatita Bay,
Mexico, May 8, 1939.
Remarks.—This species is closely related to O. violacea Liitken, from
which it differs by the single tentacle scale, the grooved under arm plates,
and the more complex rosette arrangement on the disk. From O. tabogae
Nielsen, it differs by the shape of the radial shields, semirosette arrange-
ment of the disk scales, and the unique under arm plates. The curious
sculpturing of the basal under arm plates and the very small oral shields
will distinguish this species from any other known member of the genus.
The name sculftilis has reference to the characteristic basal under arm
plates.
Genus OPHIOCNIDA Lyman
Ophiocnida californica, new species
Plate 5, Figs. 4-6
Description—Disk diameter, 6 mm.; arms, 34 mm. long. Disk pen-
tagonal, thick, covered with irregular scales radiating from a central
primary plate. Scales overlapping, those surrounding the radial shields
26 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
largest. A number of the plates bear a short, heavy, tapering spinelet.
There are from 15 to 20 spinelets irregularly placed on the disk, but they
are even more numerous along the sides and margin of the disk. Radial
shields long and narrow, about 3 times as long as broad, separated by a
series of scales proximally, distal half fully in contact. Surface of radial
shields finely granulate. Genital plates in contact with the distal margin
of the radial shields and visible from above.
Upper arm plates triangular, twice as broad as long, an obtuse angle
proximally, sides slightly convex, distal margin variable according to the
number of pieces in the upper plate. Upper plates variable, sometimes in a
single piece but often 2 distinct lateral sections and from one to 6 median
pieces, usually one small triangular piece distally, 2 small median triangu-
lar pieces or often additional pieces proximally. The upper plates are in
contact basally, but distally they become separated by the side arm plates.
Side arm plates with a prominent spine ridge bearing 3 heavy, tapering,
bluntly tipped arm spines. Middle spine longest, about as long as an arm
segment. Spines on the first arm plate very short and stubby; they gradu-
ally increase in length to the 6th arm segment, from whence they are more
or less uniform distally.
Oral shields diamond shaped, slightly wider than long, with an acute
proximal angle. Adoral plates twice as broad as long, meeting within,
widest distally. Jaws long and narrow. Three oral papillae on a side,
outermost rectangular, longer than wide, with a serrate cutting margin;
apical blocklike, well separated, longer than broad, cutting edges concave;
median heavy, triangular, about as long as wide. Four broad, flat teeth,
lowermost shortest, uppermost longest. No tooth papillae.
Interbrachial spaces below covered with thick, overlapping, crowded
scales, a few bearing short, heavy, sharp spinelets, irregularly distributed.
Genital slits bordered by a series of scales. First under arm plate small,
longer than wide, with an acutely convex distal margin. Second and suc-
ceeding plates pentagonal, wider than long, with an obtuse proximal angle
and straight lateral and distal margins. Plates barely in contact basally,
separated distally by the large, coarsely granulated side arm plates. Iwo
long, narrow tentacle scales attached on the distal side of the tentacle
pore.
Color.—Dried specimens had dark cream-colored disks with darker
radial shields. Arms pinkish buff with irregular maroon bands and mark-
ings. Interbrachial areas below and mouth parts grayish white, under
arms pinkish buff with no banding.
NO. 2 ZIESENHENNE: NEW OPHIURANS 27
Types.—Holotype, AHF no. 13; 15 paratypes.
Type locality—Holotype and 5 paratypes from Station 529-36, 165
fms., off San Francisquito Bay, Lower California, March 1, 1936. Ten
paratypes as follows: 2 from Station 526-36, 3-5 fms., off Mangles
Anchorage, Lower California, February 28, 1936; 5 from Station 534-
36, 125 fms., off San Francisquito Bay, Lower California, March 2,
1936; and 3 from Station 750-37, 60 fms., Outer Gorda Bank, Lower
California, April 4, 1937.
Remarks.—In the smaller specimens the dorsal arm plates are not
divided. O. californica can be distinguished from O. hispida (Le Conte)
by the oral shields, which are about as long as broad; the outermost oral
papilla, which is longer than broad; radial shields half in contact ; upper
arm plates triangular, divided into several pieces in the adults; and the
arm length, which is 5 times the disk diameter, while the arm length of
O. hispida is 10 times the disk diameter. From the other members of the
genus, O. californica can be readily distinguished by the unique upper arm
plates and the oral shields. It has been named californica, as the range of
this species seems to be confined to the Gulf of California.
Genus AMPHIOPLUS Verrill
Amphioplus philohelminthius, new species
Plate 5, Figs. 7-9
Description —Disk diameter, 4 mm.; arms, 51 mm. long. Disk small,
pentagonal, with a central and 5 radial primary disk plates; disk covered
with small uniform scales, those bordering the radial shields larger.
Radial shields large, the length more than half the disk radius, about 3
times as long as broad, slightly separated proximally but in contact dis-
tally ; somewhat swollen and raised above the remainder of the disk.
First upper arm plate wider than long. Succeeding plates oval, slightly
wider than long, becoming more so distally, widest proximally, with a
convex margin, distal margin sharply rounded. At the arm extremities the
plates become longer than wide and rounded pentagonal or triangular.
The plates are separated by the side arm plates throughout the arm
length.
Side arm plates 3 times as broad as long, separated from each other
by a naked membrane. Spine ridge prominent, bearing 4 flattened, lanceo-
late, blunt-tipped spines on the 10 basal arm segments, 3 subequal spines
on the remainder of the arm, median spines shorter than upper and
lowermost.
28 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Oral shields about as long as broad, the inner half much wider than
the outer half, proximal margins broadly rounded, lateral margins con-
cave, distal and truncate. Adoral plates triangular, twice as broad as
long, widest distally, not quite meeting within. Jaws long and narrow.
Four oral papillae on a side, apical largest, rounded and blocklike, longer
than wide, tapering slightly proximally to give a triangular appearance
to some specimens; other papillae flat and rounded, longer than broad,
outermost slightly largest. Four bluntly rounded, broad teeth, lower-
most shortest, uppermost longest.
Interbrachial spaces below covered with small overlapping scales
distally, but fewer proximally. Genital slits very large, bordered by large
and wide genital plates. First under arm plate triangular, longer than
wide, lateral margins deeply concave producing lateral lobes, distal mar-
gin convex. Succeeding plates rectangular, longer than wide, lateral
margins parallel, distal margin concave, proximal margin convex. Dis-
tally the proximal margins are produced in a right angle, and near the
arm extremities the plates become triangular. Basally the plates are in
contact, but distally they become separated by the side arm plates. Tenta-
cle scales, 2 on the basal pores, but the one on the under arm plate is
exceedingly small and soon disappears altogether.
Color.—In dry specimens the disk and arms are light gray with faint
brown markings on the arms. Mouth parts and under arms light yellow.
T ypes.—Holotype, AHF no. 6; 32 paratypes, 16 in M.C.Z.
Type locality—Station 823-38, 30-40 fms., San Juan Bay, Peru,
February 7, 1938. An additional paratype in the Museum of Compara-
tive Zodlogy from Station 826-38, 20-30 fms., San Juan Bay, Peru,
February 7, 1938, and one from Station 833-38, 8 fms., Independencia
Bay, Peru, February 10, 1938.
Remarks.—This species lives in great numbers in a soft mud which
teems with small, white nematode worms, a most repulsive environment,
to which, however, it is well adapted by its small flat disk and long, very
slender arms. It is the first Amphioplus to be reported from the western
coast of South America and differs obviously from the 3 species from
the Panamic region described by Liitken and Mortensen in 1899. The
oral papillae and disk scaling are quite distinctive. The species has been
named philohelminthius because of its fondness for living with little
worms.
NO. 2 ZIESENHENNE: NEW OPHIURANS 29
Ophiochitonidae
Genus OPHIONEREIS Liitken
Ophionereis dictyota, new species
Plate 6, Figs. 1-5
Description.—Disk diameter, 9 mm.; arms broken, longest about 67
mm. long. Disk covered with thick and overlapping scales which become
thicker surrounding the radial shields. Extending interradially from the
radial shields, at the margin of the disk, is a series of 6 to 8 large, distinct
scales, about 4% as long as the length of the radial shields. Radial shields
small and pennantlike, about 3 times as long as broad, proximally finely
pointed, distally rounded, surrounded by large disk scales.
First upper arm plate small, oval, proximal margin concave, distal
convex. Second plate hexagonal, twice as large as the first; third and suc-
ceeding plates larger. The plates are hexagonal, broader than long, fully
in contact. Proximal and proximal lateral margins concave, distal lateral
Margins convex, distal margin nearly straight. Supplementary plates
longer than wide, inner margin concave, outer margins convex to form a
distinct, rounded, outer distal angle. Side arm plates separated from
each other by a soft membrane. Each plate bears 3 rounded, tapering and
slightly curved, blunt-tipped arm spines, slightly longer than an arm
segment, the median longest. Upper spine flat, blunt and little curved.
Oral shields oval, longer than wide, proximally convex, distal lateral
margins concave. Adoral plates lanceolate, wider than long, enlarging
distally with a rounded margin, not meeting within. Jaws long and nar-
row. Four or 5 oral papillae on a side, outermost flat and triangular;
others blunt and angular, almost rounded; apical smallest, shorter than
others. Four broad and rounded teeth, lowermost shortest and thick,
uppermost thin and long.
Interbrachial spaces below densely covered with fine overlapping
scales. Genital slit ridge bears numerous fine stumps, longer than broad
proximally, becoming shorter and fewer distally. First under arm plate
quadrangular, proximal margin narrow and concave, lateral margins
almost straight, and distal margin decidedly rounded. Second plate much
larger, square, proximal margin little concave, plates only % in contact,
distally even less so. One large, oval, flat tentacle scale, almost 34 the
length of under arm plate.
Color.—Dried specimens had a cream-colored disk, reticulated with
brown markings which continue to the oral shields. Proximally between
30 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the genital slits is a dark brown triangular splotch. Arms light buff with
a wide (2 segments), dark brown, almost black band, every 3 to 4 seg-
ments. Mouth parts and arm spines cream colored, slight traces of the
upper arm bands continue below.
T ypes——Holotype, AHF no. 7; 5 paratypes.
Type locality—Station 778-38, tide pool, Chatham Bay, Cocos
Island, Costa Rica, January 14, 1938. Five paratypes from Station 445-
35, shore, Panama City, Panama, February 2, 1935; one from Station
447-35, coral, Secas Islands, Panama, February 4, 1935; one from Sta-
tion 512-36, shore, Ballena Bay, Espiritu Santo Island, Gulf of Califor-
nia, February 23, 1936; one from Station 515-36, shore, San Francisco
Island, Gulf of California, February 24, 1936; and one from Station
522-36, tide pool, Agua Verde Bay, Lower California, Mexico, Febru-
ary 27, 1936.
Remarks.—In life the holotype had a bluish gray disk, reticulated
with light brown, much as in the West Indian O. reticulata (Say). The
very long arms, which were much broken in capturing the specimen,
were broadly banded with dusky purple. This species differs from other
eastern Pacific O phionereis by the irregular coarse disk scaling, a uniform
series of 5 or 6 large marginal disk scales extending interradially from
the radial shields, large radial shields; and the distinct coloring of the
arms and disk. The name dictyota has been chosen because of the reticu-
lations on the disk.
Ophionereis perplexa, new species
Plate 7, Figs. 7-9
Description —Disk diameter, 8 mm.; arms, 42 mm. long. Disk more
or less depressed and irregular because of shrinkage upon drying. The
disk is covered by a membrane, which upon drying shows minute, dense,
overlapping, uniform scales, visible only under high magnification. No
primary plates are visible, and the scales bordering the radial shields and
disk margins are not enlarged or conspicuous. Radial shields small, nar-
row, more than 4 times as long as broad, acutely pointed within, widely
separated, distance apart exceeding the length of radial shields. The
minute overlapping scales obscure the outline of the radial shields.
First upper arm plate small, semilunar, wider than long, proximal
margin straight, distal margin broadly convex, overlapping next plate.
Succeeding plates large, wider than long, basally triangular with rounded
corners, proximal margin slightly concave, distal margin convex, less
NO. 2 ZIESENHENNE: NEW OPHIURANS 31
than half the width of proximal margin; distally the plates become pen-
tagonal or hexagonal with slightly concave proximal and proximal lateral
margins and convex distal margins. Supplementary plates triangular,
distal margins convex and bearing a series of parallel minute ridges con-
spicuous under high magnification. Plates 4% to % the size of upper arm
plates. Side arm plates wide, not meeting above or below, spine ridge
prominent, bearing 3 large, heavy, broad, bluntly tipped arm spines; 1%
to 2 arm segments long, middle spine longest, uppermost the shortest.
Oral shields triangular, wider than long, obtuse angle within, lateral
angles convex. Adoral plates narrow, not meeting within, broader dis-
tally. Jaws small and triangular. Four oral papillae on a side, outermost
bluntly pointed, longer than wide; next 2 broad, rounded, and heavy;
apical heavy, blunt, and rounded, slightly longer than broad. In the
holotype and a few paratypes the 2nd outermost papilla is divided into
2 or 3 fine spiniform papillae with a common base. Four broad and
rounded teeth, uppermost longest.
Interbrachial spaces below covered by a thin membrane, which under
high magnification shows very minute scales. Genital slits large and wide;
a few scales can be seen, but no genital papillae are present. First under
arm plate small, slightly longer than wide, distal margin convex, lateral
and proximal margins almost straight. Second and succeeding plates large,
longer than wide, proximal and distal margins convex, lateral margins
almost straight, plates in contact with each other. One large, oval tentacle
scale, half the length of under arm plate, attached to the side arm plate.
Color.—Dried specimens, disk pale yellow with reticulating purplish
lines ; above the arm base and between the radial shields is a purple-brown
area spotted with cream yellow that blends in with the reticulations of
the disk proximally. Upper arms greenish gray with a purple band one
segment wide, continuing below and spaced about 3 to 5 arm segments
apart. The supplementary plates on each side of the purple band are
creamy white. Arm spines are gray with a white base and a transverse
purple band above the base and on the tip. Mouth parts and under arm
plates are creamy white with the purple arm bands extended from above.
Disk below is a brown orange in the naked membrane with the disk
colors and reticulations extending onto the interbrachial area.
Types—Holotype, AHF no. 4; 88 paratypes.
Type locality—Holotype and 3 paratypes from Station 333-35,
shore, James Island, Galapagos, December 11, 1934. Eighty-five addi-
tional paratypes as follows: one from Station 73-33, shore, Cartago Bay,
32 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Albemarle Island, Galapagos, February 13, 1933; 2 from Station 202-34,
shore, Osborn Island, Galapagos, January 31, 1934; one from Station
313-35, shore, Black Beach Anchorage, Charles Island, Galapagos, De-
cember 6, 1934; 3 from Station 342-35, shore, Bartholomew Island, Ga-
lapagos, December 12, 1934; 12 from Station 350-35, shore, South Sey-
mour Island, Galapagos, December 13, 1934; 7 from Station 351-35,
shore, Charles Island, Galapagos, December 14, 1934; one from Station
515-36, shore, San Francisco Island, Gulf of California, February 24,
1936; 6 from Station 530-36, 10-20 fms., off San Francisquito Bay, Low-
er California, Mexico, March 1, 1936; 2 from Station 719-37, 10-25
fms., Consag Rock, Lower California, Mexico, March 24, 1937; 5 from
Station 782-38, shore, Darwin Bay, Tower Island, Galapagos, January
16, 1938; 29 from Station 784-38, shore, Tower Island, Galapagos, Jan-
uary 17, 1938; 3 from Station 789-38, shore, South Seymour Island,
Galapagos, January 19, 1938; one from Station 795-38, 30-40 fms.,
Sulivan Bay, James Island, Galapagos, January 21, 1938; 2 from Station
796-38, shore, Sulivan Bay, James Island, Galapagos, January 21, 1938;
4 from Station 800-38, shore, Cartago Bay, Albemarle Island, Galapagos,
January 22, 1938; 3 from Station 803-38, shore, Black Beach, Charles
Island, Galapagos, January 23, 1938; one from Station 808-38, shore,
Academy Bay, Indefatigable Island, Galapagos, January 25, 1938; and
2 from Station 400-35, shore, Manta, Ecuador, January 19, 1935.
Remarks.—This species is rather common in the Galapagos Islands
and differs from the other members of the genus by its exceedingly fine
disk scaling, radial shields which are separated more than their length
apart, lack of genital papillae, absence of enlarged scales about the disk
margin and radial shields, and coloring. From O. annulata (Le Conte)
and O. dictyota it differs by lacking large disk scales about the radial
shields and disk margins. Its 3 arm spines distinguish it from O. albo-
maculata Smith and O. eurybrachiplax H. L. Clark, which basally have
4, From its closest ally, O. nuda Liitken and Mortensen, it can be dis-
tinguished by its arm length, 4 oral papillae, presence of fine scales, lack
of genital papillae, and uniquely separated radial shields. In a series of
64 are a number of variations; in some specimens the disk below is cov-
ered by a naked membrane, and in others the upper disk scaling is continu-
ous to the adoral plates. Of the 64 specimens, 45 had 4 oral papillae on
each side; 7 had 3 on one or more jaw sides and 4 on the remaining sides ;
and 4 had 4 oral papillae with either of the 2 middle papillae divided into
2 or 3 spiniform papillae, with a common base. The name ferflexa has
NO. 2 ZIESENHENNE: NEW OPHIURANS 33
been selected because of the difficulty of distinguishing this species from
other Pacific forms.
OPHIODESMUS, new genus
Similar to Ophionereis, but the disk scaling is coarse and compacted
to such a degree that only a small portion of some of the disk scales is
visible. The supplementary arm plates are very small, confined mainly to
the side of the arm and not conspicuous from above. Upper arm plates
oval, slightly wider than long. This genus seems to be a connecting link
between Ophionereis of the Ophiochitonidae and Ophiactis of the Amphi-
uridae. Since it shows characteristics of both families and the under side
is decidedly similar to Ophionereis, this genus is placed in the Ophiochi-
tonidae. The name Ophiodesmus has reference to its connecting-link po-
sition between Amphiuridae and Ophiochitonidae.
Genotype: Ophiodesmus amphilogus, new species.
Ophiodesmus amphilogus, new species
Plate 9, Figs. 4-6
Description.—Disk diameter, 4.5 mm.; arms, short and slender, grad-
ually tapering, about 11 mm. long. Disk circular and flat, covered by
numerous compacted, overlapping, irregular scales. Some of the scales are
compacted and overlap to such an extent that only an elliptical portion or
narrow margins are exposed. Surrounding the larger scales are several
irregularly placed smaller scales. There are 7 to 9 irregular series of
scales in each interradius. Primary plates are not conspicuous. Radial
shields small, oval shaped, about twice as long as wide, widely separated,
about their length apart, by several large and numerous small disk scales.
Length of shields about %4 of the disk radius.
First upper arm plate small, wider than long, distal margin convex.
Second plate oval shaped, larger than the first, slightly wider than long,
distal margin strongly convex. Third and succeeding plates larger, oval
shaped, basally wider than long, distally becoming longer than wide, mar-
gins convex. Basally the plates overlap; distally they become separated by
the side arm plates. Under high magnification the upper arm plates appear
smoother in texture than the side arm plates. Side arm plates wider than
long, spine ridge produced and prominent, bearing 3 heavy, flattened,
broad, short, subequal arm spines, as long as an arm segment basally, and
gradually reducing in length distally to the arm tips where they are
34 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
only % the length of the arm segment. Median spine slightly the longest ;
upper spine heaviest and blunt. Supplementary arm plates small and tri-
angular, about 149 the size of the upper arm plate; inner margin concave,
distal and outer margins convex. The supplementary plate is inserted
under the upper and proximal side arm plates and overlaps the distal side
arm plate.
Oral shields triangular, about as long as broad, inner margins almost
straight. Adoral plates triangular, broader than long, widest distally,
meeting within, inner margin slightly concave, other margins straight or
slightly concave. Jaws small, triangular. Four oral papillae, about equal
in length, on a side. Outermost largest, flat, longer than broad, cutting
margin rounded. Inner 3 papillae narrow, flat, longer than broad, bluntly
pointed; inner pair heaviest. Distal to the outer papillae is the first ten-
tacle scale, set deep in the mouth, which might easily be taken for an oral
papilla. Four broad teeth of equal length, cutting margins convex.
Interbrachial spaces below covered with unequal, overlapping, crowd-
ed scales. The largest scales are centrally located ; those along the genital
slits are smaller and crowded so only a portion of the scales are visible.
Genital slits short, equal in length to 2 arm segments. First under arm
plate small, longer than broad, distal margin convex. Second and suc-
ceeding plates large, octagonal basally, triangular distally, longer than
wide; proximal lateral margins straight, lateral margins concave, distal
margins broadly convex. Plates in contact basally, separated distally by
the side arm plates. One large, oval tentacle scale, longer than wide, not
quite half the length of the under arm plate, attached to the side arm
plate.
Color.—Dried from alcohol, disk white with dark gray circles at the
arm base. Arms dark gray with irregularly spaced white and light gray
bands. Under side white with irregular gray bands on the arms. Two of
the paratypes differ by having the disk uniformly mottled with white,
light and dark gray. Arms irregularly banded with white, light and dark
gray. Under side white with several gray bands on the arms.
Types—Holotype, AHF no. 22; 3 paratypes.
Type locality—Station 287-34, 10-15 fms., Cerros Island, Lower
California, Mexico, March 10, 1934.
Remarks.—This species differs from Ophionereis annulata (Le
Conte) of equal size in its coarser disk scaling, lack of genital papillae,
arm length twice the disk diameter, and exceedingly small supplementary
plates. From the young of Ophionereis eurybrachiplax H. L. Clark, it
No. 2 ZIESENHENNE: NEW OPHIURANS 35
differs in having only 3 basal arm spines, shorter and stouter arm spines,
arms only twice the disk diameter, as compared to 5 times the disk di-
ameter, small, oval upper arm plates as compared to large, broad, rec-
tangular upper arm plates. The above characteristics distinguish this
species from other known related forms. In color it differs from Ophio-
nereis, which has regularly dark banded arms. The name amphilogus has
reference to the doubtful position of this species.
Ophiolepididae
Genus OPHIOMISIDIUM Koehler
Ophiomisidium leurum, new species
Plate 7, Figs. 1-3
Description —Disk diameter, 4 mm.; arms, 4 mm. long. Disk large;
arms short, tapering rapidly. Disk covered with 37 large, granulated
plates. Central plate pentagonal, surrounded by 5 quadrangular plates,
whose acute distal angles separate the proximal portion of the radial
shields. Separating the radial shields interradially are 2 plates, a rectan-
gular proximal plate and, distally, a produced, rounded, swollen plate
that fills the interbrachial margin and side of the disk. A rounded plate
and an oblong plate separate the distal margins of the radial shields. A
small plate (distal end of 2nd side arm plate) at the distal margin of
each radial shield, bearing 3 large spines, completes the side margins of
the disk.
Radial shields oblong, almost twice as long as wide, separated proxi-
mally by the primary radial plate, broadly in contact medially but sep-
arated distally by 2 plates, one rounded and swollen above the surface of
the radial shields, the other much broader, not elevated, lying on the side
arm plates at the arm base.
First upper arm plate very small, diamond shaped, succeeding plates
becoming still smaller distally until none can be found after the 4th arm
segment; plates widely separated by the side arm plates. First side arm
plates somewhat swollen, wider than long, meeting within, not visible
from above; 2nd plates are visible from above just distal to the radial
shields. Succeeding side arm plates narrower, becoming longer distally
and fully in contact. There are apparently not more than 10 or 12 arm
segments. Arm spines small, only 2 on first arm segment; there are 3 on
the next arm segment, well separated, uppermost heaviest, triangular,
blunt, longer than wide, median one or 2 shorter, lowermost shortest,
36 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
slightly tapering. Three uniform short arm spines on next 2 arm seg-
ments, blunt and uniformly tapering. Two very short and peglike arm
spines on the remaining arm segments.
Oral shields diamond shaped, longer than broad. Adoral plates dia-
mond shaped, slightly broader than long, meeting fully within. Jaws
quadrilateral, longer than broad. Four oral papillae, broad and blocklike,
broader than long, their outlines cannot be seen distinctly.
Interbrachial spaces below covered by 2 long, narrow plates, distal of
the oral shields, and separated at the disk margin by a triangular plate,
the acute angle within and a broadly rounded margin without that ex-
tends beyond the disk margin, appearing much like a horizontal keel on
the large interbrachial plate. Genital slits not visible. First under arm
plate large, shield shaped, succeeding plates similar but becoming smaller
distally, disappearing entirely on the 6th arm segment. Under plates
widely separated by the side arm plates. Tentacle scales only on the
basal 5 arm segments, 2, sometimes 3, small oval tentacle scales on each
pore.
Color.—All specimens dried from alcohol are bleached nearly white.
Types—Holotype, AHF no. 16; 3 paratypes, one in M.C.Z.
Type locality —Station 810-38, 73 fms., off Barrington Island, Gala-
pagos, January 26, 1938. Paratypes: one from Station 767-38, 45 fms.,
off Chacahua Bay, Oaxaca, Mexico, January 9, 1938; and 2, one in the
Museum of Comparative Zodlogy, from Station 792-38, 75 fms., off
Daphne Minor Island, Galapagos, January 20, 1939.
Remarks.—This little brittle star differs markedly from other known
members of the genus by its smooth, flat disk with 10 marginal knobs (or
at least elevations) on the 5 radial and 5 interradial marginal plates. The
name Jeurum has been selected with reference to the very flattened form
of the entire body.
Genus AMPHIOPHIURA Matsumoto
Amphiophiura irregularis, new species
Plate 7, Figs. 4-6
Description ——Disk diameter, 7 mm.; vertical diameter, 1.3 mm.;
arms, 18 mm. long. Disk pentagonal, flat, composed of 24 flat plates,
including the radial shields; 8 irregular plates surround the central plate,
a large oblong interradial plate separates the radial shields. The number
of disk plates ranges from 20 to 26, including radial shields, in the 16
NO. 2 ZIESENHENNE: NEW OPHIURANS 37
paratypes. A central plate is sometimes found, but a symmetrical rosette
has not been seen in the 17 specimens. In several specimens radial shields
and interradial plates are in contact with the central plate at one point.
Interbrachially is a large, transversely oval plate in contact with the inter-
radial plate and separated from the radial shields by one or 2 smaller
plates. Radial shields large, hexagonal, about half the disk radius, slightly
longer than wide, broadly in contact.
First upper arm plate small, triangular, with the peak within, bearing
2 to 5 fine spinelets on each lateral margin along the genital slit. Second
plate hexagonal, swollen, broader than long, distal and proximal margins
straight, lateral margins little concave, bearing 4 to 8 fine spinelets along
the genital slit. Third plate hexagonal, about as long as broad, swollen,
with straight proximal and distal margins. Succeeding plates pentagonal,
then quadrangular, becoming triangular distally, longer than broad, lat-
eral margins straight, distal margins convex. Basal 8 plates in contact,
distally they become farther separated by the side arm plates until they
are reduced to % the length of an arm segment at the extremities. All
plates are finely granulated. First side arm plate greatly reduced, bearing
2 short, broad tentacle scales and 2 long, conical arm spines on the genital
slit. Third plate slightly larger, bearing 3 short, broad tentacle scales and
4 long, slender, pointed, well-spaced arm spines along the genital slit.
Fourth to 8th plate bearing 2 small, flat, long tentacle scales or modified
arm spines, succeeding plates bearing only one. Fourth and succeeding
plates bear 5 then 4 long, finely tapering, well-spaced arm spines, about
Y the length of the arm segment, median spine longest, upper and lower-
most shortest.
Oral shields large, occupying the interradius below, oval, longer than
wide, with an acute angle within and broad, convex lateral and distal
margins. Adoral plates quadrangular, little broader than long, meeting
within. Jaws wider than long, scarcely larger than adoral plates. Four
or 5 oral papillae on a side, outermost twice as broad as long; next broader
than long; next delicate, longer than broad, apical broad, heavy, pointed,
longer than broad. Three narrow pointed teeth, undermost shortest.
Interbrachial spaces below covered by the large, oval oral shields and
2 or more small interbrachial plates. Genital slits long, extending from
the adoral plates to the radial shields. Genital plates long and narrow, in
contact with the distal margin of the radial shields; each plate bears a
comb of 18 to 26 fine, long, tapering, pointed spines, well separated at
their bases, lowermost becoming shorter and blunt. First under arm plate
38 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
largest, longer than wide, proximal margin convex, distal margin deeply
concave; lateral margins concave, bearing 3 or 4 reduced tentacle scales,
outermost being the largest. Second plate rectangular, broader than long,
proximal margin almost straight, distal margin concave, lateral margins
bearing 2 short, broad tentacle scales. Third plate rectangular, only
slightly broader than long, lateral margins bearing 2 short tentacle scales,
outermost largest. Fourth to 8th plates hexagonal, much wider than long;
Ath, 5th, and 6th bearing a minute tentacle scale on the distal lateral
angle. Succeeding plates pentagonal or oblong, small, becoming triangular
and smaller distally, until they occupy only % of the arm segment or less
at the extremities. The 6 basal plates are separated by a deep concavity,
succeeding plates separated by side arm plates; distally the separation be-
comes even greater.
Color.—Dried from alcohol, disk white with a lavender tinge, arms
light cream color. In life, upper surface variegated with light and dark
red, or red and yellowish, or red and white; oral surface white. ‘The
larger specimens are not deeply colored.
Types—Holotype, AHF no. 12; 60 paratypes, 16 in M.C.Z.
Type locality—Holotype and 33 paratypes, 16 in the Museum of
Comparative Zodlogy, from Station 792-38, 70-80 fms., off Daphne
Minor Island, Galapagos, January 20, 1938; 2 paratypes from Station
191-34, 70 fms., Lat. 0° 55’ S., Long. 90° 39’ W., January 26, 1934;
and 25 paratypes from Station 190-34, 58-60 fms., Lat. 0° 55’ S., Long.
90° 30’ W., January 26, 1934.
Remarks.—This species is closely related to 4. paucisquama but dif-
fers as follows:
A. paucisquama
A symmetrical rosette on the disk
composed of a central pentago-
nal plate and 5 larger, equal ra-
dial plates
A. irregularis
No symmetrical rosette on disk
All plates finely granular
Plates flat or slightly swollen
Radial shields large
Disk flat, 1.5 mm. thick
Arms about 2.2 times disk diame-
ter, not slender distally
Disk and basal arm plates deeply
pitted
Plates convex, basal upper arm
plates almost hemispherical
Radial shields small
Disk elevated, 3.5 mm. thick
Arms short, 1.8 times disk diame-
ter, becoming very slender dis-
tally
NO. 2 ZIESENHENNE: NEW OPHIURANS 39
The name irregularis has been chosen for this species because of the
irregular arrangement of the disk scales.
Amphiophiura paucisquama, new species
Plate 8, Figs. 1-3
Description —Disk diameter, 7 mm.; thickness of disk, 3.5 mm.;
arms, 13 mm. long. Disk large, arms relatively short, tapering rapidly.
Disk covered with 21 very large pitted plates including the radial shields.
A large, swollen, pentagonal central plate is surrounded by 5 slightly
larger plates that are in contact with the single, large, interradial plate
and the radial shields. The interradial plate is pentagonal with the proxi-
mal margins concave, forming a peak, lateral and distal margins almost
straight. Viewed under an ordinary hand lens all plates are deeply pitted,
resembling the mesh on a golf ball. Radial shields much smaller than
disk plates, pentagonal, fully in contact, decidedly broader than long,
deeply pitted as other disk plates.
First upper arm plate oval, broader than long, fully in contact with
the second arm plate and radial shields, lateral margins bear genital slit
papillae. Distally the plate bears an elevated transverse ridge which is
deeply pitted. Second plate oval, broader than long, with a high convex
surface, deeply pitted, in contact with the first upper arm plate. Third
plate oval, slightly broader than long and rather high, elevated to re-
semble a golf ball, deeply reticulated. Succeeding plates diamond shaped,
becoming smaller, flatter, and more widely separated by the side arm
plates distally. Side arm plates twice as broad as long, swollen, in contact
below from the 5th arm segment out and in contact above from the 3rd
segment out. Three short, peglike arm spines about % the length of the
side arm plates, widely separated, upper longest and heaviest, under short-
est. The arm plates are separated from each other or appear that way be-
cause of the prominent spine ridge. All plates are deeply pitted by an
irregular reticulation similar to that on golf balls.
Oral shields large, triangular, truncated within and broadly convex
without, longer than wide, occupying the entire ventral interradius. Ad-
oral plates small, diamond shaped, fully in contact and separated from the
oral shield by a furrow. Jaws diamond shaped, twice as long as wide. Four
or 5 oral papillae on a side, outermost largest, rectangular, broader than
long; 2nd and 3rd outermost rectangular, only slightly broader than long.
Inner papillae ovate, pointed, flat, longer than broad. Three bluntly
rounded teeth.
40 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Interbrachial spaces below covered by the large oral shield surrounded
by a few smaller plates. In the interbrachial area is a large plate in con-
tact with the oral shield and the upper interradial plate. First under arm
plate large, slightly longer than wide, with a convex proximal margin and
concave distal margin, widely separated from the 2nd under arm plate
by a groove. Second plate rectangular, 3rd and succeeding plates longer
than broad, becoming diamond shaped distally, reduced in size and far-
ther separated by the side arm plates, which meet below between the 4th
and 5th under arm plates. The first 5 arm plates are separated below by a
deep groove. Plates swollen and deeply pitted. Genital slits large, extend-
ing from the radial shields to the adoral plates. Thirteen to 18 spines on
the genital ridge, those above long, blunt, and well spaced, undermost
become shorter, broader, flatter, and compacted. Three broad, flat ten-
tacle scales, outermost largest on 1st under plate. Two broad, flat ten-
tacle scales on the adoral plate. Three flat, broad tentacle scales on the
2nd and 3rd under plates. Three broad flat tentacle scales on the 1st and
2nd side arm plates. Two tentacle scales on the next 3 under arm plates
and then only one for 3 plates before disappearing. Two tentacle scales on
the 3rd to 5th side arm plate and then only one to the end of the arm.
Color—Dried from alcohol, white with a lavender tinge, stained
brownish drab about the base of the arms.
Types.—Holotype, AHF no. 8; 13 paratypes, 5 in M.C.Z.
Type locality—Station 818-38, 300 fms., south of Hood Island,
Galapagos, January 20, 1938. Thirteen additional paratypes, 5 in the
Museum of Comparative Zodlogy, from Station 786-38, 392 fms., N.E.
of Indefatigable Island, Galapagos, January 18, 1938.
Remarks.—The thick rounded disk, short rapidly tapering arms, sym-
metrical disk rosette, and unique deep pits on the primary plates, espe-
cially the basal upper arm plates, which resemble the mesh on a golf ball,
easily distinguish the adults of this species from other known members of
the genus. Unfortunately, in small specimens the pitted surface is not
noticeable under an ordinary lens but requires a magnification of 40
diameters or more. It approaches Stegophiura in the shape of the arms,
but the disk scales and arm spines are more like Amphiophiura. This
species has been named paucisquama in reference to the small number of
disk plates.
No. 2 ZIESENHENNE: NEW OPHIURANS 41
Genus OPHIOLEPIS Miiller and Troschel
Ophiolepis plateia, new species
Plate 8, Figs. 4-6
Description—Disk diameter, 13.5 mm.; arms, 38 mm. long. Disk
covered with large primary plates, each surrounded by a row of single
smaller, regularly arranged scales. Disk very flat and depressed; 2
rounded protruding plates on the interbrachial margin, most conspicuous
from the under side. Arms flat, tapering distally; base of arm 4.5 mm.
wide, width midway of the arm only 2.5 mm. Radial shields large and
distinct, longer than broad, separated from each other and interradially
by a primary plate or plates, each surrounded by a single row of small
disk scales.
Upper arm plates rectangular, wider than long basally, becoming tri-
angular distally and longer than wide; lateral margins convex basally,
concave distally, distal margin concave basally, almost convex distally.
Basally 2 triangular supplementary plates are inserted between the upper
ends of the side arm plates; distally they are replaced by a single supple-
mentary plate. Side arm plates large, separated above by the supplemen-
tary plates. There are 4 short, blunt, conical, well-spaced arm spines
basally, but only 3 distally, placed low down on the side arm plates. Arm
spines less than half the length of arm segment.
Oral shields narrow, twice as long as wide, proximal angle acute,
lateral margins concave, distal margin convex. Adoral plates rectangular,
widest without, meeting within, slightly prolonged between the oral
shield and the inner side arm plate. Jaws triangular, longer than wide,
smaller than the adoral plates. Five oral papillae on a side, the triangular
outermost reaching inside the next papilla, which is angular, wider than
long; next squarish, followed by an angular papilla, then a pointed pa-
pilla; apical papilla flat and oval. Four equally long teeth, the lowermost
broad, cutting edge rounded, uppermost narrow, cutting edge straight.
Interbrachial spaces below covered by 2 long genital plates which are
separated by irregular scales, outermost largest, and 2 large rounded
interbrachial plates. Two small genital slits 14 the disk radius in length.
Under arm plates broader than long basally, becoming longer than broad
distally, proximal margins convex, distal and lateral margins concave,
plates fully in contact. Two wedge-shaped tentacle scales, closely in con-
tact, abradial scale slightly larger. There may also be a small accessory
scale on the basal arm segments.
42 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Color.—Dried specimen, disk and arms above dull dark gray and
olive buff irregularly mottled. On the disk scales the color is lighter and
more distinct. Interbrachial disk plates have a central, light cream color
area encircled with a narrow dark brown band. Entire under surface
cream color, general appearance light drab gray.
Types.—Holotype, AHF no. 14; 222 paratypes, 11 in M.C.Z.
Type locality—Holotype and 191 paratypes, 9 in the Museum of
Comparative Zodlogy, from Station 483-35, 6 fms., Tenacatita Bay,
Mexico, February 15, 1935. Thirty-one paratypes as follows: 24, 2 in
Museum of Comparative Zodlogy, from Station 116-33, 2 fms., Cocos
Bay, Puerto Culebra, Costa Rica, March 13, 1933; 6 from Station 965-
39, 8-15 fms., Tenacatita Bay, Mexico, May 8, 1939, and one from
Station 964-39, 2-8 fms., Tenacatita Bay, Mexico, May 8, 1939.
Remarks.—This species is related to O. variegata Liitken and
O. crassa Nielsen, differing from both by its depressed disk, broad, short,
and markedly tapering arms, the large, rounded interbrachial plates,
larger and broader oral shields, and color pattern. These characters are so
conspicuous that O. plateia cannot be confused with any known species of
the genus. The name selected, plateia, refers to the unusually depressed
disk.
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44 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 2
Sigsbeia laevis, new species
Fic. 1. Upper view
Fic. 2. Under view
Fic. 3. Side arm view
Ophiacantha phragma, new species
Fic. 4. Under view
Fic. 5. Upper view
Fic. 6. Side arm view
VOL. 8
NO. 2 ZIESENHENNE: NEW OPHIURANS PL. 2
a
SS
oe zmm.
46 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 3
Ophiomyces multispinus, new species
Fic. 1. Under view
Fic. 2. Upper view
Fic. 3. Side arm view
Fic. 4. Side view of disk
Ophiophragmus ophiactoides, new species
Fic. 5. Under view
Fic. 6. Upper view
Fic. 7. Side arm view
VOL. 8
PL. 3
ZIESENHENNE: NEW OPHIURANS
, eee
BO (| Keipsrees
Be LY esa
aa
48 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 4
Ophiophragmus papillatus, new species
Fic. 1. Under view
Fic. 2. Upper view
Fic. 3. Side arm view
Ophiophragmus lonchophorus, new species
Fic. 4. Upper view
Fic. 5. Under view
Fic. 6. Side arm view
Amphichondrius laevis, new species
Fic. 7. Under view
Fic. 8. Upper view
Fic.9. Side arm view
VOL. 8
NO. 2 ZIESENHENNE: NEW OPHIURANS PL. 4
Ma;
sia Doss
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50 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 5
Amphiodia sculptilis, new species
Fic. 1. Under view
Fic. 2. Upper view
Fic. 3. Side arm view
Ophiocnida californica, new species
Fic. 4. Upper view
Fic. 5. Under view
Fic. 6. Side arm view
Amphioplus philohelminthius, new species
Fic. 7. Under view
Fic. 8. Upper view
Fic. 9. Side arm view
NO. 2 ZIESENHENNE: NEW OPHIURANS PL. 5
TTS AN
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52 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 6
Ophionereis dictyota, new species
Fic. 1.
Fic. 2.
Fic. 3.
Fic. 4.
Fic. 5.
Under view
Upper view
Side arm view
Side arm view, mid arm
Upper arm view, mid arm
Ophiophragmus stellatus, new species
Fic. 6.
Fie: 7.
Fic. 8.
Fic. 9.
Upper view
Under view
Side arm view
Side view of disk showing
marginal disk papillae
PL.6
ZIESENHENNE: NEW OPHIURANS
NO. 2
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Tue UNIVERSITY OF SOUTHERN CALIFORNIA PUBLICATIONS
ALLAN HANCOCK PACIFIC EXPEDITIONS
VoLuME 8, NUMBER 3
IssUED JUNE 20, 1941
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
Los ANGELES, CALIFORNIA
THE HOLOTHURIOIDEA COLLECTED BY THE VELERO
III DURING THE YEARS 1932 to 1938
Part I, DENDROCHIROTA
(PLATES 10-30)
ELISABETH DEICHMANN
INTRODUCTION
The Panamic region, that is, the stretch of coast which extends from
Cerros Island on the western coast of Lower California and southward
(including the entire Gulf of California) to Ecuador (with the Galapa-
gos Islands) and Peru, forms zoogeographically a natural unit. As far as
holothurians are concerned, it has been almost unknown, undoubtedly
because the coast in most localities is so exposed that many species live at a
greater depth than in other places, as, for example, in the more sheltered
bays and coves of the West Indies.
The study of the fauna has tempted zoologists for many years, and as
early as 1866 Verrill began to compile lists of Echinoderms and Coelen-
terates from east and west of Central America. The purpose was chiefly
to accumulate data to show the relation of the two faunas to each other.
It was Alexander Agassiz’s idea that, since so many forms, especially
Echinoderms, in the two areas were closely related, one had here a chance
to study the effect of separation, or isolation, on the origin of new species,
since it was almost certain that the two faunas had once been identical—
before the isthmus had been formed. Verrill’s lists included 7 species from
the Panamic region; of these, 3 are Dendrochirotes. Since then a few
more have been described by various workers. ‘The general impression
gained was merely that the holothurian fauna was extremely poor and
showed some affinities to that of the West Indian seas. It is only recently
that the Allan Hancock Expeditions with the Velero III have been able
to explode this myth and show that the holothurian fauna actually is as
rich as that of the West Indies. During the past six years the Velero III
has made extensive collections in the Panamic region and particularly
dredgings in that zone hitherto so sadly neglected, between 10-100
fathoms. Almost all the species hitherto described from the region during
the last hundred years have been secured and in addition a large number
of new species. Thanks to the thorough and methodical way in which
[61]
62 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the collecting has been undertaken—in most cases under the able leader-
ship of Dr. Waldo L. Schmitt, with the enthusiastic support of several
young collectors—the material now assembled appears to be so complete
that one dares assert that very few additional species are to be expected.
Future investigators must be content to secure more material of the
species now known or additional information about the distribution of
the individual species.
The collections consist chiefly of material collected in less than 100
fathoms, or what is usually referred to as shallow water, in contrast to
the truly deep sea. The species are distributed among the following four
orders—Dendrochirota, Aspidochirota, Molpadonia, and Apoda; the
fifth order, Elasipoda, made up exclusively of deepwater forms and a few
Arctic species from shallow water, was not met with by the Velero. The
Dendrochirota and Aspidochirota have the greatest numbers of repre-
sentatives. The material of the former order has proved to be the most
interesting, because it includes a large number of new species which are
restricted to the Panamic region or have close relatives in the West Indies,
while the representatives of the Aspidochirota are mostly widespread
species. The Molpadonia and Apoda have so few representatives that it is
dificult to make any general statements about their relations to other
faunas.
In the following account all the Dendrochirota hitherto known from
the Panamic region are monographically treated. Besides the material
secured by the Velero III, a few undescribed specimens from the M.C.Z.
have been included to make the work complete. Much use has also been
made of Selenka’s old types and other material from the Panamic region
which I have had occasion to examine at various times. Special attention
has been paid to comparison with the holothurians from adjacent waters,
that is, the West Indian, Californian, Chilean, and Hawaiian seas. As far
as the Dendrochirotes are concerned, one may say that the affinities are
definitely with the West Indian fauna, while a few forms have related
representatives in Californian waters. Several species are also known
from the Chilean coast, but relationship to the Hawaiian fauna seems to
be nil. More extensive explorations may, however, show that a somewhat
richer Dendrochirote fauna exists in Hawaii than has hitherto been sup-
posed.
It is a great pleasure to thank the Allan Hancock Foundation for the
privilege of studying this valuable collection, thus enabling me to realize
my long-felt desire to write a monograph on the holothurians of the
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 63
Panamic region. Particularly do I wish to thank Dr. Irene McCulloch
for her unflagging interest in this paper and for the amount of her own
valuable time which she unstintingly has given whenever I worked in
Los Angeles.
COMPOSITION OF THE FAUNA
From the Panamic region 15 species of Dendrochirota, valid and
dubious, were described up to 1936; during the following years 6 new
species were added, thus bringing the total number up to 21.
The present monograph treats 43 species which are valid and 3 which
are so poorly described that they must be completely rejected, unless the
types should be rediscovered. Three of the 43 valid species are probably
extraterritorial—Pachythyone rubra (H. L. Clark) and Thyonepsolus
nutriens H. L. Clark, not known with certainty south of Santa Cruz
Island, southern California, and Psolus patagonicus Ekman from the
Cape Horn region. All the valid species have been examined ; of these, 34
are represented in the Allan Hancock Foundation collections, while the
remaining 9 have been studied in the M.C.Z. and in the U.S.N.M.; in
five cases the types have been available. Of the 34 species in the Hancock
collections 16 are new, while the 17th new species came from the collec-
tions of the Stranger, which are deposited in the M.C.Z.
List OF DENDROCHIROTA FROM THE PANAMIC REGION, INCLUDING A
FEW FROM THE ADJACENT NORTHERN AND SOUTHERN WATERS}
Cucumaria californica Semper p. 79 Leptopentacta nina, new species p. 97
Cucumaria chilensis Ludwig p. 80 Pentacta peruana (Semper) p. 99
Cucumaria dubiosa Semper p. 81 Thyonacta mexicana, new species p. 101
Cucumaria crax, new species p. $3 Thyone neofusus, new species p. 104
Cucumaria godeffroyi Semper (0)* p.83 Thyone bidentata, new species p. 105
Pentamera zacae Deichmann (0) p. 85 Thyone parafusus, new species p. 106
Pentamera beebei Deichmann (0) p. 86 Thyone strangeri, new species p. 107
Pentamera chierchia (Ludwig) p. 86 Neothyone gibber (Selenka) p. 109
Pentamera chiloensis (Ludwig) p. 88 Neothyone panamensis (Ludwig) p. 112
Pentamera panamensis Verrill (0)* p.89 | Neothyone gibbosa, new species p. 113
Neopentamera anexigua, new genus, Pachythyone rubra (H.L. Clark) (0) p.115
new species p. 90 Pachythyone lugubris (Deichmann) p. 116
Apentamera lepra, new genus, new Pachythyone pseudolugubris, new
species p. 92 species p. 116
Leptopentacta nova, new species p. 95 Athyone glasselli (Deichmann) (0) p. 119
Leptopentacta panamica, new species p. 96 Anaperus peruviana (Lesson) (0)* p. 120
+ Dubious species are marked with an *; those not secured by the Allan
Hancock Expeditions are marked with (0).
64 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Euthyonidium ovulum (Selenka) p. 124 Thyonepsolus hancocki, new species p. 140
Euthyonidium veleronis, new species p. 126 Psolidium dorsipes Ludwig p. 143
Athyonidium chilensis (Semper) p. 127 Psolidium eubullatum, new species p. 144
Pattalus mollis Selenka p. 130 Psolidium ekmani, new species p. 145
Phyllophorus aculeatus Ludwig (0) p. 133 Psolidium planum, new species p. 145
Phyllophorus zacae Deichmann p. 134 Psolus squamatus (Koren), var.
Thyonepsolus nutriens H. L. Clark p. 138 Segregatus Perrier (0) p. 147
Thyonepsolus beebei Deichmann p. 139 Psolus patagonicus Ekman (0) p. 148
Thyonepsolus veleronis, new species p. 140 Psolus diomedeae Ludwig p. 149
List oF STATIONS AT WHICH DENDROCHIROTE HOLOTHURIANS WERE
COLLECTED
11-32 (?). January 12, 1932. Conway Bay, Indefatigable Island,
shore, Galapagos.
Thyonepsolus nutriens H. L. Clark
(Possibly some error in the station number.)
73-33. February 13, 1933. Albemarle Island, Cartago Bay, shore,
Galapagos.
Pentamera chierchia (Ludwig)
80-33. February 15, 1933. Duncan Island, Smitty’s Shore, collecting.
In coral.
Pentamera chierchia (Ludwig)
132-34. January 4, 1934. Braithwaite Bay, Socorro Island, Mex.,
40 fms.
Pentamera chierchia (Ludwig)
Neothyone gibber (Selenka)
143-34. January 11, 1934. Wenman Island, Galapagos, 100-150 fms.
Psolus diomedeae Ludwig
145-34. January 12, 1934. North end of Albemarle Island, Galapa-
gos, 6-7 fms.
Pachythyone pseudolugubris, new species
147-34. January 13, 1934. South of Tagus Cove, Albemarle Island,
Galapagos, 30 fms.
Thyonepsolus veleronis, new species
166-34. January 19, 1934. Charles Island, Galapagos; shore collect-
ing on Black Beach.
Neothyone gibbosa, new species
189-34. January 25, 1934. Cartago Bay, Albemarle Island, Galapagos ;
shore, among Porites.
Pentamera chierchia (Ludwig)
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI,DENDROCHIROTA 65
190-34. January 26, 1934. Lat. 0°55’S., Long. 90°30’W., 58-60 fms.
Sand.
Pentamera chiloensis (Ludwig)
Thyone neofusus, new species
Phyllophorus zacae Deichmann
209-34. February 9, 1934. North of Point Sta. Elena, La Libertad,
Ecuador, 8-10 fms.
Pentamera chiloensis (Ludwig)
212-34. February 10, 1934. La Plata Island, Ecuador, dredging,
45-55 fms. Sand, shale, rock, mud.
Psolidium eubullatum, new species
213-34. February 10, 1934. La Plata Island, Ecuador, 7-10 fms.
Pentamera chiloensis (Ludwig)
Thyonepsolus hancocki, new species
Psolidium eubullatum, new species
Psolidium ekmani, new species
Psolus diomedeae Ludwig
244-34. February 21, 1934. Bahia Honda, Panama, dredging between
Medidor and Pacora islands, 30-35 fms.
Thyonacta mexicana, new species
245-34. February 21, 1934. Bahia Honda, Panama, dredging off north-
west point of Pacora Island, 15-25 fms.
Thyone bidentata, new species
249-34. February 22, 1934. Bahia Honda, Panama, dredging in 15-20
fms. Outside of island, south of bay. Rock.
Thyone bidentata, new species
251-34. February 22, 1934. Secas Islands, Panama, dredging south and
west of group in 15 fms. Rock.
Thyone bidentata, new species
259-34. February 28, 1934. Tangola Tangola, Mex., 15-20 fms.
Leptopentacta panamica, new species
261-34. March 1, 1934. Tangola Tangola, Mex., 15-20 fms.
In coral.
Cucumaria californica Semper
Euthyonidium ovulum (Selenka)
Thyonepsolus beebei Deichmann
66 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
264-34. March 2, 1934. South and west of White Friar Island, Petat-
lan Bay, Mex., 25 fms.
Pachythyone pseudolugubris, new species
Pentacta peruana (Semper)
272-34. March 4, 1934. Tenacatita Bay, Mex., dredging in 25 fms.,
between white rocks and bay. Coarse sand.
Thyone parafusus, new species
275-34. March 4, 1934. Off Navidad Head, Tenacatita Bay, Mex.,
25-35 fms.
Leptopentacta panamica, new species
Thyone bidentata, new species
Thyone parafusus, new species
281-34. March 7, 1934. West coast of L. Calif., Santa Maria Bay,
35-40 fms., 1 mile west of Hughes Point.
Leptopentacta nova, new species
283-34. March 9, 1934. Off Thurloe Point, Thurloe Bay, west coast
of L. Calif., 8-10 fms.
Cucumaria crax, new species
Pachythyone lugubris (Deichmann)
Thyonepsolus beebei Deichmann
286-34. March 9, 1934. Thurloe Point, west coast of L. Calif. Shore
collecting.
Cucumaria californica Semper
287-34. March 10, 1934. Cerros Island, west coast of L. Calif., 10-15
fms. (close to kelp beds).
Pachythyone lugubris (Deichmann)
315-35. December 8, 1934. Indefatigable Island, opposite Gordon
Rocks. In corals.
Pentamera chierchia (Ludwig)
365-35. January 10, 1935. Off southeast corner of Lorenzo Island, Cal-
lao, Peru, 10 fms.
Cucumaria dubiosa Semper
366-35. January 10, 1935. Between rocks south of Lorenzo Island,
Callao, Peru.
Cucumaria dubiosa Semper
375-35. January 13, 1935. Independencia Bay, Peru, shore.
Pattalus mollis Selenka
384-35. January 14, 1935. Independencia Bay, Peru, 5 fms., 34 mile off
shore, east side of bay.
Cucumaria dubiosa Semper
NO.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 67
391-35. January 17, 1935. Lobos de Afuera, Peru, shore of main island
with lighthouse, rocks.
Neothyone gibber (Selenka)
Neothyone gibbosa, new species
Pattalus mollis Selenka
401-35. January 19, 1935. Manta, Ecuador, 1 fm., below first rocky
point.
Pentamera chierchia (Ludwig)
423-35. January 25, 1935. Port Utria, Colombia, 20 fms., close to
shore.
Leptopentacta panamica, new species
Thyone bidentata, new species
429-35. January 27, 1935. Octavia Bay, Colombia, 30-35 fms., north
end of channel. Coarse sand and gravel.
Psolus diomedeae Ludwig
431-35. January 27, 1935. Octavia Bay, Colombia, 45 fms., north of
Octavia Strait, south end of channel.
Psolus diomedeae Ludwig
464-35. February 8, 1935. Playa Blanca, Costa Rica, 2 fms., in coral,
south shore of bay.
Pentamera chierchia (Ludwig)
465-35. February 8, 1935. Playa Blanca, Costa Rica, shale between
beach and rocky reef.
Euthyonidium veleronis, new species
Neothyone gibbosa, new species
466-35. February 9, 1935. Parker Bay, Costa Rica, shore of small is-
land at north side of bay.
Cucumaria californica Semper
Cucumaria dubiosa Semper
Neothyone gibber Selenka
467-35. February 9, 1935. Parker Bay, Costa Rica, 2 fms. Rock and
algae.
Pentamera chierchia (Ludwig)
473-35. February 9, 1935. Parker Bay, Costa Rica, 2 fms., in coral.
Pentamera chierchia (Ludwig)
Neothyone gibber (Selenka)
517-36. February 25, 1936. East side of San Francisco Island, Gulf of
Calif., 15 fms.
Phyllophorus zacae Deichmann
68 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
521-36. February 27, 1936. Agua Verde Bay, Gulf of Calif., 5-10 fms.
Thyonepsolus hancocki, new species
527-36. February 28, 1936. South of Mangles Anchorage, Gulf of
Calif., shore.
Cucumaria californica Semper
529-36. March 1, 1936. Off San Francisquito Bay, Gulf of Calif.,
165 fms.
Thyonepsolus hancocki, new species
Psolus diomedeae Ludwig
537-36. March 2, 1936. Spit, north of mill site, Angeles Bay, Gulf of
Calif., shore.
Cucumaria californica Semper
540-36. March 3, 1936. Puerto Refugio, Angel de la Guardia Island,
Gulf of Calif., shore.
Neothyone panamensis (Ludwig)
545-36. March 4, 1936. Puerto Refugio, Angel de la Guardia Island,
Gulf of Calif., west of rock spit at river wash, shore.
Neothyone gibbosa, new species
554-36. March 8, 1936. Parallel to Angel de la Guardia Island, Gulf
of Calif., east side, 10 fms.
Thyonacta mexicana, new species
557-36. March 8, 1936. Off White Rock, Isla Partida, Gulf of Calif.,
45 fms.
A pentamera lepra, new species
562-36. March 10, 1936. North of San Esteban Island, Gulf of Calif,
20-70 fms.
Thyonacta mexicana, new species
591-36. March 16, 1936. Port Escondido, shore, L. Calif., rock.
Neothyone gibbosa, new species
Euthyonidium ovulum (Selenka)
596-36. March 16, 1936. Port Escondido, Gulf of Calif., 20 fms.
Pentamera chierchia (Ludwig)
617-37. March 2, 1937. San Juanico Bay, southern end of L. Calif.,
24 fms.
Cucumaria californica Semper
623-37. March 4, 1937. Cabeza Ballena, east of Cape San Lucas, L.
Calif., shore.
Cucumaria californica Semper
NO.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 69
633-37. March 6, 1937. San Gabriel Bay, Espiritu Santo Island, Gulf
of Calif., 18 fms.
Thyonacta mexicana, new species
677-37. March 15, 1937. Ildefonso Island, Gulf of Calif., 50 fms.
Pentamera chiloensis (Ludwig)
679-37. March 15, 1937. Outside of Concepcion Bay, Gulf of Calif.,
30 fms.
Thyone bidentata, new species
683-37. March 15, 1937. Outside of Concepcion Bay, Gulf of Calif.,
12 fms.
Cucumaria californica Semper
Pentamera chierchia (Ludwig)
Neopentamera anexigua, new species
Thyone bidentata, new species
Pachythyone pseudolugubris, new species
686-37. March 16, 1937. Concepcion Bay, L. Calif., 12 fms.
Thyone bidentata, new species
699-37. March 19, 1937, Angeles Channel, Gulf of Calif., 30 fms.
Cucumaria chilensis Ludwig
719-37. March 24, 1937. Consag Rock, Gulf of Calif., 6-8 fms.
Thyonacta mexicana, new species
744-37. April 1, 1937. Near Point Piaxtla, Sinaloa, Gulf of Calif.,
6-8 fms.
Pachythyone pseudolugubris, new species
Psolidium dorsipes Ludwig
780-38. January 14, 1938. Chatham Bay, Cocos Island, Costa Rica,
40-47 fms.
Psolus diomedeae Ludwig
784-38. January 17, 1938. Darwin Bay, Tower Island, Galapagos,
Middle Beach, shore.
Pentamera chierchia (Ludwig)
792-38. January 20, 1938. Off Daphne Minor Island, Galapagos, 70-
80 fms.
Psolus diomedeae Ludwig
814-38. January 28, 1938. North of Hood Island, Galapagos, 20-40
fms.
Thyone neofusus, new species
70 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
816-38. January 29, 1938. North of Hood Island, Galapagos, 50-100
fms.
Phyllophorus zacae Deichmann
820-38. February 6, 1938. San Nicholas Bay, Peru, 10-25 fms.
Pattalus mollis Selenka
824-38. February 7, 1938. San Juan Bay, Peru, 15-20 fms.
Cucumaria californica Semper (poor specimen)
828-38. February 8, 1938. San Juan Bay, Peru, shore.
Athyonidium chilensis (Semper)
Pattalus mollis Selenka
831-38. February 9, 1938. Independencia Bay, Peru, east side of bay,
shore.
Athyonidium chilensis (Semper)
833-28. February 10, 1938. Independencia Bay, Peru, off north en-
trance, 8 fms.
Pentamera chiloensis (Ludwig)
837-38. February 11, 1938. North Chincha Island, Peru, shore.
Pattalus mollis Selenka
844-38. February 14, 1938. Lobos de Afuera Island, Peru, shore.
Neothyone gibber (Selenka)
Neothyone panamensis (Ludwig)
Neothyone gibbosa, new species
Euthyonidium ovulum (Selenka)
Pattalus mollis Selenka
845-38. February 15, 1938. Sechura Bay, Peru, 9.5 fms.
Psolidium planum, new species
850-38. February 23, 1938. Cape San Francisco, Ecuador, 15 fms.
Leptopentacta nina, new species
854-38. February 24, 1938. Gorgona Island, Colombia, north of island,
mud, rocks, 40-60 fms.
Thyonepsolus hancocki, new species
Psolidium planum, new species
Psolus diomedeae Ludwig
863-38. March 1, 1938. Bahia Honda, Panama, off North Island,
30-50 fms.
A pentamera lepra, new species
Psolidium dorsipes Ludwig
Psolidium ekmani, new species
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 71
Order DENDROCHIROTA
Diagnosis.—Plankton feeding holothurians with dendritic tentacles,
rarely fingerlike; number varying from 10, the 2 ventral often smaller,
in some cases less than 10, or 12-20, up to 30 in a few genera. Feet ar-
ranged along the ambulacra or also scattered in the interambulacra; ter-
minal and dorsal ones often modified into papillae.
Internally a calcareous ring, simple, or complex, with shorter or
longer posterior prolongations. One or more stone canals, one or more
Polian vesicles; usually a well-developed muscle stomach; intestine with
the 3 loops attached by mesenteries; that of the third loop may run along
the right or the left side of the midventral muscle band. Retractor muscles
well developed, distinctly separated from the longitudinal muscles. Re-
spiratory trees mostly well developed, usually attached to the lateral inter-
ambulacra. Gonads forming 2 tufts with a shorter or longer genital
stolon, and shorter or longer tubes, usually divided near the base.
Spicules tables, cups, buttons, or plates, et cetera, in some cases large
scales covering the entire body, or the dorsal side and the ends. Feet with
large end plate, or a vestige, or end plate completely reduced. Tentacles
with larger or smaller plates or rods, often rosettes, or no spicules at all.
In many forms the spicules disappear normally with advancing age, partly
or completely.
Remarks.—The order seems to be comparatively young. The members
occur with few exceptions in shallow water and the distribution of the
individual species is rather restricted aside from the deepwater forms and
certain Arctic and Antarctic forms.
Key TO THE FAMILIES OF DENDROCHIROTA
1. Part of the ventral side developed as a thin-walled creeping sole.
EN ts : : III. Psolidae p. 135
id No part r ee ae et side ereinen as a thin-walled creeping sole.
2. Tentacles 10 in number or less. . . . . i Cases p. 76
2. ‘Tentacles more than 10 in number (12-20, 30).
II. Phyllophoridae p. 122
Remarks.—The old division of dekachirote and polychirote forms,
designating those with 10 tentacles (or less) and those with more than
10, has been abandoned, as it is completely artificial.
72 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
CHARACTERS USED FOR IDENTIFICATION
Macroscopical characters
Size-—Where full-grown specimens are available, size constitutes a
fairly good character, although some caution must be used, as many species
are able to contract their bodies to an unbelievable degree, while others
rarely change much. Most Dendrochirotes are small forms, that is, less
than 10 cm. long, a number measure between 10-15 cm., and few are
large, above 20 cm. The largest forms are known to reach a length of
50 cm. when fully expanded. The chief difficulty about using size for
identification is that one so often encounters young individuals. Speci-
mens 1 cm. long are usually difficult to identify; the feet are frequently
not distributed in their typical pattern, and the spicules may be very
different from what they are in more mature individuals. Where very
young specimens are involved, the characters, “feet ambulacral” or “feet
interambulacral,” cannot be used, as all the forms with interambulacral
feet begin with feet in the ambulacra only. In some species the inter-
ambulacral feet begin to appear when the animal is less than 1 cm. long,
but in others they appear much later, when the animal is 2-3 cm. long.
For that matter, very little is known about the growth rate of the
Dendrochirotes or any order of seacucumbers. Mitsukuri found that the
Japanese Aspidochirote form Stichopus japonicus reaches the unbelievable
length of 25 cm. in a little over a year, but such a rapid growth is prob-
ably an exception, and under no circumstances can it be assumed to be
the normal in the Dendrochirotes without definite proof. Smaller species
may probably reach their full length in one or two years; larger forms,
as, for example, Thyone briareus (Lesueur), require undoubtedly four
to five years to reach their full size.
Tentacles —The full number is quickly reached in the “dekachirote”
forms. In the ‘“‘polychirote” forms (those which have tentacles arranged
in 2 well-defined circles and definitely of two sizes) seem to develop the
full number before they have reached the length of about 1 cm., while
in the species which have the 2 circles indistinctly set off and with ten-
tacles of varying size the full number seems to be reached rather late.
In many “dekachirote” forms the two ventral tentacles are much smaller
than the others, and the difference in size is retained throughout the
animal’s life. In a few species the number of tentacles falls below ten in
various deep sea forms and in that case the tentacles are often unbranched.
Tube feet—The arrangement of the tube feet is of importance in
reasonably mature individuals and constant for the different species. The
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 73
number of feet increases with advancing age. Many descriptions, ‘feet
in 3-4 rows,” “single row, etc.,” are of little value except when the ani-
mal is well expanded and its size also is given. In some species the feet
are heavily armed with spicules and unable to retract; in others equally
well armed they may be withdrawn. Where the feet contain few spicules,
the feet are often completely retractile. In younger individuals the ar-
rangement of the feet is often untypical, as mentioned above.
Inner organs —Much space has usually been devoted to the inner
anatomy of the Dendrochirotes, but with the exception of the calcareous
ring, the stone canal, Polian vesicle, and gonads, all other organs may
usually be ignored. A few species are said to lack a muscle stomach, but
I am not sure that the observation is correct. The course of the intestine
and the attachments of the mesenteries to the body wall are fairly stable
characters, although some variation has been noted, but the contracted,
contorted condition wherein the material often is preserved, or the com-
plete loss of inner organs, makes it impossible to use these structures
consistently. The Cucumariidae seem typically to have the third mesen-
tery attached on the right side of the midventral muscle band, the Phy/-
lophoridae seem to have it attached to the left, and the Psolidae have it
either way. The thickness or thinness of the longitudinal muscles, the
place of attachment for the retractors, the shape of the branches of the
respiratory trees, et cetera, are all characters of rather dubious value and
the use of them has often caused misidentification.
Calcareous ring—The calcareous ring presents a character of utmost
importance, as certain types invariably seem to be combined with a cer-
tain arrangement of the tube feet and certain types of spicules.
The calcareous ring is designated as simple when the radials and
interradials have fairly low, broad basal portions with shorter or longer
anteriorly projecting teeth, while the posterior margin may be almost
straight or undulated, sometimes with faint protuberances on the radials.
In the complex ring the radials have distinct posterior prolongations, the
so-called “‘tails”; and the body of the radials may be deeply cleft, some-
times almost to the base of the anterior tooth. In some species the tails may
be secondarily more or less resorbed or hidden in the tissue and hence less
noticeable. The interradials either are fairly broad, overlapping the ra-
dials as heart-shaped or diamond-shaped pieces, or are narrow, often
quite tall, rectangular, and firmly united with the radials; rarely are they
vestigial; the anterior tooth may be short or long. The complex cal-
careous ring may furthermore be composed of several smaller pieces as a
74 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
mosaic, but this “polyplacous” condition is perhaps not always an entirely
reliable character, as in certain species some specimens have polyplacous
rings, while in others the condition is little or not at all pronounced—
observed in Pentamera calcigera (Stimpson).
Stone canal and Polian vesicle—In most Dendrochirota there is a
single dorsal stone canal with round or oblong head, attached to the dor-
sal mesentery, or free, and one or two ventral Polian vesicles. In a few
species, particularly among the “polychirote” forms, there are several
free stone canals and often a large number of Polian vesicles also. In
some species the Polian vesicles have been found to be branching, but in
most cases they are simple.
Gonads.—Only few and scattered observations have been made on
the gonads in the holothurians and here also too much emphasis has been
laid on the size of these organs in different animals. In Stichopus japoni-
cus Mitsukuri observed that the sexually mature individuals, about 3
years old, in short time developed few long tubes of eggs and sperms,
which were shed, and the tubes shriveled up and new ones developed in
the following year. This condition is probably the typical one in the
aspidochirote forms. In the larger Dendrochirota it seems as if the gonad
tubes increase in size over a period of years. It is possible that some eggs
or sperms are developed before the full size of the tubes is reached. In
Thyone briareus (Lesueur) it has been observed (Kille, 1939) that the
number of tubes increases slightly with advancing age. The increase takes
place in the anterior part of the gonads, while a few shriveled-up tubes
are found posteriorly. A similar observation has been made on A thyonia-
ium chilensis (Semper). Very likely smaller, immature tubes will be
found to be present in most Dendrochirotes, which continue their growth
for several years. It is also possible that some of the so-called ‘“hermaphro-
ditic gonads” merely represent the mature and immature portions of a
unisexual gonad. In the larger species the number of tubes is large, more
than 100 in each tuft; in the smaller and supposedly more short-lived
forms the number of tubes in each tuft is low, varying between 5-20 or
thereabout, and no immature portion seems to be present. More observa-
tions are, however, needed before any definite statements can be made.
Microscopical characters
The spicules in the integument, feet, introvert, and tentacles form
valuable characters for identification of Dendrochirota as well as most
other holothurians. Certain types of spicules or derivates of these occur,
usually together, and accompany certain anatomical features. The spicules
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA tbe)
change considerably during the animal’s life, either becoming more com-
plex or degenerating; in some cases they disintegrate almost completely,
with the exception of the end plate which normally seems to persist in
the species where it is well developed from the beginning; when it is
rudimentary in the young individual, it quickly becomes resorbed.
In the integument two layers of spicules are typically present, an ex-
ternal, the function of which is presumably to give roughness to the skin,
and an inner layer, which gives stiffness. The spicules in the outer layer
seem definitely fixed in their position, usually with the external side
rough with spines, often perforating the skin, while the spicules in the
inner layer are able to slide over each other when the animal contracts—
as the cards in a deck—and these spicules are therefore more smooth and
flat. Either layer or both may be reduced or lacking; the inner layer is
never present in very young individuals, while the outer layer frequently
is lost or reduced with age. Usually the spicules in the 2 layers are ex-
tremely different, in other cases they are less sharply differentiated. Com-
mon types in the external layer are: the table, a basal plate with a num-
ber of pillars (1-6) ending in a varying number of teeth, or the basket
(or cup)—a hollow body usually with a number of spines developed on
the edge. Other types are oval buttons and plates, often with spines or a
rough reticulated mass on the external side. The inner layer consists of
regular or irregular buttons or plates, smooth or knobbed. Sometimes also
large and more complex bodies may be found.
A few forms have the body covered partly or completely by scales. In
that case the external layer of spicules is usually quite insignificant,
although it may contain rather interesting types of deposits.
In the tube feet a large end plate is almost invariably present in the
species which have well-developed tails on the calcareous ring. Likewise,
a well-developed end plate is usually accompanied by supporting tables
with a curved disk and a spire of varying development—sometimes the
spire is totally absent except in very young individuals, but it is usually
possible to ascertain whether a supporting rod is derived from the table
type or not. In more papilliform appendages the end plate is usually
small or lacking.
No end plate, or a vestigial end plate is usually combined with a
simple calcareous ring, and the supporting rods are either simple or pro-
vided with a third arm, more or less platelike, often irregularly star
shaped.
76 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
The introvert (the thin-walled skin between the base of the tenta-
cles and the anteriormost tube feet) often contains characteristic spicules.
In the species where the spicules are reduced gradually they are fre-
quently retained here. They usually resemble those found in the external
layer of the skin but are often more delicate and more complex. The
tentacles may be filled with large perforated plates or rods, or contain
minute rods or delicate plates, rarely a mixture of both. Rosettes seem
typical of certain species; they are apparently restricted to tropical and
subtropical forms and occur often in both the introvert and the tentacles.
In many cases the spicules in the tentacles become completely reduced
with advancing age.
Family I. Cucumariidae
Kery TO THE GENERA OF CUCUMARIIDAE KNOWN FROM THE
PANAMIC REGION
1. Body more or less globose, covered by large, reticulated scales.
Deepwater forms. . . . . . . Sphaerothuria Ludwig
1. Body not globose, not covered by large, reticulated scales. . 2
2. Spicules spinous crosses. Deepwater forms. . ‘
Unaware) telyrks. arg Staurocucumaria eee
Spicules notuspmous crosses: 6s 92.7 we Se ee
2s 3
3. Meet! restricted to ambulacra:. <1) “s \le ce fas le alls 4
3. Feet also found in the interambulacra. : GT
4. Feet retractile, soft, with end plate reduced or ipeeiae Seu
plates or buttons, often reduced. ;
ERIN aig 1. Cucumaria Blainville ey p 77
4. Feet not retractile . . . . Saawie es
5. Skin covered by huge reticulated ee aa sates spicules.
Calcareous ring at most with short posterior prolongations.
: y 5. Leptopentacta H. L. Clark p. 92
oF Skin not es by huge reticulated bodies. Calcareous ring
with long posterior prolongations. . . Pk de 6
6. Spicules 2-pillared tables, spire ae a sedeed Feet ith
large end plate. Spicules rarely reduced. 2. Pentamera Ayres p. 84
6. Spicules knobbed buttons, or irregular. Feet with large end
plat. . . . . . =. . . 3. Neopentamera, new genus p. 90
7. Feet occur in interambulacra but chiefly in the dorsal ones. 8
77
No.3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA
7. Feet occur in interambulacra but chiefly in the ventral ones.
(iilivanie Sole.) ae ce aes it Sas a ae ea
8. Dorsal feet normally developed, soft, retractile. Spicules plates
or buttons, often reduced. 1. Cucumaria Blainville (partim) p. 77
8. Dorsal feet mostly papilliform. Spicules few or numerous. 10
9. Dorsal interradial feet few, small, easily overlooked; radial
feet numerous. Resembles Pentamera.
4. Apentamera, new genus p. 91
9. Dorsal feet partly large, wartlike or papilliform. Spicules
crowded. . .... . . . . 6. Pentacta Goldfuss p. 96
10. Spicules numerous, forming a crowded layer. 11
10. Spicules more or less reduced, scattered, or lacking. 14
11. Spicules as 2- or 4-pillared tables. Calcareous ring with long
posterior prolongations. . . . . . . 8. Thyone Oken p. 102
11. Spicules as knobbed buttons. 12
12. Feet at first comparatively few, in distinct rows along the am-
bulacra, later crowded in the interambulacra, mostly papilli-
forme. sw ee Pe yonacta, new genus-p. 101
12. Feet numerous, but in almost indistinct rows along the am-
DULAGrale ni at eee Ore Wes) Sut gee dae Pay eee. Ves tee LO
13. No external plates covered by reticulum. Either baskets or
buttons with spinous handle. . 9. Neothyone, new genus p. 108
#3. sExternalplatescovered by a reticulum: <0 9/666 6.4% 3.
10. Pachythyone, new genus p. 114
14. Spicules swollen buttons, often obviously derived from tables.
Calcareous ring with long, well-developed posterior prolonga-
fianssit. ert! ie Lowe as Wh. Athyone, new genus p. 118
14. Spicules apparently lacking except the end plate in the feet.
Calcareous ring with short posterior prolongations sometimes
reduced... 2) 2 ./ es. « «& 12. Anaperus Troschel p. 120
Genus 1. CUCUMARIA Blainville
Diagnosis —Dendrochirotes with 10 tentacles, of equal size or the
ventral ones smaller. Feet large, soft, in 5 bands, in some forms also scat-
tered in the interradii, particularly dorsally. Calcareous ring low, simple.
78 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Spicules plates or buttons, knobbed or smooth, often reduced. Feet with
a vestige of an end plate or none at all; walls supported by rods, often
3 armed, or plates. Introvert and tentacles with perforated plates and
rods. The spicules show tendency to become reduced in many forms.
Type species —Cucumaria frondosa (Gunnerus).
Remarks.—The diagnosis embraces only the members of the genus
Cucumaria sensu strictiore. The well-known type species is the only rep-
resentative of the genus in the northern Atlantic, while the northern
Pacific, including Bering Sea, harbors 3 large species: C. japonica Semper,
C. fallax Ludwig, and C. miniata Brandt. From the west coast of North
America a number of smaller forms have been described: C. vegae Théel,
C. lubrica H. L. Clark, C. curata Cowles, C. pseudocurata Deichmann.
From the Panamic and Peruvian-Chilean region 4 species are reported
with certainty, while a fifth species, from Chile, imperfectly described, is
briefly discussed below.
Key TO THE SPECIES OF Cucumaria s. str. KNOWN FROM THE PANAMIC
REGION
1. Spicules small, crackerlike buttons or plates with marginal
holes or incisions. . . . 4. Cucumaria crax, new species
1. Spicules larger plates and buttons, knobbed to smooth, often
reduced in older individuals /, (5) 6 bo. ase 2 ee
2. Feet not restricted to the ambulacra, fairly numerous in the
dorsal interambulacra. . . 3. Cucumaria dubiosa Semper
2. (eet restricted'to the ambulacra. ©: 3’. 2 2. 2s & OS
3. Tentacles soft, bushy, of equal size. Spicules usually almost
lacking in larger individuals (6-10 cm. long), in young indi-
viduals as knobbed oblong plates, often with a dentate handle;
supporting rods in feet predominantly 3-armed rods. . . .
1. Cucumaria californica Semper
3. Tentacles rigid with spicules, not bushy, ventral ones small.
Spicules strongly knobbed circular plates or smooth ones, usu-
ally without a spinous handle. Feet with narrow, bandlike sup-
porting rods. . . . . . 2. Cucumaria chilensis Ludwig
Remarks.—Not included in the key is Cucumaria godeffroyi Semper,
which possibly is a strictly southern form. See p. 83.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 79
1. Cucumaria californica Semper
Plate 10, Figs. 6-8
Cucumaria californica Semper, 1868, p. 235, pl. 39, fig. 16; pl. 40, fig.
10.—Lampert, 1885, p. 147.—Théel, 1886, p. 109; 1886a, p. 8.
Nec Cucumaria californica Edwards, 1910, p. 601 (i.e., C. fallax Lud-
wig—an Arctic species, known from Bering Sea).
Diagnosis—Medium-sized form (10 cm.), with soft, bushy tentacles
of equal size. Skin soft, slippery; feet large, soft, completely retractile,
restricted to the ambulacra. Calcareous ring simple, other anatomical
features very nearly as in the type species. Spicules oblong knobbed plates,
often with a spinous handle, and 4-holed buttons, frequently reduced or
lacking. Feet with a vestige of an end plate and numerous 3-armed per-
forated supporting rods. Introvert and tentacles with perforated plates
or rods, often completely reduced. Color varying from dark slate gray
or black to almost white; tentacles seem always to be darkly pigmented.
Holotype.—Possibly in Germany.
Type locality —Mazatlan, Mexico.
Distribution.—West coast of Mexico and Central America, possibly
to Peru.
Depth.—F rom shore to about 100 fms.
Specimens examined.—Numerous in the U.S.N.M., and the follow-
ing from the Allan Hancock Expeditions:
261-34. Tangola Tangola, Mex., 15-20 fms., March 1, 1934, 4 speci-
mens.
286-34. Thurloe Bay, Thurloe Point, L. Calif., Mex., shore, March 9,
1934, 8 specimens.
466-35. Parker Bay, Costa Rica, shore, February 9, 1935, 3 specimens.
527-36. South of Mangles Anchorage, L. Calif., Mex., shore, Feb-
ruary 28, 1936, 1 specimen.
537-36. Spit, north of mill site, Angeles Bay, L. Calif., Mex., shore,
March 2, 1936, 10 specimens.
617-37. San Juanico Bay, L. Calif., Mex., 24 fms., March 2, 1937, 2
specimens.
623-37. Cabeza Ballena, east of Cape San Lucas, shore, March 4, 1937,
2 specimens.
683-37. Concepcion Bay, L. Calif., Mex., 12 fms., March 15, 1937,
1 specimen.
824-38. San Juan Bay, Peru, 15-20 fms., February 7, 1938, 1 specimen.
80 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Remarks.—The species seems to be common in the Gulf of Cali-
fornia and along the west coast of Mexico and Central America. A single
record exists from Peru, possibly an error in labeling. Its soft skin, bushy
tentacles, and usually dark coloration make it easy to recognize. In life
it is often bright red; in alcohol it fades to white with dark tentacles.
The name californica has unfortunately been applied to the common
frondosa-like form from Bering Sea, C. fallax Ludwig. During the
Albatross expeditions a single specimen of this Arctic form evidently got
mixed up with some material from the Galapagos Islands; Edwards,
who worked over the material, gave the name californica to this specimen
with the dubious locality, as well as to the larger individuals of C. fallax
from Bering Sea (while the smaller individuals were identified as C.
fallax). ‘The supposedly Galapagos specimen, which shows strong signs
of having been dried up, is a typical huge, brownish, frondosa-like form
and cannot be separated from the individuals of C. fallax of the same
size collected in the Bering Sea.1 Cucumaria californica, on the other
hand, is much smaller and either completely slate gray or black, or the
tentacles and part of the anterior end are dark colored. Its spicules are
entirely different from those found in C. fallax.
2. Cucumaria chilensis Ludwig
Plate 11, Figs. 3-5
Cucumaria chilensis Ludwig, 1875, pl. 6, fig. 11 (used only in the plate
explanation).
Cucumaria exigua (partim) Ludwig, 1874, p. 84.—Lampert, 1885, p.
145.—Théel, 1886, p. 108.
Diagnosis—Medium-sized form. Feet restricted to ambulacra, re-
tractile. Tentacles not bushy, filled with spicules, the 2 ventral ones small.
Calcareous ring simple; other anatomical features apparently typical of
the genus.
Spicules 4-holed buttons and smooth to knobbed plates. Feet with
rudimentary end plate and numerous slender supporting rods, slightly
shorter in the dorsal appendages. Introvert and tentacles with large per-
forated plates and rods. Color yellow, with or without irregular spots,
or mottled gray and brown.
Holotype-—Hamburg Museum.
Type locality—Chile.
Distribution.—Various localities in Chile.
1 Material in U.S.N.M.
NO.3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA 81
Depth.—Not recorded for the type and paratypes.
Specimens examined.—A single specimen from Velero III Station
699-37, Angeles Channel, Gulf of California, 30 fms., March 18, 1937.
Remarks.—The name chilensis has been re-established, since it is al-
most unbelievable that the species from China is identical with the
Chilean form. Ludwig evidently considered the 2 species as different and
then decided to unite them in spite of the differences which they ex-
hibited. It is not clear from his description whether he found any cross-
shaped bodies in the Chilean specimens (present in C. exigua from
China). The combination of characters, viz., simple ring and small ven-
tral tentacles and the unusually narrow supporting rods in the feet, is
such that it seems almost certain that the present species is identical with
the one described by Ludwig. Most of the plates found in the Velero
specimen are considerably more complex than indicated by Ludwig, but
simple 4-holed buttons are also present, and it is well known that the
spicules are often more or less reduced in this group. (The earlier
authors, often on general principle, avoided figuring the more complex
spicules, as study of certain holotypes has revealed, and figured only those
which could be easily reproduced.)
The Velero III specimen measures 3 cm. in length, but is so strongly
retracted that it probably measured 10 cm. or more when expanded. Lud-
wig’s types measured about the same, but it is not known whether they
were very strongly contracted or not.
3. Cucumaria dubiosa Semper
Plate 11, Figs. 1-2
Cucumaria dubiosa Semper, 1868, p. 238, pl. 39, fig. 19.—Lampert,
1885, p. 151.—Théel, 1886, p. 111.
Cucumaria leonina Ludwig, 1898, p. 36.—Ekman 1925, p. 52, text fig-
ure 10.
Nec Cucumaria leonina Semper, 1868, p. 53, pl. 15, fig. 9.
Diagnosis—Medium-sized form, length up to 10 cm. Feet in 5
bands and numerous in the dorsal interambulacra. Tentacles bushy, of
unequal size. Color uniformly parchmentlike (in alcohol). Calcareous
ring simple, other features as in the typical forms. Spicules numerous,
oblong, knobbed plates, often with spinous handle? and 4-holed buttons.
2In very young specimens (1 cm. long) a number of almost circular plates
with a long narrow handle are found (see Ekman’s text figure). Such plates with
handles have not been discovered in Cucumaria lubrica from the western coast
of North America.
82 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Feet with rudimentary end plate, or none at all, and numerous support-
ing rods, mostly 3 armed. Introvert and tentacles with perforated plates
and rods.
Type.—Possibly in Germany.
Type locality.—Coast of Peru.
Distribution—From Peru to the south end of South America, in-
cluding Falkland Islands, also Costa Rica. See Ekman, 1925.
Depth.—Not recorded for the type.
Specimens examined—Numerous from the following stations of
Velero III:
365-35. Callao, Peru, off southeast corner of Lorenzo Island, 10 fms.,
January 10, 1935, 21 specimens.
366-35. Callao, Peru, between rocks, south of Lorenzo Island, 8 fms.,
January 10, 1935, 1 young specimen.
384-35. Independencia Bay, Peru, 34 mile off shore, east side of bay,
5 fms., January 14, 1935, 1 young specimen.
466-35. Parker Bay, Costa Rica, small island at north side of bay,
shore, February 9, 1935, 1 specimen. This record so far north
is unusual.
Remarks.—The examined specimens range in size from 1.5 cm. to 6
cm. Even the smallest individual has a number of tube feet scattered
in the dorsal interambulacra and therefore cannot be mistaken for C.
californica of the same size; the latter species has, moreover, darkly pig-
mented tentacles.
The spicules resemble those found in Cucumaria lubrica H. L. Clark
(including C. fisheri Wells) from the west coast of North America, and
it is rather difficult to express the differences in such variable forms. The
northern form seems always to have numerous 4-holed buttons which
apparently sometimes are lacking in the southern form.
The name dubiosa has been preferred to /eonina against such authori-
ties as Ludwig (1887 and 1898) and Ekman (1925). Semper’s leonina
is supposed to have come from Singapore, and it is only a surmise that
it is identical with C. dubiosa. Since it is very unlikely that the same
species occurs in both Singapore and Chile and since the Chilean (and
Peruvian) locality is above reproach, it has been concluded that the
locality Singapore was wrong. But it is quite possible that C. Jeonina came
from Singapore; species with similar spicules are known from other
places, for example, C. ollikeri from the Mediterranean Sea and C.
salmini from Celebes. The latter was listed by Lampert, 1885, as occur-
No.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 83
ring in the Strait of Magellan, and since then it was concluded that the
locality Celebes was wrong. Very likely salmini from Celebes is a syno-
nym of Jeonina from Singapore but different from the species from Peru
and Chile.
4. Cucumaria crax, new species
Plate 10, Figs. 1-5
Diagnosis—Small form (?). Skin soft, smooth; feet large, soft,
completely retractile, arranged in 5 bands, apparently not present in
the interambulacra. Tentacles soft, bushy, of equal size. Calcareous ring
low, simple, strongly undulated posteriorly; radials deeply incised; a
single stone canal attached in the dorsal mesentery, 2 ventral Polian
vesicles. Retractors short, fleshy, gonads attached near the middle of the
body. Spicules, a scattered layer of small biscuit-shaped bodies, usually
with 4-8 marginal holes, frequently incompletely closed, and knobbed
margin. Feet with rudimentary end plate and short 3-armed supporting
rods. Introvert and tentacles, respectively, with buttons and numerous
narrow rods with perforated ends, large in the stem, small and almost
hair fine in the ends of the tentacles. Color mottled brown with paler
ambulacra; tentacles black.
Type.—Holotype, AHF no. 23, 6 paratypes.
Type locality—Station 283-34, Thurloe Point, Thurloe Bay, west
coast of L. Calif., 8-10 fms., March 9, 1934.
Distribution—Known from the type locality.
Depth.—F rom 8-10 fms.
Specimens examined.—The type and 6 paratypes.
Remarks.—The type measures 1.5 cm. in length and is strongly con-
tracted. The gonads are well developed, but this fact in itself constitutes
no clue to the size which the species may attain when full grown.
Superficially it resembles a small specimen of C. californica Semper,
but the spicules are entirely different. If the spicules disappear completely
as the animals grow larger, it may become impossible to distinguish be-
tween the two forms.
5. Cucumaria godeffroyi Semper
Cucumaria godeffroyi Semper, 1868, p. 53, pl. 15, figs. 12, 14.—Lam-
pert, 1885, p. 144.—Théel, 1886, p. 99.—Clark, 1910, p. 352.
84 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Nec Holothuria crocea Lesson, 1830, p. 153, pl. 52. Suggested by Semper
but not accepted by later writers.
Description ——Imperfectly known. Presumably the feet are restricted
to the ambulacra and the tentacles probably of equal size. The calcareous
ring is simple. The Polian vesicle is single; the stone canal attached in
the dorsal mesentery. Spicules perforated plates with laciniated edge and
one end narrower with long marginal spines, this end projecting through
the skin.
Type.—Hamburg.
Type locality.—Iquique, Chile.
Distribution.—Coast of Chile.
Depth.—Not noted.
Specimens examined.—None.
Remarks.—No specimen seems to have been secured since the type
was described. The type measured 3.5 cm. in length, possibly strongly
contracted. The interambulacral feet may have been overlooked. The
spicule figured could be interpreted as a degenerate plate from an aged
C. dubiosa.
Genus 2. PENTAMERA Ayres, 1852
Pentamera Ayres, 1852, p. 207.—Deichmann, 1938, p. 373; 1938a,
p. LOD.
Diagnosis—Small to medium-sized forms; ventral tentacles small;
feet long, nonretractile, arranged in 5 bands but never scattered in the
interambulacra. Calcareous ring with long posterior prolongations on
the radials. Spicules 2-pillared tables or derivatives from these with the
spire reduced or developed as acornlike bodies. Feet with large end plate
and supporting tables, usually with well-developed spire; in some forms
the spire is more or less completely reduced. ‘Tentacles with rods or
plates, in some forms devoid of spicules, at least in the older individuals.
Spicules in most forms numerous throughout the animal’s life, in some
species the spicules are few and degenerate.
Type species —Pentamera pulcherrima Ayres.
Remarks.—The diagnosis has been modified to include also Pentam-
era chierchia Ludwig and P. zacae Deichmann, the former with few
and mostly reduced spicules, the latter with peculiar acorn-shaped bodies
—as it seems unwise for the present moment to segregate these two
forms which otherwise conform so well with the typical members of the
genus.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 85
‘The type species seems to be the only representative known from
West Indian waters, while P. calcigera (Stimpson) is widespread in the
northern Atlantic and Pacific oceans, and 7 related species have devel-
oped along the western coast of North America (see Deichmann, 1938).
From the Panamic region (and Chile) 4 species are known. One of
these is closely related to the type species.
Key TO THE SPECIES OF Pentamera KNOWN FROM THE PANAMIC
REGION
1. Spicules in skin acorn-shaped bodies, possibly derived from
tables. Feet with end plate and curved pales tables with
well-developed spire. Color white. . . . ‘ a
erates eee Pa ep as eer Pentamera-sacde Deickane
1. Spicules in skin 2-pillared tables or derivatives of these (with
spire reduced to knobs-or spines). : . << 2... +... 2
2. Spicules scattered, tables reduced to disks with spinous edge
and mostly with 2 knobs or spines indicating the spire. Feet
with large end plate and a few spectacle-shaped rods, rarely
any trace of a spire. Color dark brown to black. . . . .
foe eS oe se 2 © od. Pentamera chierchia (huaweiet
2. Spicules crowded tables with well-developed spire, disk with
smooth edge. Feet with end plate and supporting tables with
well-developed to excessively developed spires . . . . . 3
3. Supporting tables in feet partly with excessively tall spire.
Tables small. . . . . . 2. Pentamera beebei Deichmann
3. Supporting tables in feet with moderately tall spire. Tables
not small. . . . . . 4. Pentamera chiloensis (Ludwig)
1. Pentamera zacae Deichmann
Plate 12, Figs. 10-17
Pentamera zacae Deichmann, 1938, p. 375, text fig. 9.
Diagnosis.—Small form (few cm. long) body strongly curved, taper-
ing toward both ends. Feet cylindrical, nonretractile, in 5 bands, most
numerous on the ventrum, more scattered on the dorsum and toward the
oral and anal ends. Spicules a crowded layer of peculiar small bodies re-
sembling acorns, with a tapering spire, mostly composed of 2 rods and a
basal cup-shaped part. Feet with large end plate and numerous support-
86 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
ing tables with curved disk with 4 central holes and a small hole in each
end, spire with 2 pillars and ending in a few lobes or blunt teeth which
lie in one plane. Color white.
Type.—M.C.Z.
Type locality.—Zaca Sta. 196 D-17.
Distribution —Tangola Tangola Bay, Mex.
Depth—From 23 fms.
Specimens examined.—The type.
Remarks.—The type measures about 3 cm. in length. The skin is
rigid with spicules. The oral end with the tentacles and calcareous ring
is lost as well as most of the inner organs. It is therefore only tentatively
that the species is referred to Pentamera.
2. Pentamera beebei Deichmann
Plate 12, Figs. 1-9
Pentamera beebei Deichmann, 1938, p. 374, text fig. 7.
Diagnosis.—Typical small pentamerid with minute tables (diameter
0.03 mm.) with mostly 4 larger holes and 4 smaller ones; spire 2 pil-
lared, ending in a tuft of spines. Feet with large end plate and curved
supporting tables with from 2-7 crossbeams in the spire and a terminal
tuft of flat lobes. Color white.
Type—M.C.Z.
Type locality —Zaca Sta. 213 D-15.
Distribution Known from Ballena Bay, Gulf of Costa Rica.
Depth.—F rom 40 fms.
Specimens examined.—The type.
Remarks.—The type measures few cm. in length. Its striking spicules
set it apart from all other species known from the Panamic waters.
3. Pentamera chierchia (Ludwig)
Plate 13, Figs. 19-21
Cucumaria chierchia Ludwig, 1887, p. 13, pl. 1, fig. 5.
Pentamera chierchia Deichmann, 1938, p. 374, text fig. 8.
Diagnosis —Small form (about 3-4 cm. long) with numerous feet in
5 narrow bands. Calcareous ring with long posterior prolongations.
Spicules in varying numbers, in some individuals almost completely lack-
ing. Skin with 4-holed tables with dentate margin and 2-pillared spire
with few irregular teeth; often the spire is reduced to knobs or com-
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 87
pletely lacking. Feet with large end plate and a varying number of mostly
spectacle-shaped rods, often with 4 central holes incomplete or complete,
and a number of smaller holes in the ends. Introvert with a few tables.
Tentacles with curved perforated plates and rods. Color black, rarely
brown with pale underside.
Type.—Possibly in Germany.
Type locality—Coast of one of the islands in the Gulf of Panama.
A single specimen secured.
Distribution—From Ecuador and Galapagos to L. Calif.
Depth.—Shore to 40 fms.
Specimens examined.—The following from the Allan Hancock Expe-
ditions:
73-33. North Beach, Cartago Bay, Albemarle Island, Galapagos,
shore, February 13, 1933, 22 specimens.
80-33. Smitty’s Shore, Duncan Island, Galapagos, shore, February 15,
1933, 15 specimens.
132-34. Braithwaite Bay, Socorro Island, Mex., 40 fms., January 4,
1934, 13 young specimens.
189-34. Cartago Bay, Albemarle Island, Galapagos, shore, January 25,
1934, 5 specimens.
315-35. Indefatigable Island, opposite Gordon Rocks, coral, 1-2 fms.,
December 8, 1934, 3 specimens.
401-35. Manta, Ecuador, below first rocky point, 1 fm., January 19,
1935, 5 specimens.
464-35. Playa Blanca, Costa Rica, 2 clumps of coral, south shore of bay,
shore, February 8, 1935, 2 specimens.
467-35. Parker Bay, Costa Rica, rock and algae, 2 fms., February 9,
1935, 2 specimens.
473-35. Parker Bay, Costa Rica, coral, 2 fms., February 9, 1935, 39
specimens.
596-36. Port Escondido, L. Calif., 20 fms., March 16, 1936, 1 speci-
men.
683-37. Outside of Concepcion Bay, L. Calif., Mex., 12 fms., March
15, 1937, 17 specimens.
784-38. Darwin Bay, Tower Island, Galapagos, Middle Beach, shore,
January 17, 1938, 7 specimens.
Also several from the Zaca expedition, 1938.
Remarks.—Apparently one of the most common forms in the Pan-
amic region.
88 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
4. Pentamera chiloensis (Ludwig)
Plate 13, Figs. 15-18; text figure 1
Cucumaria chiloensis Ludwig, 1887, p. 12, pl. 1, fig. 4—Deichmann,
1938, p. 373, passim.
Cucumaria tabulifera R. Perrier, 1904, p. 14; 1905, p. 17, pl. 1, figs.
4-5; pl. 3, figs. 1-5.
Diagnosis—Typical pentamerid, resembles the type species. Spicules
tables with oval to squarish disk (diameter 0.06 mm.) with 4 or 8 holes;
spires 2 pillared with one or two crossbeams and a few short teeth at
the top. Feet with large end plate and numerous curved supporting tables
with low spire with 2 pillars and a few flat teeth on the top. Introvert
with rosettes; tentacles with delicate rods. Color white.
Type.—Possibly in Germany.
Type locality —Chiloe Island, Chile.
Distribution —Cape Horn and vicinity to L. Calif.
Depth.—The type came from 40 m. depth; the Yel/ero specimens were
taken at from 8-60 fms. depth.
Specimens examined.—The following material from the Hancock
Expeditions:
190-34. Lat. 0° 55’ S., 90° 30’ W., Galapagos, 58-60 fms., January
26, 1934, 1 specimen.
209-34. La Libertad, Ecuador, north of Point Sta. Elena, 8-10 fms.,
February 9, 1934, 1 specimen.
213-34. La Plata Island, Ecuador, north of anchorage, 7-10 fms., Feb-
ruary 10, 1934, 1 specimen.
677-37. Ildefonso Island, L. Calif., Mex., 50 fms., March 15, 1937,
3 specimens.
833-38. Independencia Bay, Peru, off north entrance, sand and shell, 8
fms., February 10, 1938, 1 specimen.
Remarks.—Vhe species resembles the type species from the West
Indies. Ludwig’s specimen measured only 9 mm., while Perrier had two
specimens, 20 and 23 mm. long. The Velero material falls about within
these limits. The species may be expected to reach a length of about 5
cm.; the tables in younger individuals may possibly have larger disks
with 2 circles of holes; some tables have a dentate edge as if one row
of holes had been resorbed. It differs from C. pulcherrima in the lack of
perforated plates in the tentacles and the presence of rosettes in the
introvert, and the smaller size—characters which may prove to be abso-
No.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 89
Text fig. 1. Pentamera chiloensis (Ludwig) ?
Spicules from juvenile individual.
Scale divisions indicate 499 mm.
lutely worthless. A small specimen from Sta. 190-34, 1.5 cm. long, had
the genital tubes packed with ripe eggs.
The very young individual from Station 209-34 had much smaller
spicules than the other individuals, but it can hardly be doubted that it
represents P. chiloensis.
Doubtful form:
Pentamera panamensis (Verrill)
Ocnus panamensis Verrill, 1867, p. 321—DLampert, 1885, p. 132.—
Théel, 1886, p. 116.
From Verrill’s incomplete description it can only be learned that the
type measured about 4 cm. in length; the feet were restricted to the
ambulacra, in double rows on the ventrum, in single rows toward the
ends and on the dorsum; the tentacles were 10 in number and the 2
ventral smaller. The skin was thin, coriaceous, filled with numerous
minute calcareous grains or plates. The color was grayish brown, the
tentacles yellowish brown.
Type.—Apparently lost.
Type locality —Gulf of Panama.
Distribution—Known only from the type locality.
Depth.——Presumably shallow water.
Specimens examined.—None.
Remarks.—This may be any of the species belonging to Pentamera,
Apentamera, or Neopentamera, except P. chierchia and A. lepra. The
90 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
few feet on the dorsal side suggest P. beebei, while the description of the
spicules suggests P. zacae. As it is completely impossible to place the
species, the name must once and for all be rejected.
Genus 3. NEOPENTAMERA, new genus
Diagnosis.—Externally resembling Pentamera Ayres, with tube feet
restricted to the 5 ambulacra (?). Calcareous ring with well-developed
posterior prolongations. Spicules irregular, 4-holed, knobbed buttons;
feet with end plate and narrow ribbonlike rods, possibly derived from
tables.
Type species—Neopentamera anexigua, new species.
Remarks.—The genus and species are based upon a number of small
specimens, apparently adult, secured by the Hancock Expeditions. As the
specimens are so small, there is of course the possibility that tube feet
may appear later in the interradials, which will necessitate a modifica-
tion of the diagnosis. The spicules bear some resemblance to those of
Eupentacta pseudoquinquesemita Deichmann from the coast of Alaska
but are so much smaller that I have felt justified in establishing a new
genus for this species. It cannot be confused with any other dendrochirote
form hitherto recorded from the Panamic region.
Neopentamera anexigua, new species
Plate 13, Figs. 11-14
Diagnosis.—Small form (few cm. long), externally as internally a
typical pentamerid. Spicules numerous, irregularly knobbed buttons or
plates. Feet with large end plate and ribbonlike supporting rods with no
indications of a spire. Introvert with plates and rosettes. ‘Tentacles with
rosettes and delicate rods derived from rosettes. ‘Tentacles dark brown.
Color white.
Type.—Holotype, AHF no. 24, one paratype.
Type locality—Velero III Station 683-37, outside of Concepcion
Bay, L. Calif., Mex., 12 fms., March 15, 1937.
Distribution —Known only from the type locality.
Depth.—Taken at 12 fms. depth.
Specimens examined.—The type and a paratype.
Remarks.—The types are poorly preserved and measure only 2 cm.
NO.3 DEICHMANN: HOLOTHURIOIDEA; PARTI,DENDROCHIROTA 91
in length. In the type the gonads are large; many tubes measure almost
2 cm.; presumably it is a male, as no traces of eggs were seen.
It is possible that an external layer of more delicate spicules normally
is present but has been destroyed. The spicules are slightly reminiscent
of those characteristic of Eupentacta pseudoquinquesemita Deichmann,
from the Alaskan waters, but they are smaller, and the tails on the cal-
careous ring are distinctly longer in the present species.
The name anexigua is given because it was at first glance assumed
that the present species was identical with Ludwig’s Cucumaria exigua
from China and, supposedly, also Chile (see p. 81). The species from
Chile is now called C. chilensis, as indicated in Ludwig’s figures; it was
actually rediscovered by the Allan Hancock Expedition (see p. 80).
Dissection revealed, however, that the calcareous ring was entirely dif-
ferent—with long tails—and the species must therefore be regarded as
new.
Genus 4. APENTAMERA, new genus
Diagnosis ——Small forms (few cm. long) with tube feet in 5 bands,
papilliform toward the ends where 5 valves are formed; a few smaller
papilliform tube feet are scattered in the dorsal interambulacra, may be
lacking in small individuals. Tentacles 10, the 2 ventral smaller. Skin
rigid, packed with spicules. Calcareous ring with long posterior prolon-
gations; other internal features almost as in Pentamera.
Spicules an external layer of flattened baskets almost rosettelike,
easily overlooked and may possibly be lost in older individuals, and an
inner layer of regular 4-holed knobbed buttons. Feet with end plate
smaller in the terminal appendages; walls packed with curved supporting
tables with 2 pillars and a few blunt teeth. Introvert with oblong tables
or plates with slightly knobbed edge. Tentacles packed with plates and
rods which gradually decrease in size in the branches; apparently no
rosettes.
Type species.—A pentamera lepra, new species.
Remarks.—The material of the type species was originally referred
to 2 genera related respectively to Pentamera and Pentacta, possibly with
affinities to some of the Thyone-like forms. After all other species had
been classified, it became evident that these two “genera” represented
only one, which in spite of its affinities to the T’hyone-like forms could
not be placed among these.
92 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
Apentamera lepra, new species
Plate 13, Figs. 1-10
Diagnosis —As for the genus. Color white, mottled with reddish
brown.
Type.—Holotype, AHF no. 25.
Type locality —Station 557-36, off White Rock, Isla Partida, Gulf
of Calif., in 45 fms., March 8, 1936.
Distribution.—Gulf of Calif. to Panama.
Depth.—From 30-50 fms.
Specimens examined.—The following material from the Hancock
Expeditions :
557-36. Isla Partida, Gulf of California, 45 fms., type and 2 paratypes.
863-38. Bahia Honda, Panama, off North Island, 30-50 fms., March
1, 1938, 3 specimens (M.C.Z.).
Remarks.—The type specimens measure 3.5, 3.4, and 2.0 cm. in
length; the gonads are well developed in the 3 individuals, so presumably
the animals are full grown. The material has been carefully compared
with the other forms with 4-holed knobbed buttons, but it seems impos-
sible that it represents the juvenile stages of any of these. The only form
which possibly could be considered is Neothyone gibbosa (see p. 113),
but the latter has numerous interambulacral feet when it is as large as
A. lepra, and its spicules are definitely larger; the buttons are more ob-
long and it has numerous rosettes in the tentacles.
Genus 5. LEPTOPENTACTA H. L. Clark, 1938
Synonym.—Ocnus Auctores. Nec Ocnus Forbes, 1841. See H. L. Clark,
1938, p. 453.
Diagnosis —Body slender, skin rigid. Tentacles 10 (or 8), the ven-
tral ones small (or lacking). Feet restricted to the ambulacra, rigid, filled
with spicules, arranged either in single rows with feet well spaced or in
more crowded or zigzaggy rows. Toward the oral and anal ends the feet
are scattered and papilliform, often forming 5 valves as in Pentacta. Cal-
careous ring simple, or with short posterior prolongations. Retractors
short, longitudinal musculature feebly developed. Gonads with few tubes
in each tuft, attached near the middle of the animal.
Spicules reticulated bodies or baskets, larger or smaller, reticulated
scales or grains and 4-holed buttons, swollen or knobbed. Feet with or
without end plate, with or without supporting rods. Tentacles with large
perforated rods or plates; sometimes also rosettes and smaller rods.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA 93
Type species.—L. grisea H. L. Clark, 1938.
Remarks.—The genus may possibly be further divided into two, viz.,
Leptopentacta, which includes the type species, L. panamica spec.
nov., possibly also Sluiter’s ““Ocnus” javanicus, and a species listed here as
L. nina, based on a juvenile individual from the Hancock Expeditions,—
and another, for which the name Parocnus is proposed,—to accommodate
“Ocnus” imbricatus Semper and “Ocnus” typicus Théel, which possibly
are identical, and L. nova, spec. nov.
Ce Au
1 3
Text fig. 2. 1. Calcareous ring from Lefptopentacta grisea H. L.
Clark.
2. Leptopentacta panamica, new species.
3. Leptopentacta javanica (Sluiter).
Scale, 1 mm.; Sluiter’s figure has no indication of
what magnification is used; probably it is about as
large as the other two.
“Ocnus” molpadioides Semper definitely does not belong in this genus,
while “Ocnus” pygmaeus Semper, based upon a specimen 1 cm. long, has
long and rather flexible feet and probably represents a juvenile specimen
of some species of Cucumaria or Thyone (in the broad sense). It is not
possible to ascertain whether the calcareous ring is simple or has pos-
terior prolongations. Semper figures a simple ring but mentions that the
ring has long posterior prolongations. His figures of the ring of imbricata
show no posterior prolongations, and in his description he emphasizes the
way in which the ring is cut off posteriorly. (See footnote, p. 94.)
Key TO THE KNown Speciés or Leptopentacta
1. External layer of spicules consisting of small biscuit-shaped
plates or rosettes. Feet with no supporting rods, and apparently
without end plate. Calcareous ring simple (not known in L.
DUPICA a hie lO be BN eS: shah RL ele ey ee ee
94
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
External layer of spicules consisting of baskets, reticulated
bodies, or hollow plates. Feet with supporting rods or plates,
in some species also with end plate. Calcareous ring with
Shontstailswatle gochey’ tls! ace ‘Ves 'S aii Gieti-sea baie penne
External spicules consisting of small biscuit-shaped bodies with
4-5 holes, often one side prolonged into a sharp tooth. Feet
few, scattered, 5-6 in the dorsal ambulacra, 8-10 in the ven-
tralones. . . . . . 1. Leptopentacta nova, new species
External spicules consisting of rosettes. Feet fairly numerous,
although arranged in single rows. . . arte ‘
Feet 24-26 in the ambulacra (in individuals 35- 40 mm. ae):
2 6 ew ew ew ew eh). 6Leptopentacta imbricata (Semper)
Feet 15-20 in the ambulacra (in individuals 40 mm. long).
i < s « « os » « « » Leptopentacta typica (Wheel)
External spicules consisting of simple, 4-spoked, deep baskets
with knobbed to dentate rim. Feet with numerous supporting
TOUS. GEplatese! wy. s- as. fae ey Eély Stale hat mae
External spicules consisting of sees reticulated bodies
or concave plates. . . . ee, ae eee
Buttons swollen or aaa deena Baskets comparatively
large and delicate, with 4 marginal holes. . . .
oe eal oS 5 we Leptepentacta grisea FH. ie “Clark
Buttons predominantly strongly knobbed and regular. Bas-
kets small, not delicate, and apparently never with marginal
holes. . . . . . 2. Leptopentacta panamica, new species
External spicules reticulated bodies?; feet 20-23 in ventral
rows (in individual, 40-45 mm. long). oh Siaierees ne
Mee eee es ge te are javanica (States)
External spicules large concave plates with a varying amount
of reticulated bars. Inner layer knobbed buttons slightly ir-
regular and large reticulated plate. Feet with oblong support-
ing plates smooth or knobbed, with a varying number of holes.
3. Leptopentacta nina, new species
3 Sluiter’s figure (1880, pl. 4, fig. 10) looks suspiciously as if it belonged to
a Paracaudina, His description is not very clear, and in 1901, p. 79, he writes
“javanica with typica and imbricata,” following Ludwig, 1887, p. 1221 (reprint
p. 5). If one can trust Semper’s figure (plate 14, fig. 12), imbricata must be
different from javanica, as it has a simple calcareous ring. Although Ludwig's
“imbricata” has distinct tails on the ring, it is not proved that it is identical with
Semper’s species or with Théel’s.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 95
1. Leptopentacta nova, new species
Plate 14, Figs. 13-22; text figure 3
Diagnosis.—Body slender, slightly curved with rigid body wall. Feet
few, contracted into conical protuberances, well spaced. The ventral am-
bulacra possess from 8-10 feet, while the dorsal ones each have 4 feet
anteriorly and one or two posteriorly. Skin with large cobblestone-like
scales, and the space between them filled with masses of smaller spicules.
Tentacles fingerlike (in the type which measures 2 cm. in length) and
apparently only 8 are present, but the 2 small ones may have been over-
looked. Calcareous ring low, with no indication of posterior prolonga-
tions. Inner anatomy not remarkable in any respect.
Spicules large scales or grains with reticulated structure, 4-holed but-
tons, irregular, swollen or with irregular knobs, and minute perforated
biscuit-shaped plates with 4-5 holes and frequently a long spine developed
on one side. Feet with no supporting plates and apparently no end plate.
Introvert with reticulated plates which intergrade into rosettes; tenta-
cles with curved perforated or branching rods which decrease in size
toward the ends.
Type.—Holotype, AHF no. 26, 1 paratype.
Type locality—Station 281-34, Santa Maria Bay, west of Hughes
Point, L. Calif., 35-40 fms.
MALY
Text fig. 3. Leptopentacta nova, new species.
Scale, 1 mm.
96 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Distribution —Known from the type locality.
Depth.—Dredged at 35-40 fms.
Specimens examined.—The type.
Remarks.—The type and other specimen measure 2 cm. in length
and 3 mm. in width. The surface of the skin appears mottled, because of
the large scales’ being glassy and contrasting with the solid white of the
smaller spicules which are massed between them. The tentacles which
are completely retracted are small and finger shaped; only 8 could be
counted, but 2 small ones may possibly have been overlooked. The cal-
careous ring is low with no indication of posterior prolongations; the
stone canal is small and attached as usual in the dorsal mesentery, and
the Polian vesicle is single and ventrally placed. Other anatomical fea-
tures are not remarkable. The gonads form few tubes of varying length,
and are attached near the middle of the dorsal midline.
This species seems most closely related to L. imbricata (Semper) and
L. typica (Théel). It differs in having small biscuit-shaped plates in-
stead of rosettes in the external layer of the skin, fewer feet, and, presum-
ably, in having simple tentacles. However, L. nova, L. imbricata (and L.
typica) may well be placed in a separate genus for which the name
Parocnus is proposed, with imbricata as type.
2. Leptopentacta panamica, new species
Plate 14, Figs. 6-12; text figure 2
Diagnosis —Slender form with curved body, ambulacra forming 5
indistinct ridges. Feet in scattered double rows in the middle portion of
the ambulacra, toward the ends in single rows, around the base of the
introvert forming 5 large valves as in Pentacta. Tentacles 10, bushy,
the 2 ventral smaller. Surface of skin rough on account of the numerous
spinelike projections on the spicules which pierce the skin. Calcareous
ring with short posterior prolongations. Other features also typical of
the genus.
Spicules consisting of an external layer of strongly knobbed 4-holed
buttons with minute holes and smaller buttons, less strongly knobbed and
with larger holes and numerous 4-spoked baskets, and an inner layer of
large, reticulated ovoid or spherical scales or grains. Feet possibly with an
end plate (not discovered in type)* and numerous perforated supporting
4 The type species has a large and well-developed end plate in the feet (over-
looked by H. L. Clark), and one would expect that an end plate would also be
present in the present species. Two tube feet have been examined, but the feet
are strongly contracted and no trace has been found of end plates. Still it is pos-
sible that they may be present.
NO.3 DEICHMANN: HOLOTHURIOIDEA; PARTI,DENDROCHIROTA 97
plates, and the same type of spicules as in the integument. Particularly
numerous in the appendages, although also scattered in the skin are some
peculiar oblong heavy plates with small holes and one end drawn out
into a shorter or longer spine which pierces the skin (visible with low
magnification). Introvert with rosettes, tentacles with large curved per-
forated plates and rods, also rosettes, and in the finer branches, delicate
rods with perforated ends.
Type.—Holotype, AHF no. 27.
Type locality—Port Utria, Colombia, close to shore, north of point
and west of islands, 20 fms., January 25, 1935.
Distribution —From Port Utria, Colombia, to Tenacatita Bay, Mex.
Depth—From 20-35 fms.
Specimens examined.—The type and 3 other specimens from the fol-
lowing Velero stations:
259-34. Tangola Tangola, Mex., 15-20 fms., February 28, 1934, 1
paratype.
275-34. Tenacatita Bay, Mex., off Navidad Head, 25-35 fms., 2 speci-
mens.
423-35. Port Utria, Colombia, close to shore, 20 fms., 1 specimen, type.
Remarks.—The type measures about 4 cm. in length; the other indi-
viduals are smaller. The color in alcohol is dirty yellow with large ir-
regular brown patches. The gonads are ripe and filled with eggs. (Speci-
mens collected in January. )
The species seems fairly closely related to the type species, L. grisea
H. L. Clark, from Broome, West Australia (plate 14, figures 1-5). It
differs chiefly in the presence of the peculiar, spine-bearing plates, the
strongly knobbed buttons, and the much smaller and more simple baskets.
3. Leptopentacta nina, new species
Plate 15, Figs. 1-12
Diagnosis—Small form (?), type measures 1.2 cm. with the tenta-
cles contracted. Body curved, slender; skin rigid; feet cylindrical, re-
tractile, restricted to the ambulacra (the type has about 25 tube feet in
the ventral rows and considerably fewer in the dorsal). Calcareous ring
with distinct posterior tails on the radials and long teeth, deeply grooved
on radials and interradials (resembles that of Sluiter’s species, see text
figure 2).
98 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Spicules an external layer of concave plates, often with a delicate
reticulum across the hollow surface; an inner layer of knobbed, irregular
buttons, and larger plates with knobbed surface, sometimes transformed
into large convex bodies. Feet apparently devoid of end plate; walls with
short, broad, perforated supporting rods. Introvert and tentacles not
examined. Color of the preserved specimens white.
Type.—Holotype, AHF no. 28.
Type locality—Station 850-38, Cape San Francisco, Ecuador, 15
fms., February 23, 1938.
Distribution Known only from the type locality.
Depth—From 15 fms.
Specimens.—The type.
Remarks.—The type represents undoubtedly an immature individual.
On account of its small size and its rigid skin it was almost impossible to
dissect, and no attempt was made to study the tentacles and introvert.
The inner organs were poorly preserved, and nothing can be said about
the stone canal and Polian vesicle. The musculature is feebly developed,
and the intestine and respiratory trees form a structureless mass; a well-
developed muscle stomach could, however, be seen. The calcareous ring
is highly reminiscent of the one figured by Sluiter for javanica (see text
figure 2), and it is of course remotely possible that the present species has
previously been described from the East Indies. As the various earlier
descriptions are rather unsatisfactory, as also the figures hitherto pub-
lished, it has been deemed wiser to designate the unique individual to a
new species.
L. nina differs distinctly from both L. nova and L. panamensis in its
calcareous ring as well as in its spicules. It cannot be confused with any
other species at present described from the Panamic region.
Genus 6. PENTACTA Goldfuss, 1820
Pentacta Goldfuss, 1820, p. 177.—H. L. Clark, 1923, p. 416.
Colochirus ‘Troschel, 1846, p. 64. (Type species—C. quadrangularis
‘Troschel. )
Cercodemas Selenka, 1867, p. 343. (Type species.—C. anceps Selenka.)
Diagnosis—Body with more or less flattened ventral side with the
tube feet arranged in 3 bands; dorsal side vaulted with feet of different
size, often as large papillae, frequently scattered in the interambulacra.
Around the base of the introvert and the anus the ambulacra form thick
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA 99
valves. Tentacles 10, the 2 ventral smaller. Skin rigid, filled with spicules.
Calcareous ring simple, posteriorly often strongly undulated but never
with posterior prolongations.
Spicules consisting of an external layer of either baskets or delicate
reticulated bodies, and inner layer of knobbed buttons, heavy plates, or
reticulated bodies. Feet apparently with no end plate; walls supported
by perforated rods or plates; dorsal papillae with mostly curved sup-
porting plates. Tropical shallow water forms.
Type species —Pentacta doliolum (Pallas).
Remarks.—The name has been used by various authors for members
of the genus Cucumaria s.l., and Ludwig (1899, p. 344) claims that it
ought to be used instead of Blainville’s name, from 1835. H. L. Clark
(1923, p. 416) has, however, called attention to the fact that Troschel’s
name Colochirus is a complete synonym of Pentacta, for Goldfuss desig-
nated Pallas’ doliolum from Cape of Good Hope as the type species and
the latter is generally accepted as the typical “Colochirus.”
The genus Pentacta is most abundantly represented in shallow water
in the East Indies. The type species represents the only form known from
the South African waters, and only one form is known from Brazil and
the West Indies. From the western coast of South America Semper de-
scribed a new species which was never found again and hence was re-
garded as rather dubious. There is no doubt that it is the one which the
Allan Hancock Expeditions now have brought back, thereby proving
that Semper’s specimens were correctly labeled.
The genus may possibly in the future be divided into 2, viz., one for
those with more complicated spicules, and one for those with regular,
knobbed buttons and simple baskets.
The 2 species known from the American waters both belong to the
latter group.
Pentacta peruana (Semper)
Plate 16, Figs. 1-8
Colochirus peruanus Semper, 1868, p. 239, pl. 39, fig. 20.—Lampert,
1885, p. 126.—Théel, 1886, p. 123.—H. L. Clark, 1910, p. 353.
Diagnosis—Small form, few cm. long, with cylindrical feet in 3
crowded bands on the greater part of the ventral side, transformed into
papillae toward the ends, dorsally 2 bands of papillae and a number
scattered in the interambulacra. Oral and anal ends closed by 5 distinct
valves. Inner anatomy typical of the genus. “a Pa
| Sete
wt £08 iy aie
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(LIERARY] =
100 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Spicules an external layer of regular baskets, the hollow part formed
by broad bars, the narrow rim with few teeth. An inner layer of regular
4-holed knobbed buttons and larger knobbed plates and huge reticulated
grains. Ventral feet mostly with a minute end plate and the walls packed
with supporting rods mostly 3-armed, smooth to knobbed. Dorsal ap-
pendages apparently lacking end plate but with heavier 3-armed rods.
Introvert with a few oblong buttons smooth or knobbed. Tentacles with
stems packed with narrow plates or rods, smaller, curved and often 3-
armed rods in the finer branches.
Type.—Possibly in Germany.
Type locality.—Peru, probably at comparatively low depth.
Distribution From Peru to Mexico.
Depth.—The Hancock material came from 25 fms.
Specimens examined.—The following specimens from the Allan Han-
cock Expeditions:
264-34. Petatlan Bay, Mexico, 25 fms., south and west of White
Friars Island, rock with gorgonids, 4 specimens.
Remarks.—The 4 specimens measure between 3-4 cm. in length and
are thus somewhat larger than the types which measured only 15-18 mm.
Semper’s description is not very detailed,®> and his figure of a basket
shows a more irregular type than the one which commonly occurs. He
mentions hollow knobbed bodies and large smooth plates, the latter re-
ferring possibly to the smooth supporting plates. He gives, however, no
reference to the regular 4-holed knobbed buttons, possibly because he
considers them growth stages of the larger knobbed bodies; also they
may possibly be more scarce in younger individuals. It must further be
mentioned that the ventral tentacles in the present material are very
slightly smaller in size than the other.
5 Semper’s original description: “Zehn Tentaklen, die zwei ventralen kleine.
Habitus Ascidien-artig. Auf den Riicken stehen die Ambulacral-papillen regellos
zerstreut, am Bauche in drei deutlichen Reihen, deren jede zwei Fiisschen in der
Breite zeigt. Es bleibt sich eine Art Bauchscheibe dadurch das die Fiisschen
hinten in Ambulacral-papillen tibergehen. Farbe gelblich (im Spiritus). Drei
Exsemplare, 15-18 mm. lang, 6-7 mm. breit.
Am Gefassring eine einzige Polische Blase, eine kleine dorsaler Steinkanal.
Lungen fast so lang wie der K6rper, mit wenig, aber sehr langen einfachen
Nebenasten. Geschlechtsteile zwei Biischel sehr kleiner unverdstelter Schlauche,
ihre Basis am Mesenterium sehr weit nach vorne, vor dem Wassergefassring.
Kalkring aus 10 einfachen Gliedern bestechend; die Radialen etwas breiter als
die Interradialen, nicht nach hinten verlanget. Keine Kaumagen. Die Retraktoren
inserieren sich in der Mitte des K6rpers.
Grosse glatte Kalkplatten in der ganz starre Haut, dariiber ein Schicht
durchbrochenene, etwas knotiger Halbkugeln.”
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 101
One specimen, which was opened, had few but well-developed genital
tubes. The color of the 4 individuals is light brown with darker freckles;
the general impression of the preserved material is a nondescript grayish-
brown color.
Genus 7. THYONACTA, new genus
. Diagnosis—Medium-sized to large form, 10-15 cm. long in con-
tracted condition. Skin thick, filled with spicules. Tube feet in 5 bands
and in varying number in the interambulacra, depending on the animal’s
age; toward the oral and anal ends thickened valves are developed. Many
tube feet are conical, papilliform. Tentacles 10, the 2 ventral smaller.
Calcareous ring low but with short posterior projections; stone canal
small, attached to the dorsal mesentery; Polian vesicle single, ventral in
position; retractors exceptionally short, attached closely behind the cal-
careous ring (when the crown of tentacles is withdrawn). Gonads
placed unusually far back, in the posterior third of the body.
Spicules consist of an external layer of simple baskets, and an inner
layer of 4-holed buttons of different size, also a few larger plates; mostly
regularly knobbed. Feet apparently without any end plate (at least in
larger individuals) ; walls with numerous supporting rods and plates
besides buttons and baskets. Introvert with rosettes; tentacles with large,
mostly narrow perforated plates and rods which decrease in size toward
the tips of the branches.
Type species —Thyonacta sabanillensis (Deichmann).
Remarks.—At present 2 species have been referred to this genus, both
restricted to the American waters. Very likely some of the members of
the old genus Thyone from the western Pacific and Indian oceans ought
to be transferred to this genus. The type species came from the West
Indian seas, while the other species, described below, has been secured
from various localities in the Panamic region.
Thyonacta mexicana, new species
Plate 17, Figs. 1-13
Diagnosis—As for the genus. Spicules rather stout and deep baskets
and buttons of 2 types, a large and strongly knobbed one and a smaller,
less regularly knobbed and often with accessory holes. Color varying
shades of brown.
102 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Type.—Holotype, AHF no. 29, 5 paratypes.
Type locality—Station 633-37, San Gabriel Bay, Espiritu Santo
Island, L. Calif., 18 fms., March 6, 1937.
Distribution.—Widespread in the Gulf of California and as far as
Panama.
Depth.—F rom 10-35 fms.
Specimens examined.—The type and 5 specimens from the following
Velero stations:
244-34. Bahia Honda, Panama, 30-35 fms., February 21, 1934, 1 speci-
men.
554-36. Parallel to Angel de la Guardia Island, east side, 10 fms.,
March 8, 1936, 2 specimens.
562-36. North of San Esteban Island, L. Calif., 20-70 fms., March 10,
1936, 1 specimen.
633-37. San Gabriel Bay, Espiritu Santo Island, L. Calif., 18 fms.,
March 6, 1937, 1 specimen.
719-37. Consag Rock, L. Calif., 10-25 fms., March 24, 1937, 1 speci-
men.
Remarks.—The material shows to perfection how impossible it is to
give an exact description of the general appearance of most holothurians.
Two individuals are well expanded with tapering bodies, one is con-
tracted to a short Pentacta-like form, and two resemble slightly deformed
oranges. The spicules, however, are completely identical, and a closer
study of the anatomical features, calcareous ring, etc., showed that the
specimens also in these respects are alike.
T. mexicana is closely related to the type species but differs distinctly
in the shape of the spicules. ‘The extremely deep baskets and the large
strongly knobbed buttons alone are sufficient to distinguish the present
species from all other related forms known from the Panamic region.
Genus 8. ,THYONE Oken, 1815
Thyone Oken, 1815, p. 351.
Anaperus Troschel, 1846, p. 60 (partim).
Diagnosis—Small to medium-sized forms (rarely more than 10 cm.
long, many species smaller). Body thin skinned with numerous nonre-
tractile fairly delicate feet, most numerous ventrally. Tentacles 10, the
2 ventral smaller. Calcareous ring with long posterior prolongations on
the radials.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART 1, DENDROCHIROTA 103
Spicules small 2-pillared tables or derivatives of these. Feet with
large end plate and numerous curved supporting tables usually with a
well-developed spire, which in some forms, however, is completely re-
duced. Tentacles with delicate rods and often rosettes. Spicules gradually
reduced with age in certain species.
Type species—Thyone fusus (O. F. Miller).
Remarks.—The genus is well defined and corresponds to Pentamera
in Cucumaria sensu latiore. It is sometimes almost impossible to identify
very young specimens of Pentamera and Thyone, as the latter passes
through a Pentamera-stage. Fortunately that stage seems to be so short
that one usually is able to find a few feet in the interambulacra. When
dealing with specimens less than 1 cm. long, it is, however, wise to con-
sider all the species of Pentamera and Thyone which are known in the
region under consideration. Often the spicules in the introvert and ten-
tacles differ in species which have spicules in the body wall which are
alike, but this character cannot always be relied upon.
The type species is common in the eastern and northeastern part of
the Atlantic Ocean, while a closely related species is known from the
West Indies, but not reaching the coasts of New England (T. fusus ?
Deichmann, 1930, p. 167). Another species with more robust tables,
often with a handle on the inner side of the buttons, is likewise known
from the West Indies (JT. pseudofusus Deichmann, 1930, p. 168).
From the Panamic region the Hancock Expeditions have secured 3
new species, while a fourth was in the M.C.Z., collected by the Stranger.
Of these 4 species, 2 are closely related to the West Indian species men-
tioned above.
Key TO THE SPECIES OF Thyone s. str. KNOWN FROM THE
PANAMIC REGION
1. Tables delicate with 4 to 8 holes; disk oval or squarish;
spire of medium height usually with 6 diverging teeth. Sup-
porting tables with spire completely reduced ; disk curved, with
4 central holes and ends expanded with a varying number of
minute holes. . . . . 1. Thyone neofusus, new species
1. ‘Tables more or less stout; supporting tables in tube feet with
MECIEEVGIODERD SPIE Aiece id cd) ‘isin coy cates eeews a) Deel ime aes
2. Disk of table medium-sized to large, round, squarish, or oblong,
with numerous holes and often with a few knobs or the entire
104 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
margin distinctly thickened ; spire in most cases low, in process
of becoming vestigial; supporting tables with an enormous
conical spire. . . . . . 4. Dhyone strangeri, new species
2. Disk of tables small, oval or squarish in outline, with 4-8
holes, in some cases the disk is lozenge shaped with a few small
holes in the ends;"no knobs. °. 2°.) 2... ee
3. Disk oval or lozenge shaped ; spire moderately well sie
mostly with few, indistinct teeth. Supporting tables with 2
pillars which unite into a tapering cone ending in 2 small di-
verping: teeth (often broken off). <<. °. . 2 . 2.7.
2. Thyone bidentata, new species
3: Disk ay ih 4 central holes and a “handle” on the inner
side; spire tapering with 3-4 small teeth at the top; support-
ing tables with 2-3 pillars in the spire which ends in 2-3 teeth.
oe ew ew wl ew ew he 68. Lhyone parafusus, new species
1. Thyone neofusus, new species
Plate 18, Figs. 1-6
Diagnosis—Small form (few cm. long?) ; feet delicate in 5 indis-
tinct bands and scattered in the interambulacra. Calcareous ring typical;
single small stone canal and single ventral Polian vesicle.
Spicules delicate tables with 4-8 holes and 2-pillared spire. Feet with
large end plate and numerous curved supporting plates or rods with 4
central holes and a varying number in the expanded ends and no trace
of a spire. Introvert with rosettes; tentacles with delicate rods and ro-
settes. Color of the preserved specimens white.
Type.—Holotype, AHF no. 30, 2 paratypes.
Type locality—Station 190-34, Lat. 0° 55’ S.; Long. 90° 30’ W.;
58-60 fms.
Distribution —Off the Galapagos and vicinity.
Depth—From 20-60 fms.
Specimens examined.—The type and the following specimen from the
Velero Expeditions:
190-34. Lat. 0° 55’ S.; Long. 90° 30’ W.; 58-60 fms., January 26,
1934, 1 specimen, type.
814-38. North of Hood Island, Galapagos, 20-40 fms., January 28,
1938, 2 specimens.
Remarks.—The type measures about 2 cm. with the tentacles with-
drawn. The gonads form 2 well-developed tufts; so the animal may pos-
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 105
sibly have reached its full size. Other members of the genus, as for ex-
ample Thyone fusus, reach a considerably greater length.
Thyone neofusus resembles Thyone fusus? from the West Indies
(Deichmann, 1930, p. 167) but differs in the complete lack of a spire
on the supporting plates; this may of course be a variable character. For
the present it seems natural to segregate the various forms of fusus until
different material has been gathered to permit a critical survey. So far
no closely related species have been reported from the Californian and
Chilean waters or from Hawaii.
2. Thyone bidentata, new species
Plate 18, Figs. 13-16; text figure 4
Diagnosis —Small form (few cm. long) with delicate to fairly stout
feet, often in distinct bands and also scattered in the interambulacra.
Inner anatomy typical of the genus. Spicules tables with oval to lozenge-
Text fig. 4. Thyone bidentata, new species, juv.
1-3. Tables from integument.
4. Supporting table from tube foot.
Scale, 4409 mm.
shaped disk with 4 central holes (and a few holes in the ends when the
disk is lengthened) ; spire tapering to a cone with few teeth. Supporting
tables with spire composed of 2 rods united into a slender spine, ending
in 2 diverging teeth. Introvert with tables with numerous holes and low
spire with diverging teeth; tentacles with a few rosettes. Color of pre-
served specimens pale brown.
106 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Type.—Holotype, AHF no. 31.
Type locality.—Station 275-34, Tenacatita Bay, Mex., dredging in
25-35 fms., March 4, 1934.
Distribution —From Gulf of Calif. to Colombia.
Depth.—F rom 12-30 fms.
Specimens examined.—The type and the following specimens from
the Velero expeditions:
245-34. Bahia Honda, Panama, dredging 15-25 fms., February 21,
1934, 1 specimen.
249-34. Bahia Honda, Panama, 15-20 fms., February 22, 1934, 1 speci-
men.
251-34. Secas Islands, Panama, dredging in 15 fms., February 22, 1934,
1 specimen.
275-34. Tenacatita Bay, Mex., off Navidad, 25-35 fms., March 4,
1934, 3 specimens, type and paratypes.
423-35. Port Utria, Colombia, close to shore, 20 fms., January 25,
1935, 1 specimen.
679-37. Outside of Concepcion Bay, Gulf of Calif., 30 fms., March
15, 1937, 1 specimen.
683-37. Outside of Concepcion Bay, Gulf of Calif., 12 fms., March
15, 1937, 1 specimen.
686-37. Concepcion Bay, Gulf of Calif., 12 fms., March 16, 1937, 1
specimen.
Remarks—The smallest paratype measures 0.5 cm. in length; the
type and other larger specimens measure about 2 cm. In the smallest
specimens the spicules are considerably smaller than in the larger as is so
often the case, and many of the tables have a squarish disk with 4 acces-
sory holes. In some of the larger individuals many tables are lozenge
shaped with small accessory holes in the ends, while other individuals
of the same size have almost exclusively oval disks with 4 holes.
The exterior of the animals shows considerable variation in regard
to the size of the tube feet, which vary from delicate to fairly stout.
3. Thyone parafusus, new species
Plate 18, Figs. 7-12
Diagnosis.—Small form (few cm. long) with fairly stout feet, more
or less distinctly arranged in bands and scattered in the interambulacra.
Inner anatomy typical of the genus. Spicules form a crowded layer of
NO. 3. DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 107
tables with oval disk with 4 holes and a well-developed handle on the
inner side; spire tapering, ending in a few teeth. Feet with well-devel-
oped end plate and numerous supporting tables normally with 3 rods in
the spire, which rapidly tapers into a short spine with a few teeth on the
tip. Introvert with oblong tables with a varying number of holes. Ten-
tacles packed with perforated plates of different sizes, with numerous
holes; rosettes and delicate rods are present in the terminal branches.
Type.—Holotype, AHF no. 32, 1 paratype.
Type locality—Station 272-34, Tenacatita Bay, Mex., 25 fms.,
March 4, 1934.
Distribution—Tenacatita Bay, Mex.
Depth—From 25 to 35 fms.
Specimens examined.—The type and one paratype from the Velero
Expeditions:
272-34. Tenacatita Bay, Mex., 25 fms., dredging off Navidad Head,
March 4, 1934, 1 specimen.
275-34. Tenacatita Bay, Mex., 25-35 fms., March 4, 1934, 1 specimen,
type.
Remarks.—The type measures about 2 cm. with the oral end ex-
panded. Several tentacles are torn off, and most of the inner organs
are ejected.
The spicules resemble closely those found in T. pseudofusus Deich-
mann (1930, p. 168) from Yucatan, 25 fms. depth. The spicules differ
in various respects; the West Indian form has tables with a short robust
spire with numerous teeth and lacks apparently the numerous perforated
plates in the tentacles.
4. Thyone strangeri, new species
Plate 19, Figs. 1-11
Diagnosis—Medium-sized form; numerous delicate feet scattered
all over the body wall, often with conical or wartlike base. Calcareous
ring with long posterior prolongations; interradials broad, more or less
heart shaped, united by sutures with the radials. Stone canal small with
a small head with thickened edges coiled up like a ball; Polian vesicle
single.
Spicules a crowded layer of oblong tables, often with thickened edge
and a varying number of knobs; number of holes in disk variable; spire
small, on the verge of being reduced; teeth almost completely lacking on_
=
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108 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the tip. Feet with large end plate and numerous supporting tables with
a conical spire. Introvert with oblong tables, disk with knobbed edge
and low spire. Tentacles with rosettes and delicate rods. Color brown,
tentacles dark, blackish.
Type.—M.C.Z.
Type locality—lIsle Grande, west coast of Mexico (Lat. 170° 46’
N.; Long. 101° 42’ W.).
Distribution —Known from the type locality.
Depth—From 7 fms.
Specimens examined.—The type.
Remarks.—The type measures about 4 cm. with the tentacles with-
drawn. The anterior portion is well preserved, but most of the intestine
and respiratory trees are eviscerated. The gonads form well-developed
tufts near the middle of the dorsal midline.
The spicules are extremely characteristic and different from those
found in all other species known from the Panamic region as well as the
adjacent waters. Superficially they may be mistaken for those of Phylloph-
orus aculeatus Ludwig, but comparison of the figures (plate 25) shows
that they are entirely different.
Genus 9. NEOTHYONE, new genus
Stolus Selenka (partim), 1867, p. 356.
Diagnosis —Medium-sized forms with numerous feet; skin thick,
packed with spicules. Tentacles 10, the 2 ventral smaller. Calcareous
ring with posterior prolongations on the radials, in some species fairly
short and soft, often inward curled, so that they easily escape notice.
Single stone canal, one or more Polian vesicles, ventrally placed.
Spicules an external layer of specially modified buttons, with spines
either on the outer handle or transformed into reticulated baskets, in
some cases lost with advancing age. An inner layer consisting of numerous
knobbed buttons, mostly with 4 holes; knobs distinct or almost obliter-
ated. Feet with large end plate and numerous supporting tables with or
without a spire. Introvert with a varying amount of tables or reduced
buttons; tentacles with a varying number of heavy plates or rods, some-
times reduced with age. Rosettes seem to be present in certain species, at
least in the younger individuals; possibly they represent a variable char-
acter.
Type species—Neothyone gibber (Selenka).
NO. 3 DEICHMANN: HOLOTHURIOIDEA}; PARTI, DENDROCHIROTA 109
Remarks.—The genus has been established to accommodate some of
the species which have numerous buttons and distinct posterior prolonga-
tions on the radials. For the present 2 species from the West Indian
waters and 3 from the Panamic region are placed in the genus. Of these
latter, one may possibly deserve a genus of its own. No attempt has been
made to discuss the various East Indian species with similar spicules that
possibly belong in the genus.
Key TO THE SPEcIES OF Neothyone KNOWN FROM THE
PANAMIC WATERS
1. Spicules large buttons, faintly knobbed; external layer of but-
tons with strongly spinous handle. Feet with stout supporting
tables with well-developed spire.e . . . . 2... ee
ele) ew elles 2 ew 1S Neothyone gibber: (Selenka)
1. Spicules smaller or larger buttons strongly knobbed; external
layer of buttons with few spines on the handle (often lost in
older individuals) or transformed into baskets (the button
itself is almost smooth with long delicate projections on the ex-
ternal side uniting into a delicate reticulum). . . . . . 2
2. Feet with large smooth supporting rods with no trace of spire
(except in some of the dorsal appendages). External layer of
smooth buttons modified into baskets with the external side
covered by a delicate reticulum; these deposits are apparently
retained throughout the animal’s life. . . . . ....
2 2 « « © the ht 62 Neothyone panamensis (Ludwig)
2. Feet with curved, stout supporting tables with a varying
amount of spire; spire often large and deformed in dorsal ap-
pendages; older individuals may lack spire completely, but
often one side is expanded and lobate. External layer of buttons
often with a few spines on the handle but this type is often rare
or lacking in older individuals; often a few concave buttons
with marginal spines form a simple type of baskets. . . . .
oe ew ew ew hw hel elle «(C3 Neothyone gibbosa, new species
1. Neothyone gibber (Selenka)
Plate 20, Figs. 1-9; text figure 5
Stolus gibber Selenka, 1867, p. 356.
Thyone gibber Semper, 1868, p. 66.—Théel, 1886, p. 139.—Deichmann,
1936, p. 64 (passim).
110 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Nec Thyone gibber Deichmann, 1938, p. 376, text figure 11 (Neothyone
gibbosa, spec. nov., see below).
Thyone similis Ludwig, 1887, p. 23, pl. 2, fig. 7.
Diagnosis —Medium-sized form, often with body strongly contracted
and the blunt oral and anal ends upward curved. Calcareous ring fairly
delicate with well-developed posterior prolongations on the radials;
interradials narrow. Single dorsal stone canal, usually several Polian
vesicles.
Spicules large +holed buttons varying from regularly knobbed to
swollen; the latter are especially characteristic of the older individuals.
The 2 knobs in the center of the button are often united to a handle on
both sides of the button. The external layer of buttons has normally dis-
tinct teeth developed on the handle of the external side; sometimes ac-
cessory rods from the margin of the button connect with the spines. Feet
with large end plate and numerous stout supporting tables with several
rods in the spire and a varying number of teeth on the top. Introvert with
Text fig. 5. Neothyone gibber (Selenka), juv., still in the “Pen-
tamera”’ stage.
1-2. Buttons.
3. Supporting table.
4. Table from introvert.
5. Plate from tentacle.
6-7. Rosette and rod from tentacle.
Scale, 4o9 mm.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 111
delicate tables with low spire and lacelike disk approaching rosettes in
complexity. Tentacles with perforated plates which often disappear in
the older individuals; besides more delicate rods and plates in the
branches, also rosettes. Color dirty white to almost black; oral end or at
least the tentacles and the introvert are always dark purplish to black in
color.
Type.—M.C.Z.; paratypes in Gottingen.
Type locality—Panama, depth not reported.
Distribution—Known from Acapulco, Mex., to Lobos de Afuera
Island, Peru, often in the same locations as N. panamensis (Ludwig).
Depth.—Tidemark to few fms., clinging to rocks.
Specimens examined.—The type and several specimens in various
collections (from Acapulco, Panama, etc.).
From the Velero Expeditions the following specimens have been ex-
amined :
132-34. Braithwaite Bay, Socorro Island, Mex., 40 fms., January 4,
1934, 5 small specimens.
391-35. Lobos de Afuera Island, Peru, shore of main island with light-
house, rocks, January 17, 1935, 34 specimens.
466-35. Parker Bay, Costa Rica, shore, February 9, 1935, 14 specimens.
473-35. Parker Bay, Costa Rica, rocks and algae, 2 fms., February 9,
1935, 2 specimens.
844-38. Lobos de Afuera Island, Peru, shore, February 14, 1938, 14
specimens.
Remarks.—Selenka’s description is exceedingly brief and not accom-
panied by any figures. A good description and excellent figures are given
by Ludwig, who described the species as Thyone similis, also from
Panama.
Neothyone gibber differs from the other species with knobbed buttons
from the Panamic region in its larger buttons, with strong tendency to
obliteration of the knobs and the presence of numerous spines on the ex-
ternal handle. Deichmann referred erroneously a specimen of Neothyone
gibbosa to this species (see p. 113), but the latter has very distinct knobs
on the buttons, and these are also much smaller as shown by the figures
here given.
The smallest individuals, from Station 132-34, measure less than 0.5
cm., and the feet are restricted to the ambulacra. The spicules are con-
siderably smaller than in the larger individuals.
112 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
2. Neothyone panamensis (Ludwig)
Plate 21, Figs. 1-6
Thyone panamensis Ludwig, 1887, p. 22.—Deichmann, 1936, p. 64
(passim) ; 1938, p. 378 (passim).
Diagnosis —Medium-sized form with numerous cylindrical tube feet
on the ventrum, fewer and most papilliform on the dorsum. Calcareous
ring with relatively broad radials with short posterior prolongations,
often inward curved, so that they are easily overlooked. Other anatomical
features as in the other species.
Spicules an external layer of baskets derived from oval smooth but-
tons, which on their external side develop a number of delicate projections
that unite into a fragile meshwork. Spicules in the inner layer small regu-
lar 4-holed buttons, strongly knobbed. Ventral feet with large end plate
and numerous large, almost flat, smooth supporting plates or rods, ob-
viously derived from supporting tables but with no trace of spire. In the
dorsal appendages the end plate is smaller, and the supporting tables often
show traces of a spire and are more curved. Introvert with delicate but-
tons with various projections on the external side; tentacles normally with
heavy rods or plates with numerous small holes and in the finer branches
smaller, more delicate rods and plates. Apparently no rosettes. Color of
dorsum and tentacles black; ventral side mostly white except near the
oral and anal ends.
Type.—Possibly in Italy.
Type locality —Coast of one of the islands in the Gulf of Panama.
Distribution —Ranging from Panama to Peru, often taken in num-
bers with Neothyone gibber.
Depth—From tidemark to few fms. depth.
Specimens examined.—The following individuals secured by the
Velero Expeditions:
391-35. Lobos de Afuera Island, Peru, shore of main island with light-
house, rocks, January 17, 1935, 2 specimens.
540-36. Puerto Refugio, Angel de la Guardia, Gulf of Calif., shore,
March 3, 1936, 1 specimen.
844-38. Lobos de Afuera Island, Peru, shore, February 14, 1938, 13
specimens.
Remarks.—The type measured only 23 mm., while the present mate-
rial (which is well expanded) measures up to 6 cm. in length. It is with
some doubt that the material is referred to Ludwig’s species, as he main-
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 113
tains that the calcareous ring lacks posterior prolongations, and he does
not mention the difference in the form of the dorsal and ventral append-
ages and the striking black and white coloration. The calcareous ring
does, however, superficially look as if it lacks posterior prolongations, and
the differences between the dorsal and ventral appendages and the color-
ing may quite well be less pronounced in younger individuals. On the
other hand, the description of the spicules shows no discrepancies, and the
present species is the only “7 hyone’-like form with such spicules as those
described by Ludwig. Also it speaks in favor of the identification that it
is usually found in the same localities as Neothyone gibber.
Neothyone panamensis is easily recognized on account of its color and
the arrangements of its feet. Also the spicules are very striking. The bas-
kets seem to be retained throughout the animal’s life and are numerous in
all preparations, while the large, smooth supporting plates from the ven-
tral feet form another character which is not easily overlooked.
The Velero specimens have large tufts of gonads and appear to be
adult individuals.
3. Neothyone gibbosa, new species
Plate 21, Figs. 7-11; Plate 22, Figs. 1-8
Thyone gibber Deichmann, 1938, p. 376, text figure 11. Nec Thyone
(Stolus gibber Selenka, 1867, p. 356, now Neothyone gibber, see
above).
Diagnosis—Medium-sized form superficially resembling Thyone
gibber in its external and internal anatomy.
Spicules an insignificant layer of shallow knobbed baskets derived
from buttons, easily overlooked, and an inner layer of regular 4-holed
buttons strongly knobbed mostly with handle and some with a few spines
of varying size on the handle. Feet with large end plate, walls with nu-
merous heavy supporting tables, mostly with low spire composed of several
rods, occasionally more well-developed spire may be found; especially in
the dorsal feet in older individuals a number are developed as smooth
plates often with one side expanded, with scalloped edge. Introvert with a
varying number of mostly larger tables with numerous holes in the oblong
disk and a low spinous spire. Tentacles with heavy perforated plates and
rods decreasing in size in the branches; in many cases also rosettes. Color
in alcohol dirty gray or reddish, often darker mottled in the interambu-
lacra; tentacles rather darkly colored.
Type.—Holotype, AHF no. 33.
114 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Type locality.—Station 545-36, Puerto Refugio, Angel de la Guardia
Island, Gulf of Calif.
Distribution From the upper end of the Gulf of Calif. to Mazat-
lan, Mex.
Specimens examined.—The type and the following individuals from
the Velero Expeditions:
465-35. Playa Blanca, Costa Rica, shore, February 8, 1935, 1 specimen.
545-36. In Puerto Refugio, Angel de la Guardia Island, west of rock
spit at river wash, shore, March 4, 1936, type and 4 paratypes.
591-36. Port Escondido, L. Calif., Mex., shore, March 16, 1936, 2
specimens.
844-38. Lobos de Afuera Island, Peru, shore, February 14, 1938, 1
specimen.
Also a number of specimens in the M.C.Z. collection from Mazatlan,
Mex., shore, Situantanejo Bay, Mex., and Costa Rica, shore.
Remarks.—It is with some doubt that the present material has been
referred to a new species, for the spicules are—aside from the absence of
heavy plates covered by a stout reticulum—highly reminiscent of those
found in Pachythyone pseudolugubris, which is described below, and the
latter does actually occur in the same localities as Neothyone gibbosa. The
buttons have, however, more frequently 3 knobs on the external handle
and show little tendency to become elongate as in psewdolugubris where
the 2 central knobs frequently remain separate.
Genus 10. PACHYTHYONE, new genus
Diagnosis —Small forms (few cm. long) with stout feet in 5 bands
and numerous in the interambulacra; dorsally and toward the ends more
papilliform. Ventral tentacles smaller. Calcareous ring with short poste-
rior prolongations. Single stone canal, single Polian vesicle. Gonads form-
ing 2 tufts of few tubes.
Spicules an external layer of oval buttons or plates with the external
side covered by a dense reticulum; an inner layer of oblong 4-holed but-
tons regularly knobbed. Feet with end plate, reduced in the papilliform
feet; walls supported by supporting tables with numerous rods in the
reticulated spire; a few developed as plates. Introvert with oblong tables ;
tentacles with heavy perforated plates which gradually become smaller
and more delicate in the branches.
Type species.—Pachythyone rubra (H. L. Clark).
NO.3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 115
Remarks.—Only 3 species have been referred to the genus; all are
known from the western coast of North and Central America and two
are viviparous.
Key TO THE SPECIES OF Pachythyone
1. Color white ventrum and red dorsal side and ends. Spicules
slightly larger and robust; plates in tentacles mostly knobbed.
Dae ue sie a oe le Pachythyone rubra. (HL. Clark)
1. Color mottled brown. Spicules more delicate; plates in ten-
facles ‘miestly smooth: . . 94 6 6 6°. se el tle
2. Supporting tables with strongly branching spire, even in the
ventral feet. . . . 2. Pachythyone lugubris (Deichmann)
2. Supporting tables with low spire, at least in the ventral feet.
. . . « « « 3. Pachythyone pseudolugubris, new species
1. Pachythyone rubra (H. L. Clark)
Plate 23, Figs. 10-12
Thyone rubra (H. L. Clark), 1901, p. 166; text figures 1-5; 1901a,
p. 494.—Deichmann, 1939, p. 5 (passim); text figure 3.
Diagnosis—Small, robust form, few cm. long, with stout cylindrical
feet in 5 longitudinal bands and scattered in the interambulacra; dorsally
the feet show some tendency to become papilliform. External and in-
ternal anatomy typical of the genus. Spicules an outer layer of oval
plates covered on the external side by a large reticulum; an inner layer
of mostly regular 4-holed buttons with 10 marginal knobs and usually
2 central knobs; often the button is lozenge shaped with an accessory
hole in each end. Feet with large end plate, reduced in some of the
dorsal feet, and numerous large supporting tables with a huge reticu-
lated spire; in some of the dorsal feet it covers almost the entire disk of
the supporting table. Introvert with oblong tables with more or less com-
pletely reduced 2-pillared spire, sometimes with a few teeth. Tentacles
with knobbed plates or rods. Apparently rosettes are not present. Color
bright orange-red dorsum and ends, ventrum pure white. (Color retained
in alcohol.) Viviparous; the body cavity of the female contains often
about half a dozen embryos of different developmental stages.
Type—M.C.Z.
Type locality—Monterey Bay.
116 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Distribution From Monterey Bay to Santa Cruz Islands, off Santa
Barbara County, southern California.
Specimens examined.—The type and numerous individuals, preserved
as well as living, from Monterey to Santa Cruz Islands.
Remarks.—Differs from the other 2 species in the color and the much
heavier spicules.
2. Pachythyone lugubris (Deichmann)
Plate 23, Figs. 7-9; text figure 6
Thyone lugubris Deichmann, 1939, p. 1, text figures 1-2.
Diagnosis.—As for the genus. Color mottled brown. Spicules slightly
smaller and more delicate than in P. rubra; tentacles without knobbed
plates. Rosettes absent or rare. Viviparous.
Holotype-—U.S.N.M.
Type locality—Magdalena Bay, west coast of L. Calif., 10-15 fms.
Distribution Between Magdalena Bay and Cerros Island, west
coast of L. Calif.
Depth.—10-40 fms.
Specimens examined.—The types and the following specimens from
the Velero Expeditions:
283-34. Thurloe Bay, L. Calif., 8-10 fms., off Thurloe Point, March 9,
1934, 5 specimens.
287-34. South Bay, Cerros Island, Mex., 10-15 fms., March 10, 1934,
12 specimens.
Remarks.—The type, a female, contained 13 embryos (collected in
June). In the Hancock material collected earlier in the year no embryos
were discovered.
While the type material appeared to be devoid of rosettes in the ten-
tacles, a few were found in the Hancock material which came from a
much more northern locality.
3. Pachythyone pseudolugubris, new species
Plate 22, Figs. 9-10; Plate 23, Figs. 1-6; text figure 6
Diagnosis—Medium-sized form (up to 5 cm. long). Resembles
lugubris in the external and inner anatomical features. Spicules an exter-
nal layer of reticulated plates, an inner layer of regularly knobbed 4-holed
buttons rarely elongate and with accessory holes in the ends. Feet with
well-developed end plate ; walls supported by numerous curved supporting
tables with a low spire or none in the ventral feet; in the dorsal a number
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA 117
have a well-developed complex spire, although rarely as large as in
lugubris. Introvert with large tables with numerous holes and reduced
spire. Tentacles exclusively with delicate rods or plates perforated by a
large number of holes. Color mostly mottled brown, sometimes almost
black ; occasionally a pale faded specimen may be encountered in the pre-
served material.
Type.—Holotype, AHF no. 34.
Type locality—Station 744-37, near Point Piaxtla, Sinaloa, Mex.,
Gulf of Calif., April 1, 1937, 6-8 fms.
Distribution From the upper end of the Gulf of Calif. to the Gala-
pagos.
Depth—From 6-25 fms.
Specimens examined.—The type and paratypes and a number of indi-
viduals secured at the following /’elero stations:
145-34. North end of Albemarle Island, 6-7 fms., January 12, 1934,
1 specimen.
Text fig. 6. Pachythyone lugubris (Deichmann).
1-2. Plate from tentacle and table from introvert.
Pachythyone pseudolugubris, new species.
3-4. Plate from tentacle and table from introvert.
Scale, 499 mm.
118 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
264-34. Petatlan Bay, Mex., 25 fms., March 2, 1934, 1 specimen.
683-37. Outside of Concepcion Bay, Mex., March 15, 1937, 30 speci-
mens.
744-37. Near Point Piaxtla, Sinaloa, Mex., Gulf of Calif., April 1,
1937, 7 specimens.
Remarks.—The species resembles /ugubris to an extraordinary degree,
and future workers may decide that it merely represents a variety of that
form. It is easily distinguished by the more simple supporting tables in the
ventral feet, the shorter buttons which seem to lack all tendency to become
lozenge shaped with accessory holes in the ends, and in the larger tables
in the introvert and the absence of heavier rods and plates in the stem of
the tentacles. The larger size and the darker color are, of course, quite
useless characters, as Jugubris possibly may be discovered to reach the
same size and, although a dark color is common, paler individuals have
been found. In no case have embryos been discovered, so presumably this
species is not viviparous as are ugubris and rubra.
The species has in some localities been found with the larger and
paler Neothyone gibbosa, and the possibility has been considered that the
latter represents senescent specimens of pseudolugubris, in which the ex-
ternal layer of reticulated plates has been lost. The spicules seem, how-
ever, to be definitely different. Pachythyone pseudolugubris shows little
tendency to form handles, and the buttons have rarely more than 2 central
knobs (except in the lozenge-shaped buttons), while Neothyone gibbosa
normally forms handles, and as often as not the external handle has 3
knobs. Also the latter species shows definitely a tendency to develop large, ©
smooth supporting plates in the ventral feet, while such plates are almost
unknown in Pachythyone pseudolugubris, where the few supporting plates
which lack a spire are narrow, thick, and in most cases knobbed.
Genus 11. ATHYONE, new genus
Diagnosis—Medium-sized to large forms. Feet numerous, fairly
stout. Tentacles large, bushy, the 2 ventral smaller. Skin soft, with few
spicules. Calcareous ring stout with broad radials with distinct posterior
prolongations and tall interradials with short anterior tooth.
Spicules scattered swollen 4-holed buttons, often incomplete. Feet
with loose-meshed end plate and numerous supporting rods, undoubtedly
derived from tables. Introvert and tentacles with rosettes.
Type species —A thyone glasselli (Deichmann).
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 119
Remarks.—The diagnosis is based on the type of glasselli, a single
adult individual. More material is needed, especially of younger individ-
uals, to complete the diagnosis. The genus is not related to any other form
known from the Panamic region. From the west coast of North America
a form with somewhat similar spicules is known, Thyone benti Deich-
mann, including a variety, var. zacae, from off Cerros Island, Lower Cal-
ifornia, but that species has a taller, more delicate calcareous ring,
exceedingly small tentacles, and larger spicules.
Athyone glasselli (Deichmann)
Thyone glasselli Deichmann, 1936, p. 63, text figure 1 ; 1937, p. 170, text
figure 2. (Comparison with 7. benti.)
Diagnosis.—As for the genus. Color brownish. Resembles superficially
Thyone briareus (Lesueur) from the coast of North America. Shallow-
water form.
Type.—M.C.Z., cat. no. 736.
Type locality—Punta Penasco, Sonora, Mexico.
Distribution—Known only from the type locality.
Depth.—Shore, buried in sand.
Specimens examined.—The type.
Remarks.—The strongly contorted type specimen must have meas-
ured more than 10 cm. in expanded, normal condition. More material is
necessary to explain from what type of spicules the buttons and rods are
derived. Very likely younger individuals may have distinct spire on the
buttons and the supporting rods and thus be derived from tables.
The species is not closely related to any other species known from the
Panamic region. Thyone benti, from Puget Sound, and var. zacae from
Cerros Island have similar spicules but much larger and definitely derived
from tables; the calcareous ring is much more tubular and more delicate
and the tentacles are unusually small. From the southern part of South
America Thyone lechleri Lampert is known to have similar swollen but-
tons, but the calcareous ring is entirely different with no posterior pro-
longations at all.
The only other soft-skinned Thyone with few spicules described from
the Panamic region is the dubious Anaperus peruana Troschel, possibly
identical with Lesson’s species (see discussion, p. 120).
Regarding Thyone ovulum Selenka, which Deichmann (1936, p. 65)
mentioned as possibly the juvenile stage of 4. peruana, the type has been
re-examined and found to be a phyllophorid (see p. 125).
120 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Genus 12. ANAPERUS Troschel
Anaperus Troschel (partim), 1846, p. 60.
Diagnosis —Large form with soft skin and numerous feet distributed
without order. Tentacles 10, the 2 ventral smaller. Anus with calcareous
teeth. Calcareous ring with posterior prolongations (apparently short) ;
retractor muscles fleshy. Spicules not numerous except in the stem of the
tentacles.
Type species.—A naperus peruviana (Lesson).
Remarks.—As type species for this dubious genus Lesson’s large den-
drochirote species from Payta, Peru, has been chosen since it is a question
whether Thyone briareus (Lesueur)from the western Atlantic belongs in
the genus; Thyone fusus (O. F. Miiller)—the type species for Thyone
s. str.—does definitely not belong here, and the fourth species, listed by
Troschel as Thyone cigaro Troschel, from Labrador, cannot be identified.
It is furthermore doubtful whether Lesson’s and Troschel’s species
are identical. Very likely Lesson’s species is a phyllophorid (Pattalus
mollis (Selenka) ), while Troschel’s species is a soft-skinned Thyone
which resembles T. briareus; (Selenka actually united the 2 species after
having examined Troschel’s material in Berlin).
The only other soft-skinned large Thyone known from the upper end
of the Gulf of California is Athyone glasselli (Deichmann), which
possibly may occur as far south as Peru. If we discount Troschel’s state-
ments about the color and size of his 4. peruana—as they seem to be
borrowed from Lesson’s description—the assumption is not unreasonable.
If re-examination of Troschel’s material in Berlin should prove that that
is the case, the name Athyone becomes a synonym of Anaperus; in that
case Thyone briareus is excluded from the genus, and Troschel’s species
must be named Anaperus glasselli (Deichmann), syn. 4. peruana Tro-
schel, nec Holothuria peruviana Lesson.
Another possibility is that Troschel’s species is simply material of
Thyone briareus which has been wrongly labeled. Such errors were by no
means unusual in older times.
Anaperus peruviana (Lesson)
Holothuria peruviana Lesson, 1830, p. 124, pl. 46, fig. 1.
Trepang peruana Jaeger, 1833, p. 25.—Brandt, 1835, p. 57.
Anaperus peruanus Troschel, 1846, p. 61.—J. Miiller, 1854, pl. 9, fig. 9.
NO. 3. DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 121
Thyone peruana Selenka, 1867, p. 354.—Lampert, 1885, p. 160.—Sem-
per, 1868, p. 242.—Théel, 1886, p. 140.—Deichmann, 1936, p.
65 (passim).
Nec Anaperus carolinus Troschel, 1846, p. 62, nec Thyone tenella Selen-
ka, 1867, p. 354, pl. 20, figs. 113-114. Referred to Thyone peru-
ana by Selenka, 1868, p. 118 (both are synonyms of Thyone bri-
areus and both came from the Atlantic coasts of North America).
Diagnosis—No diagnosis can be given, as Lesson and Troschel pos-
sibly had two different species before them.
Type.—Probably lost, or in Paris.
Type locality—Payta, Peru.
Remarks.—Lesson’s description and figure refer to an exceptionally
large form, 6 inches long, with 8 bushy tentacles, numerous feet, and
soft skin. The color of the animal was deep purplish.
Troschel’s numerous specimens in Berlin, likewise from Peru, refer
apparently to a typical Thyone. They seem to resemble the West In-
dian Thyone briareus Lesueur and have similar short posterior pro-
longations on the radials and distinct anal teeth; the 2 ventral tentacles
are smaller (according to Troschel they were probably overlooked by
Lesson, since he figured only 8). The calcareous spicules were not
studied ; Troschel mentions merely that they were numerous in the stem
of the tentacles. The size and color given by Troschel may very likely be
quoted directly from Lesson.
Selenka re-examined Troschel’s material and decided that Anaperus
carolinus Troschel and his own Thyone tenella from, respectively, Caro-
lina and Texas were synonyms of the Peruvian form (both names are
synonyms of Thyone briareus Lesueur, the common large Thyone from
the eastern coasts of the United States).
Lampert may possibly have examined some specimens of 7". briareus
and based his few remarks concerning Thyone peruana on a study of this
material but not on the true Peruvian form; J. Miiller discusses (1854,
p. 85) the stone canal and figures (p. 9), one from 4. peruana, possibly
from Troschel’s material, but nothing of importance can be gleaned from
his drawing.
Whether Selenka for once was hasty in his identification or whether
Troschel’s material was mislabeled and actually represented material of
Thyone briareus is impossible to decide. If the latter is true, it is obvious
that the statements regarding the size and color came from Lesson’s de-
122 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
scription. So far, no species which resembles Thyone briareus is known
from the Pacific coasts of America except 4. glassellt.
To solve the question it is necessary to re-examine Troschel’s material
in Berlin and possibly Lesson’s type, if still extant. Very likely Lesson de-
scribed Pattalus mollis, the only large purplish form, and overlooked
some of the tentacles, while Troschel had either mislabeled material of
Thyone briareus before him, an undescribed Thyone, or Athyone glas-
selli,
Family II. Phyllophoridae
Diagnosis —Dendrochirota with more than 10 tentacles, the number
ranging from 15 to 30. The tentacles either definitely of two sizes, large
in the external circle, small in the inner, and appearing almost simultane-
ously, or the sizes intergrade, the 2 circles being indistinctly set off from
each other, and the full number is reached comparatively late in life.
Feet restricted to the ambulacra, or scattered also in the interambulacra.
Calcareous ring simple, or with distinct posterior prolongations; stone
canal and Polian vesicle often in large numbers.
Spicules of various kinds, mostly as tables, often reduced with age.
Feet with end plate; walls with or without supporting tables or rods.
Introvert with tables or plates, tentacles with or without spicules. The
majority are shallow-water forms.
Remarks.—A preliminary revision has been undertaken (Deichmann,
1938) which includes all the genera known from the American waters.
To this list is now added Athyonidium, established for the species for-
merly known as Thyone chilensis Semper. The latter is synonymous with
Eucyclus duplicatus, new genus, new species, established by Lampert,
1885, pp. 290-292. Lampert was fully aware of the close similarity to
Semper’s species Thyone chilensis from Chile. Théel discussed the ques-
tion; he believed the two species were identical but concluded that the
number of tentacles possibly was more variable in the polychirote forms
than generally known. Ludwig concurred in Théel’s opinion and assumed
that the tentacles were merely overlooked by Semper. He rejected, how-
ever, the genus Eucyclus as superfluous and united it with Thyonidium
under Phyllophorus. The Velero III has procured ample material of a
form which unquestionably refers to the species which Semper examined.
As in the case of Thyone ovulum Selenka—now Euthyonidium—in which
Selenka overlooked the inner circle of smaller tentacles, so here the inner
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 123
circle of smaller tentacles was completely missed by Semper and the spe-
cies therefore likewise referred to Thyone.6 The genus resembles the
northern genus T'hyonidium but is distinguished by the almost completely
resorbed interradials. As Eucyclus is preoccupied, a new genus had to be
established.
Key TO THE GENERA OF PHYLLOPHORIDAE KNOWN FROM THE
PANAMIC REGION AND CHILEAN WATERS
All the genera known from this region have at utmost 20 tentacles,
and the feet are numerous in the interambulacra as in the genus Thyone.
1. ‘Tentacles distinctly of 2 sizes, i.e., 10 large ones in the external
circle and either 5 small ones, more or less deeply divided, or 5
small pairs of equal size inthe inner circle. . . .... 2
1. ‘Tentacles of varying number and size, sometimes of almost the
same size; the 2 circles are more or less confluent. . . . . 3
2. Calcareous ring low, simple (or with insignificant posterior
projections) ; interradials well developed, often pentagonal or
heart-shaped—in many cases overlapping the radials. Spicules
derived from 2- to 4-pillared tables, often reduced to plates
with 2-4 knobs or spines indicating the pillars. Feet with end
plate but no supporting tables, sometimes a few rods. Tentacles
with rosettes and delicate rods. 1. Euthyonidium Deichmann p. 124
2. Calcareous ring low, simple; radials with long anterior tooth,
interradials completely vestigial, easily overlooked. Spicules
apparently lacking except for the end plate and a few branching
rods. . ... .. =. . J Athyonidium, new genus p. 127
3. Calcareous ring simple; radials tall, posteriorly slightly incised ;
interradials well developed. Spicules almost totally lacking,
except for end plate and a few perforated plates. Tentacles 20,
large, of almost equal size. Large reddish species. . . .
3. Pattalus Selenka p. 129
3. Calcareous ring with long posterior prolongations. Tentacles
of different size, fairly delicate. Spicules 2- to 4-pillared tables
sometimes scarce in older individuals. Small- to medium-sized
TOrS. 2 =. . © + « » «+ 4: Phyllophorus Grubep. 152
6 In other specimens Semper noticed the inner circle of smaller tentacles and
these he referred to Pattalus with a comment on the variability of the tentacles in
this genus.
124 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Genus 1. EUTHYONIDIUM Deichmann
Euthyonidium Deichmann, 1938, p. 380.
Diagnosis.—Medium-sized form (length 10 cm. or less). Tentacles
definitely arranged as 10 large ones in 5 external pairs and 5 inner pairs
(or 5 single ones?)? which are smaller. Apparently all the tentacles ap-
pear at an early stage. Calcareous ring simple with tall rectangular radials,
posteriorly with short protuberances which may be interpreted as rudi-
mentary prolongations. Interradials well developed.
Spicules small tables with oval to squarish disk, often with dentate or
branching margin; spire low, 2-4 pillared often reduced to knobs or spines
or totally resorbed. Feet with large end plate and often a few small sup-
porting rods, rarely with trace of being derived from tables. Introvert
with tables with numerous holes in the disk and spire often reduced.
Tentacles with rosettes and delicate rods. Spicules often reduced with
advancing age. Shallow-water forms, restricted to the tropical waters.
Type species —E. seguroensis (Deichmann).
Remarks.—F rom the West Indies 2 species are known, viz., the type
species and E. occidentalis (Ludwig). From the Panamic region 2 species
are now known; one represents the species which formerly was known as
Thyone ovulum Selenka, and the other, which is new, is closely related
to E. occidentalis from the West Indies.
Key TO THE SPECIES OF Euthyonidium KNOWN FROM THE
PANAMIC REGION
1. Spicules oblong tables with dentate edge; spire 2 pillared, low,
often reduced to knobs or spines, or lacking. Color grayish. .
. . . . . «. 2. Euthyonidium veleronis, new species
1. Spicules large tables with cross-shaped disk, often irregularly
laciniated ; spire mostly reduced to a few knobs or spines. Color
reddish brown. . . . 1. Euthyonidium ovulum (Selenka)
1. Euthyonidium ovulum (Selenka)
Plate 24, Figs. 1-7
Stolus ovulum Selenka, 1867, p. 356, pl. 20, fig. 117—Semper, 1868,
p. 66.—Lampert, 1885, p. 162.—T héel, 1886, p. 138.
7 The inner smaller pairs may possibly in some forms be found to be 5 single
tentacles which are deeply divided as Heding (1936, p. 23) has found it to be the
case in the type species of the closely related genus Thyonidium from the northern
seas.
a
_ -
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 125
Thyone ovulum Deichmann, 1936, p. 64 (passim).
Euthyonidium ovulum Deichmann, 1938, p. 380, text figure 12.
Diagnosis—Medium-sized form (probably about 10 cm. long) with
soft, thick skin with numerous soft feet. Tentacles 10 large external ones
and 10 inner ones which are much smaller and may be completely re-
tracted into small pockets. Calcareous ring with radials posteriorly broad
and deeply incised and tapering toward the anterior end; interradials
unusually broad and heart shaped with a slight posterior incision, ante-
riorly tapering to a point. Stone canals numerous and free, also several
Polian vesicles. Inner anatomy otherwise normal.
Spicules a scattered layer of tables with irregular disk varying from
cross-shaped to a more or less branching or laciniated plate. Spire almost
always reduced to a few spines. Feet with large end plate; supporting rods
are apparently totally lacking. Introvert seems to lack spicules; tentacles
with few rosettes and delicate rods. Spicules more or less completely re-
duced with advancing age and, except for the end plate, likely to be com-
pletely overlooked.
Type.—M.C.Z.
Type locality—Acapulco, Mexico.
Distribution—From Lower California to Peru.
Specimens examined.—Selenka’s type specimens (3) and several spec-
imens from the Allan Hancock Expeditions:
261-34. angola Tangola, Mex., 1 fm., March 1, 1934, 1 specimen.
591-36. Port Escondido, L. Calif., Mex., shore, March 16, 1936, 40
specimens.
844-38. Lobos de Afuera Island, Peru, shore, February 14, 1938, 1
specimen.
Remarks.—Re-examination of Selenka’s type material of Stolus ovu-
lum showed that Selenka had overlooked the inner circle of tentacles
which were completely withdrawn into little pockets and that it was
identical with various individuals collected by the Hancock Expeditions,
and also by the Zaca.
Euthyonidium ovulum has the same reddish-brown color which is
extracted in alcohol as the much larger species Pattalus mollis Selenka
from Peru and Chile. It would therefore be correct to consider whether
the two forms are merely growth stages of the form which possibly ought
to be called Pattalus peruana (Lesson)—see discussion under Pattalus
and Anaperus. Deichmann (1936) considered ovulum as possibly being
identical with Lesson’s species (which was figured as having 8 tentacles)
126 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
but was at that time unaware that ovulum was not a true Thyone. It
would, however, be a hitherto unknown phenomenon that the smaller
tentacles in the inner circle of a phyllophorid should be able to catch up
in size with the large ones, and for that reason alone it would seem
unlikely that ovulum could grow into Pattalus mollis usually with about
20 tentacles of almost identical size. Furthermore, the latter shows great
variation in the number of tentacles, while ovu/um invariably seems to
acquire the full number at an early age.
2. Euthyonidium veleronis, new species
Plate 24, Figs. 8-11
Diagnosis—Medium-sized form, up to 10 cm. long; body tapering
toward both ends with numerous soft feet scattered over the interambu-
lacra; toward the oral and anal ends arranged in a more orderly manner
along the ambulacra. Tentacles 5 large external pairs, and 5 inner smaller
pairs closely appressed.§ Calcareous ring with 5, almost rectangular radi-
als; posteriorly with short projections; interradials more or less heart
shaped, with the point forward directed, and overlapping the radials.
Stone canal single, attached in the dorsal mesentery; Polian vesicles usu-
ally 3 in number, muscle stomach delicate; intestine with normal course;
retractors attached near the middle of the body where the 2 tufts of
gonads are also attached.
Spicules a scattered layer of small tables with oval to squarish disk
with about 8 holes and dentate edge; spire low, mostly with 2 pillars,
often reduced to knobs. Feet with large end plate and occasionally a few
supporting rods. Introvert with tables; tentacles with rosettes and small
curved rods, mostly with perforated ends. Color mottled grayish.
Type.—Holotype, AHF no. 35.
Type locality—Station 465-35, Playa Blanca, Costa Rica, shale
beach between sand beach and rock reef, shore, February 8, 1935, 1 speci-
men.
Distribution.—The type locality.
Depth.—Shore.
Specimens examined.—The type.
Remarks.—The species is closely related to E. occidentalis (Ludwig)
known from the West Indies.
8 As only the type was available, the tentacles in the inner circle were not dis-
sected out, but very likely this species has only 5 tentacles, deeply cleft, in the inner
circle.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 127
Genus 2. ATHYONIDIUM, new genus
Eucyclus Lampert, 1885, p. 290.—Théel, 1886, p. 268.—Ludwig, 1887,
p. 1239 (reprint p. 23). Name preoccupied.
Diagnosis—Large forms (25 cm. or more) with thick, soft skin and
numerous stout feet. Tentacles 5 large external pairs and 5 small inner
pairs. Calcareous ring with 5 stout, almost rectangular radials and 5 in-
significant interradials, often completely concealed in the tissue.
One large dorsal stone canal, often branched and one or two tufts of
smaller stone canals with minute heads. Numerous tufts of branched
Polian vesicles. Inner anatomy seems otherwise quite typical of the family.
Spicules large well-developed end plates, a few perforated spinous
rods; tentacles with few rosettes in the younger individuals. Color gray-
ish mottled to almost black, ventrum paler, tentacles dark. Shallow-water
forms.
Type species—A thyonidium chilensis (Semper).
Remarks.—The genus is monotypic and related to Thyonidium and
Euthyonidium, but the strikingly reduced interradials give it a position
almost unique among the Phyllophoridae. Very little can be said about the
spicules, as they are reduced even in small individuals (5 cm. long). They
may be present in still younger specimens, and one would expect them to
be tables, possibly with the spire reduced.
Athyonidium chilensis (Semper)
Thyone (Stolus) chilensis Semper, 1868, p. 241, pl. 40, figs. 3-6—Lam-
pert, 1885, p. 156.—Théel, 1886, p. 139.
Eucyclus duplicatus Lampert, 1885, p. 250.—Théel, 1886, p. 268.
Diagnosis.—As for the genus.
Type.—Possibly in Germany.
Type locality—Coast of Chile.
Distribution—From Chile to Ancon, Peru.
Depth.—Shallow water, at tidemark, in sand.
Specimens examined.—The following material from the Allan Han-
cock Expeditions:
828-38. San Juan Bay, Peru, shore, February 8, 1938, numerous speci-
mens.
831-38. Independencia Bay, Peru, shore, February 9, 1938, 3 specimens.
Also the following specimens in the M.C.Z.: 21 from Talcahuana,
Chile; 3 thrown up on the beach in Ancon, Peru, with Pattalus mollis.
128 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Remarks.—I have ventured to transfer Semper’s Thyone chilensis to
this genus, as his description of that unusual form agrees exactly with the
numerous specimens I have had occasion to examine, and the species
differs distinctly from Pattalus mollis Selenka, of which I have likewise
had the opportunity to study a long series. My results are in complete
agreement with Ludwig's conclusions, that Lampert’s species is a syno-
nym of Semper’s Thyone chilensis (and probably some or all of his
Pattalus mollis). But contrary to Semper’s and Théel’s belief, the num-
ber of tentacles is constant in this species.
The inner circle of tentacles is often completely withdrawn so that
only the 10 larger ones are visible. The inner tentacles become quite stout
with advancing age but remain, nevertheless, always distinctly smaller
than the external ones, and the inner circle is well separated from the
outer one. The radials, as Semper describes them, are slipper shaped, al-
most rectangular, broadest posteriorly with a large depression below the
attachment for the retractors; the interradials are small in the young indi-
viduals; in the older they are completely concealed in the thick tissue, and
the calcareous ring appears to be composed of 5 stout radials. Semper
noticed only one tuft of small accessory stone canals but may possibly
have failed to examine both sides of the mesentery. The Polian vesicles
are distinctly arranged in tufts, and most of them are branched.
Except for the presence of end plate and a few rosettes in the tentacles,
spicules are so rare that they often are difficult to find. Semper figures a
few branching rods perforated by a few holes, and such spicules have been
found.
The color is strikingly different from that of Pattalus mollis even in
preserved individuals. It is mottled gray, sometimes almost black on the
dorsal side, with paler ventral side and black tentacles. In freshly col-
lected individuals the tube feet may have a reddish tinge, possibly caused
by the hemoglobin contained in the water vascular system. The skin seems
never to contain a reddish pigment as Pattalus mollis, nor does it ever
fade to that uniform dull reddish brown which is so characteristic of
Selenka’s species.
Athyonidium chilensis seems to be the most common phyllophorid
known from Chile. On the coast of Peru it is often known to occur in the
same localities as Pattalus mollis, but buried in sand with the tentacle
crown expanded on the surface, while the former clings to the rocks
(H. L. Clark’s observations).
No.3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 129
The Velero material measures from 5 to 20 cm.; the largest individu-
als in the M.C.Z. measure more than 25 cm. in length.
The occurrence of this typical Chilean species in Peruvian waters
makes it impossible to decide whether this species or Pattalus mollis rep-
resents Lesson’s troublesome form, Holothuria peruana. Possibly a care-
ful survey at Payta, the type locality for perwana, may show that only one
of the 2 species occurs there. Pattalus mollis Selenka does occur in
Payta (material in the M.C.Z.), but 4. chilensis also may possibly be
found there, for both species are known to occur in the same localities,
i.e., at Ancon, Peru, and at Station 828-38.
Genus 3. PATTALUS Selenka
Pattalus Selenka, 1868, p. 113.
Diagnosis —Large form (20 cm. or more) with up to 20 large tenta-
cles of almost equal size. Skin thick with numerous tube feet. Calcareous
ring with 5 tall rectangular radials, with posterior corners rounded into
low protuberances; interradials with broad low basal part and a long
anterior tooth, often concealed in the thick tissue. Dorsally a long stone
canal with small head; Polian vesicles numerous, hanging as a fringe
around the ring canal, a few are dichotomously divided. Other anatomical
features typical of the family.
Spicules a few branched rods, feet with large end plate. Color reddish
brown with a pigment which is dissolved in alcohol ; specimens which have
been preserved in alcohol for a long time are uniformly dull brown. Shal-
low water.
Type species —Pattalus mollis Selenka.
Remarks.—The genus is monotypic and seems not related to any other
form. It has in the course of time been confused with Athyonidium chil-
ensis (Semper) but is definitely different from that species. It may pos-
sibly be identical with Euthyonidium ovulum (Selenka), although it
seems impossible that the smaller inner tentacles should ever be able to
attain the size of those in the external circle. The 2 species have the same
reddish color with pigment which is extracted in alcohol. They differ as
far as our present material shows in the more heart-shaped interradials in
E. ovulum, with a short anterior tooth, while P. mollis has a remarkably
long anterior tooth on the interradials. Also E. ovulum seems to lack a
large, dorsally attached, stone canal which is so characteristic a feature in
P. mollis. The large P. mollis is extremely common in Peru and is also
130 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
known from Concha, Chile, while the smaller E. ovulum is known from
various localities in the Panamic region and has once been reported from
Lobos de Afuera Island, Peru (see above, p. 125). (The M.C.Z. has fur-
ther two lots marked “locality dubious” from, respectively, Hawaii, and
Mazatlan, Mexico.) If Pattalus mollis is assumed to represent the full-
grown stage of Euthyonidium ovulum, it is impossible to explain how that
large species could have escaped discovery in the Panamic region. (See
also under E. ovulum.)
Pattalus mollis Selenka
Pattalus mollis Selenka, 1868, p. 113, pl. 8, figs. 4-5.
Thyonidium peruanum Semper, 1868, p. 67, pl. 15, fig. 17.
Thyonidium mollie Semper, 1868, p. 243.—Ludwig, 1874, p. 4; 1883,
p. 163; 1887, p. 25.—Lampert, 1885, p. 174 (partim).—Theel,
1886, p. 144.
Anaperus peruana Verrill, 1867, p. 322.
Pattalus peruana Verrill, 1867, p. 376.
Phyllophorus peruvianus H. L. Clark, 1910, p. 351 (partim).
Diagnosis—As for the genus. Color reddish purplish, pigment ex-
tracted in alcohol; preserved specimens usually dull brown.
Type.— Berlin.
Type locality —Peru and Chile.
Distribution —Common in Chile and Peru.
Depth.—Shallow water, to 25 fms., often collected at tidemark, cling-
ing to rocks (H. L. Clark).
Specimens examined.—A large series in U.S.N.M. and M.C.Z. Also
several specimens from the Allan Hancock Expeditions:
375-35. Independencia Bay, Peru, shore, January 13, 1935, 13 speci-
mens.
391-35. Lobos de Afuera Island, shore, January 17, 1935, 8 specimens.
820-38. San Nicholas Bay, Peru, 10-25 fms., February 6, 1938, 4 speci-
mens.
828-38. San Juan Bay, Peru, shore, February 8, 1938, 9 specimens.
831-38. Independencia Bay, Peru, shore, February 9, 1938, 3 specimens.
837-38. North Chincha Island, Peru, shore, February 11, 1938, 6 speci-
mens.
844-38. Lobos de Afuera Island, Peru, shore, February 14, 1938, 2
specimens.
NO. 3. DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 131
Remarks.—The species seems to be the most characteristic holothurian
from the coasts of Peru and certain localities in Chile. In some cases it
occurs in the same localities as Athyonidium chilensis, but the 2 species
have an entirely different ecology. Pattalus mollis is found completely
exposed, clinging to rocks, while Athyonidium chilensis is found buried
in sand with its tentacles extending flatly on the surface (H. L. Clark,
observed on the Hancock Expeditions in the spring of 1938). In pre-
served condition the 2 species are entirely unlike except in size; P. mollis
is uniformly purplish red or faded brown, while the other species is mot-
tled gray, sometimes with almost black dorsum, black tentacles, and more
lightly colored ventrum. Internally P. mollis has large well-developed
interradials with long anterior tooth, while Athyonidium chilensis has
broad radials and the interradials are almost completely resorbed or hid-
den in the leathery tissue.
It would be tempting to select Lesson’s name peruanus for this species
so typical of Peru, especially since material in the M.C.Z. shows that
Pattalus mollis does occur in Payta, Peru, the type locality for Lesson’s
species. But there is still the possibility that Lesson did have a true deka-
chirote form before him, with 2 smaller ventral tentacles which were
overlooked, a form which has not as yet been rediscovered except possibly
by Troschel. Furthermore, both P. mollis and A. chilensis often occur in
the same localities, and Lesson’s description and figures may refer to a
composite species, the low number of tentacles—although incorrect—
being taken from J. chilensis, while the color note “rouge violet éclatant
et foncé” refers to P. mollis.
No other species has caused so much confusion in the literature as this
species (including A. chilensis). It has usually been identified with Les-
son’s Holothuria peruana, although, if Troschel is correct, the latter is
one of the members of the genus Thyone s.l.
Semper obviously referred some of his material of 4. chilensis—those
in which he observed the 10 inner small tentacles—to this species and
commented on the variability of the number and size of the tentacles in
this genus; Lampert did the same, as also H. L. Clark.
Verrill referred his material of mollis to Anaperus peruanus Troschel
but realized the discrepancy in the number of tentacles and corrected the
name to “Pattalus peruanus—closely related to Selenka’s P. mollis,”
without indicating wherein the 2 species differed.
The reasons why P. mollis cannot represent the mature stage of
Euthyonidium ovulum Selenka are mentioned above (under that species).
132 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Genus 4. PHYLLOPHORUS Grube, 1840
Phyllophorus Auctores. Nec Phyllophorus Heding, 1936 (Thyonidium
Diiben & Koren).
Thyonidium Auctores. Nec Thyonidium Diiben & Koren, 1844, p. 303.
Diagnosis —Medium-sized forms (6-15 cm.) with feet distributed in
the interambulacra. Tentacles up to 20 in number, of varying size, not
sharply set off into 2 circles; the complete number is reached late in the
animal’s life. Calcareous ring with distinct posterior prolongations on
the radials; interradials well developed, of varying shape; ring often com-
posed of smaller pieces, mosaiclike.
Spicules 2- to 4-pillared tables, spire usually retained. Feet with large
end plate; walls with or without supporting tables, or a few rods. Intro-
vert usually with tables with low spire and numerous perforations in the
disk; tentacles with either perforated plates or delicate rods; rosettes
seem normally to be present in the introvert and tentacles. Shallow-water
forms, tropical and subtropical.
Type species—Phyllophorus urna Grube.
Remarks.—The type species is common in the Mediterranean Sea.
Grube’s description and figures are not particularly convincing, but an
excellent account is given by Sars (1846). Unfortunately it is written in
the Norwegian language and is therefore not so well known as it deserves.
A number of tropical species belong to this genus. (See Engel, 1933, and
Deichmann, 1930 and 1938.)
From American waters 5 species are reported from the West Indies
and coast of Brazil. From the tropical west coast of America 2 species
are known. One was described by Ludwig in 1894, the other by Deich-
mann in 1938. The latter species is closely related to one of the West
Indian forms, while that of Ludwig seems completely unrelated to all
other members of the genus. No true Phyllophorus is known from the
west coast of North America, or from Chile or Hawaii.
Key TO THE SPECIES OF Phyllophorus KNOWN FROM THE
PANAMIC REGION
1. Spicules tables with oval disk with an average of 8 marginal
holes; spire distinctly 4 pillared, usually with a single cross-
beam and ending in a large number of spines. Feet with large
end plate and numerous supporting tables with broad elongate
disk with numerous holes and a huge spire ending in a conical
No.3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA 133
tooth which often pierces the skin. :
1. Phyllophorus ees rm ae
1. Spicules tables with oval disk with 4 large and 4 smaller holes,
the latter often wedge shaped ; spire usually with 2 pillars and
ending in 2 tufts of few spines, sometimes ending in 4 spines.
Feet with large end plate and a few supporting tables with
broad elongate disk with numerous holes and comparatively
low spire with a single crossbeam and ending in 2 diverging
spines or horns, rarely in a single point. ;
See 6 ee We eb hey oe Pepliaphorus sarac Dehn
1. Phyllophorus aculeatus Ludwig
Plate 25, Figs. 1-8
Phyllophorus aculeatus Ludwig, 1894, p. 128, pl. 13, figs. 6-11.—Deich-
mann, 1938, p. 381 (passim).
Diagnosis —Medium-sized form (type measured 6.5 cm. but was
strongly contracted). Feet numerous, scattered almost uniformly over
the body. Tentacles of varying size, up to 20 in number. Calcareous ring
with long posterior prolongations on the radials; interradials with heart-
shaped to almost rhomboid base and long anterior tooth; the basal part
overlaps the radials.
Spicules numerous tables with oval disk with a varying number of
marginal holes, usually 8; spire mostly with 4 pillars, a single crossbeam,
and a wreath of small spines on the top. Feet with large end plate and
heavy supporting tables with broad oval disk with numerous holes and an
enormous spire which tapers into a conical tooth. Introvert with tables
with oval, lacelike disk and spire composed of 2 to 4 pillars. Tentacles
with a few rosettes and numerous delicate rods, rarely with the ends per-
forated. Color grayish with purplish tentacles.
Type.—U.S.N.M.
Type locality —Gulf of Panama, shore.
Distribution Known from the type locality only.
Depth.—Shallow water.
Specimens examined.—The type, the only specimen known in exist-
ence in any collection.
Remarks.—The strongly contracted type measured probably more
than 10 cm. in length in expanded condition. ‘The inner organs are strong-
ly distorted, and very little can be said about them. The gonads are well
developed, forming long tubes, dichotomously divided near the base.
134 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
The spicules show some similarity to those characteristic of Thyone
strangeri (see pl. 19), and for a while I thought that Ludwig had made
a mistake in counting the tentacles. Re-examination of the type and com-
parison of the spicules showed beyond doubt that Ludwig was right and
that the 2 species cannot possibly be confused as a glance at the figures of
the spicules will show.
It is remarkable that this well-defined species has never been taken
again, especially since the type locality is one of the most frequently vis-
ited collecting grounds in the Panamic waters.
2. Phyllophorus zacae Deichmann
Plate 24, Figs. 12-21
Phyllophorus zacae Deichmann, 1938, p. 381.
Diagnosis —Medium-sized form, feet numerous scattered almost uni-
formly over the body, not crowded. Tentacles 15-20 in number, long,
delicate, of varying size. Calcareous ring with well-developed posterior
prolongations on the radials; the anterior tooth is long rectangular with
a constriction at its base; the interradials are more or less heart shaped
with a broad base, overlapping the radials and with a short tooth.
Spicules delicate tables with oval disk with 4 larger and 4 smaller
holes, the latter often wedge shaped; spire with 2 to 4 pillars, a single
crossbeam, and ending in 4 tufts of few spines, sometimes reduced to 4
undivided spines. Feet with large end plate and a few supporting tables
with broad oblong disk with numerous holes; spire low or moderately
high with one crossbeam, and usually ending in 2 diverging spines or
horns or in a single point. Introvert with delicate tables; tentacles with
delicate rods and rosettes. Color reddish to white.
Type.-—M.C.Z.
Type locality —Tangola Tangola Bay, Mex.
Distribution Ranging from L. Calif. to the Galapagos.
Depth.—The type came from 10 fms. depth.
Specimens examined.—The type, and 4 individuals from the follow-
ing Velero III stations:
190-34. Galapagos, Lat. 0° 55’ S.; Long. 90° 30’ W., 58-60 fms., Jan-
uary 26, 1934, 2 young.
517-36. East side of San Francisco Island, Gulf of Calif., 15 fms., Feb-
ruary 25, 1936, 1 specimen, oral end missing.
816-38. North of Hood Island, Galapagos, 50-100 fms., January 29,
1938, 1 specimen.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 135
Remarks.—The species is closely related to Phyllophorus destichadus
Deichmann from the West Indian waters. It differs in the presence of
supporting tables in the feet and in minor details of the calcareous ring
and the spicules.
A species with somewhat similar spicules was described from Hawaii
by Fisher, 1907, and is now referred to Neothyonidium (Deichmann,
1938, p. 382). The Hawaiian species has tentacles in 2 separate circles
and of 2 different sizes, and the calcareous ring is of another type, with
long narrow interradials which are firmly joined to the radials, not over-
lapping them.
Family III. Psolidae
Diagnosis —Dendrochirota with part of the ventral side developed as
a thin-walled sole, with feet along the margin and in many forms also
along the odd ambulacrum of the sole. Dorsal side (including the oral
and anal ends) covered more or less completely by scales; in certain
genera some scales are perforated for the passage of tube feet; in one
genus no feet are developed on the dorsal side. Tentacles 10 (15 in one
genus), the 2 ventral ones are often smaller; in the few deep water forms
the tentacles are almost finger shaped, with few or no branches. Cal-
careous ring simple; retractors of unequal length; gonads in 2 tufts on
the dorsal side opening on a papilla behind the tentacles.
Spicules in sole, perforated plates or buttons, smooth or knobbed; in
some forms the external deposits may be cup shaped, in others gradually
transformed into reticulated bodies; in certain species the spicules become
scarce with age. Feet with end plate and a varying number of supporting
rods or plates. Dorsal side with or without a layer of spicules outside the
scales, as grains, cups, hourglass-shaped bodies, plates, etc. Dorsal ap-
pendages, if present, with or without end plate and supporting rods or
plates. Tentacles with or without perforated plates or rods; sometimes the
spicules disappear in older individuals; a few forms have rosettes.
Remarks.—The family comprises about half a dozen genera and
represents a rather interesting series of evolutionary steps. Of the 3 genera
which are known from the Panamic waters Thyonepsolus represents un-
doubtedly the most primitive group, indistinctly separated from Psolid-
ium, while the highest evolutionary stage is represented by the genus
Psolus. The inner anatomy of the 3 genera is very similar. The most
striking difference is that in the 2 first-named genera the third loop of the
136 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
intestine is attached in the left ventral interambulacrum, while in the
last genus it is attached in the right interambulacrum. This may indicate
that the T’hyonepsolus-Psolidium series represents an independent branch
and the Psolus series another, or it may simply be that the mesenterial
attachment has shifted completely over in the most advanced genus. Some
of the members of the genus Psolidium have very few dorsal feet, and
these have formerly partly been referred to the genus Psolus.
Key TO THE GENERA OF THE FAMILY Psolidae KNOWN FROM THE
PANAMIC REGION
1. Dorsal side (including oral and anal ends) without appendages
(except the anal papillae and in some forms a few abortive feet
on the introvert). ..... » = .«.«, «, 3: Psolus Oken.p. 146
1. Dorsal side with a varying number of tube feet, usually perfor-
ating the SC@less\) iis) bac et a ee’ Men Sea Se eee ee eee
2. Dorsal side characterized by the presence of large hourglass-
shaped bodies. Tube feet on dorsal side with walls stiffened by
numerous curved squarish or circular plates, sometimes reduced
to crosses (also a few oblong supporting plates may be pres-
ent) ; towerlike deposits present in most species, often reduced
With age: 2 . . . . «© 1 Thyonessolus BOL. Clark ps 136
2. Dorsal side without large hourglass-shaped bodies, at utmost
small cups or baskets. Dorsal tube feet in some forms supported
by squarish plates, in others by oblong plates or rods, in others
superficial deposits seem to be entirely lacking on the dorsum;
towerlike deposits sometimes present. 2. Psolidium Ludwig p. 141
Genus 1. THYONEPSOLUS H. L. Clark, 1901
Thyonepsolus H. L. Clark, 1901, p. 169.—Deichmann, 1930, p. 192;
1937p 72:
Psolidium Ludwig, 1904, p. 689.
Lissothuria Verrill, 1867, p. 322.
Diagnosis—Small to medium-sized forms with 10 tentacles, the 2
ventral smaller; sole sharply set off; feet numerous in 3 crowded bands.
Dorsal scales few to numerous, covering the back more or less completely ;
feet numerous or few; a well-developed external layer of spicules covers
the scales.
No.3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA 137
Spicules in sole perforated plates, knobbed or smooth, some may form
shallow baskets; feet with end plate and perforated supporting rods or
plates. Dorsal side with a varying number of large towerlike deposits,
reduced with age, in some species shorter, approaching the shape of grains;
also hourglass-shaped bodies and curved plates. Feet with a small end
plate or none, and the walls supported by curved squarish plates, some-
times reduced to crosses and rods. Tentacles with heavy perforated plates
and smaller plates and rods, in some species rosettes. Shallow-water forms.
Type species—Thyonepsolus nutriens H. L. Clark.
Remarks.—The genus seems to be restricted to the American waters.
The type species is known from Monterey Bay, California, and south-
ward to Santa Cruz Islands; another species is known from the West
Indian seas, while 3 occur in the Panamic region and adjacent waters.
Two of these are here described as new. The type species is known to be
viviparous; the life histories of the other species are yet unknown.
Kery TO THE SPECIES OF Thyonepsolus KNOWN FROM THE
PANAMIC REGION AND THE CALIFORNIAN WATERS
1. Dorsal scale covering incomplete in the midline (in the adult
individuals). Scales numerous and small; dorsal feet numer-
ous; external layer of spicules well developed. Sole with large
perforated plates and an outer layer of flattened baskets. Per-
forated plates in tentacles decreasing in size, no rosettes. Vivip-
arous; carries its large eggs in pits on the back. Californian
waters (and Galapagos ?). . se ae
epee rh ee aa al cre Oe nutriens H. iD ieee
f. Darsdl scale covering complete in the midline. Dorsal feet not
remarkably numerous; external layer of spicules not remark-
ably crowded, varyingly developed. Sole without external layer
of flattened baskets. Panamic waters. . . ny ener
2. Tentacles with rosettes and delicate rods eae Rees narrow
rods; plates in sole with indented margin. Midventral band
with feet crowded. . . 2. Thyonepsolus beebei Deichmann
2. Tentacles without rosettes and delicate rods, only heavy plates.
Midventral band of feet with few feet. . . . . Sac
3. Dorsal side with few (7-8) scales and fairly numerous feet.
Hourglass-shaped bodies large, irregular. Sole with knobbed
plates. (Towers were not found in the type. )
3. Thyonepsolus veleronis, new species
138 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
3. Dorsal side with numerous scales (14 between oral and anal
scales) but comparatively few feet, easily overlooked. Dorsal
side with typical hourglasses of lacelike structure and large
towers which in older individuals become shortened, so that
they are thimble shaped. Sole with 4-holed buttons and plates
with marginal knobs; holes comparatively small. . F
4. Thyonepsolus hancocki, new species
1. Thyonepsolus nutriens H. L. Clark
Plate 26, Figs. 1-5
Thyonepsolus nutriens H. L. Clark, 1901, p. 169; 1901a, p. 491, text
figures.—Deichmann, 1930, p. 193 (passim); 1937, p. 174.
Psolidium nutriens Ludwig, 1904, p. 689.
Diagnosis—Small form (length of sole 2 cm.) ; distinct sole with
numerous feet in 3 crowded bands. Dorsal side with numerous small
scales, lacking in the midline (except in very young individuals), feet
numerous, and external layer of spicules well developed. Spicules in sole
perforated plates (0.10-0.15 mm.) and smaller flattened baskets (0.06
mm.) ; feet with end plate and supporting rods. Dorsal side with scales
and perforated plates (0.18 mm.) often slightly concave, and hourglass-
shaped bodies (0.10 mm.). In most individuals a number of huge tower-
like deposits (0.20-0.30 mm.) numerous in young individuals where the
other types of external spicules may be scarce. Dorsal feet with rudimen-
tary end plate and curved perforated plates or rods in the walls. Introvert
and tentacles with perforated plates or buttons and rods.
Type—M.C.Z.
Type locality—Pacific Grove, California, in tide pools.
Distribution —From the type locality southward, Santa Cruz Islands,
off Santa Barbara, California.
Depth.—Tide pools to few fms. depth.
Specimens examined.—The type and about 50 from various localities.
The following material examined from the Allan Hancock Expedi-
tions:
11-32. Conway Bay, Indefatigable Island, Galapagos, shore, January
12, 1932, 1 specimen.
Remarks.—A single specimen in the Allan Hancock Expeditions has
been identified with this species, but the label says Station 11-32(?) which
is Conway Bay, Indefatigable Island, Galapagos, shore.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 139
As this discontinued distribution seems at most unbelievable, it is for
the present assumed that the labeling is erroneous. The record is given
here in case future investigation should prove that it is correct and a
species closely related to T. nutriens, or identical with it, does occur in
the waters around Galapagos Islands. Comparison with material from
California has failed to reveal any tangible differences in the spicules.
The living animal (observed in Hopkins Marine Station) is brilliant
red with a more pinkish sole. The inner anatomy is similar to that of
Psolidium. The third mesentery is attached in the left ventral interambu-
lacrum, close to the midventral ambulacrum. The respiratory trees are
feebly branched and attached to the lateral interambulacra; the muscula-
ture is moderately developed, the retractors short and attached near the
oral end. The genital organs form short tubes in 2 tufts near the oral
end; they open on a papilla immediately behind the tentacles; small gen-
ital organs are present in specimens 0.6 mm. long. The eggs are huge,
about 1 mm. in diameter, and reddish gray, opaque. The eggs are trans-
ferred to the back of the female and lodged in pits; about 10 eggs are
hatched at a time; when ready to leave the mother in a few weeks, the
young are pale rose colored and have their full number of tentacles and 6
large tube feet. Specimens 2 mm. long have developed a distinct sole;
those 5 mm. long have 3 rows of feet on the ventral sole. In Monterey
Bay the female has been observed brooding its eggs from March to No-
vember; very likely it broods during the remaining winter months also.
2. Thyonepsolus beebei Deichmann
Plate 28, Figs. 1-3
Thyonepsolus beebei Deichmann, 1937, p. 172, text figure 3, 1-10; 1938,
pi dsZ:
Diagnosis —Small form (length of sole few cm.). Exterior typical of
the genus; dorsal side completely covered by scales. Ventral sole with
flat plates with scalloped edge; dorsal side with large complicated lacelike
hourglass-shaped bodies, towers and curved perforated plates; tube feet
with end plate and curved plates with 4 central holes and numerous mar-
ginal ones. Tentacles with heavy rods in stem; branches with delicate rods
and rosettes.
Type.—M.C.Z.
Type locality—Off Arena Bank, L. Calif., 2.5 fms.
Distribution.—West coast of Lower Calif., Mex., Costa Rica, pos-
sibly also Panama ( ?Lissothuria ornata Verrill, 1867, p. 322).
140 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Specimens examined.—F rom the following stations:
261-34. Tangola Tangola, Mexico, 15-20 fms., March 1, 1934, 1
specimen.
283-34. Thurloe Bay, West Coast of L. Calif., 8-10 fms., March 9,
1934, 3 specimens.
3. Thyonepsolus veleronis, new species
Plate 26, Figs. 6-8; Plate 27, Figs. 8-9
Diagnosis —Small form (type less than 2 cm. long) ; distinct sole
with feet in a double row along the edge and in a scattered row along the
midventral ambulacrum; dorsal side with comparatively few large scales
and relatively numerous feet; external layer of spicules well developed.
Spicules in sole knobbed plates; dorsally huge irregular hourglass-shaped
bodies, more cup shaped and less lacelike than in the other species. (Tow-
ers were not observed in the type.) Dorsal feet apparently without end
plate, walls supported by squarish curved plates of the typical structure.
Tentacles with heavy plates and rods with small holes; branches with
delicate rods or plates; apparently no rosettes.
Type.—Holotype, AHF no. 36.
Type locality—Station 147-34, Tagus Cove, Albemarle Island, Ga-
lapagos, dredging south of Cove, 30 fms., January 13, 1934.
Distribution —Type locality.
Depth.—Dredged from 30 fms.
Specimens examined.—The type.
Remarks.—The single individual measures 1.8 cm. in length (some-
what distorted). Its exterior and its spicules are very characteristic, and
it cannot be confused with any of the other species of Thyonepsolus. The
lack of towerlike deposits may be accidental—it is not uncommon in older
individuals of Thyonepsolus nutriens.
It is a question whether this species properly belongs in Thyonepsolus
or Psolidium, but the large hourglass-shaped spicules are so much more
reminiscent of those characteristic of Thyonepsolus than of the small cups
which are found in Psolidium that I have preferred at present, at least, to
refer it to the former genus.
4. Thyonepsolus hancocki, new species
Plate 27, Figs. 1-7
Diagnosis —Comparatively large form (sole up to 4 cm. long) ; sole
sharply set off; feet in a double row along the edge and in a scattered
NO.3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 141
band along the midventral ambulacrum. Dorsal side with numerous im-
bricating scales, about 14 between oral and anal apertures in the largest
individual. Dorsal feet few, rudimentary, easily overlooked.
Spicules in sole varying from 4-holed buttons to irregular, perforated
plates with marginal knobs and occasionally a few knobs near the center.
Feet with end plate and perforated rods and plates. Dorsal side with
scales, delicate hourglass-shaped cups of lacelike structure and huge tow-
ers which in the older individuals are partly resorbed from the tip, so
that they become shorter, thimblelike. Feet apparently without end plate;
walls supported by curved squarish plates with 4 central holes and a vary-
ing number of smaller marginal ones. Tentacles with heavy plates and
rods; in the branches delicate plates and rods; apparently no rosettes.
Type.—Holotype, AHF no. 37, 19 paratypes.
Type locality —Station 521-36, Agua Verde Bay, L. Calif., north of
anchorage, 5-10 fms., February 27, 1936.
Distribution —Gulf of Calif. to Ecuador.
Depth.—From 5-165 fms.
Specimens examined.—The following individuals from the Allan
Hancock Expeditions:
213-34. La Plata Island, Ecuador, dredging north of anchorage, rocky,
7-10 fms., February 10, 1934, 3 small specimens.
521-36. Agua Verde Bay, L. Calif., Mex., 5-10 fms., February 27,
1936, 5 small specimens.
529-36. Off San Francisquito Bay, L. Calif., Mex., 165 fms., March 1,
1936, 12 large specimens.
854-38. Gorgona Island, Colombia, north of Island, 40-60 fms., Febru-
ary 24, 1938, 7 large and 2 small specimens.
Remarks.—A well-defined species which is easily recognized.
Genus 2. PSOLIDIUM Ludwig, 1887
Diagnosis —Small forms (sole few cm. long) ; sole sharply set off;
with feet along the edges and few or many in the midventral ambulacrum ;
dorsal side with numerous imbricating scales, and a varying number
of tube feet which pass out through some of the scales. Spicules in sole
plates or buttons, smooth to knobbed, usually also an external layer of
small deep cups. Feet with end plate and supporting plates or rods. Dorsal
side usually with an external layer of cups (possibly lacking in some
species) ; besides grain or towers (more or less completely reduced with
142 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
age). Dorsal tube feet with or without end plate and supporting rods or
plates in the walls (the type species has squarish plates of similar type as
those in Thyonepsolus).
Type species—Psolidium dorsipes Ludwig.
Remarks.—The genus comprises a number of species. A few of these
have formerly been referred to Psolus because the few dorsal tube feet
were overlooked. On the other hand, it seems as if Psolidium convergens
Hérouard (1901, p. XXX) ought to be transferred to some other genus,
judging from the description given by Perrier (1905, p. 38, pl. 2, figs. 2-4,
text figures D-F).
The genus is closely related to Thyonepsolus H. L. Clark; actually
Ludwig considered (1904) the latter as a synonym of Psolidium. The
similarities between the 2 genera are even greater than assumed since it
has been discovered that the curved plates in the type species are partly
supporting plates for the dorsal tube feet and likewise that the character-
istic towers also may be present (see below). Nevertheless, it seems justi-
fiable for the present to retain the 2 genera on account of the more com-
plex hourglass-shaped bodies which are typical in Thyonepsolus, while
Psolidium has small cups or baskets.
Four species are discussed here, all represented in the Allan Hancock
collections and all taken at moderate depth or even in shallow water. One
is the type species, while 3 are described as new.
Key TO THE SPECIES OF Psolidium KNOWN FROM THE PANAMIC
REGION AND ADJACENT WATERS
1. ‘Tube feet fairly numerous in the midventral ambulacrum, also
numerous on the dorsal side where the scales are small and nu-
merous. Sole with plates with fairly large holes, smooth or
knobbed, and deeply convex cups. Dorsal side with numerous
deep cups and often also towerlike deposits; tube feet with
numerous curved squarish supporting plates, a few are narrow,
rod shaped. . . . . . . 1. Psolidium dorsipes Ludwig
1. Tube feet in midventral ambulacrum few. Dorsal feet
easily overlooked. ao ay yee Bai eeiy A Cae eg eee
2. Dorsal scales with a varying number of prominent pearllike
projections. Spicules in sole 4-holed buttons or plates with
knobbed surface and varying from delicate to stout. Oral
opening with large triangular scales; many small scales inter-
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 143
calated between the larger scales. y
3. Psolidium ekmani, new species
N
Dorsal scales smooth, without pearllike projections. . . . . 3
3. Dorsal side covered by numerous thimble-shaped grains with
flat base (reduced towers). Sole with small flattened cups with
lateral spines and a few larger plates with undulated surface
orsmooth. . . . . . 4. Psolidium planum, new species
3. Dorsal side with no accessory spicules (?). Ventral side with
heavy knobbed plates and buttons. . . . ... .
2. Psolidium eubullatum, new species
1. Psolidium dorsipes Ludwig
Plate 29, Figs. 1-12
Psolidium dorsipes Ludwig, 1887, p. 9, pl. 2, fig. 9; 1894, p. 135
(passim ).—Perrier, 1905, p. 48, pl. 2, figs. 5-15, text figures G-H.
—Ekman, 1925, p. 112, text figure 26.
Diagnosis—Small form (sole 1 cm. long) ; ventral sole sharply set
off with 3 crowded bands of tube feet. Dorsal side covered by imbricating
scales, about 10 between oral and anal scales; dorsal feet numerous, often
2-3 perforating a single scale; external layer of deposits well developed.
Spicules in sole plates varying from smooth to knobbed, 2 types of
plates, one with few holes, smooth or with few knobs, one with large holes
and numerous pearllike knobs; also an external layer of cups with 4 knobs
on the strongly convex side and a marginal brim of delicate teeth. Feet
with end plate and oblong supporting plates, slightly curved, often with
few knobs. Dorsal side with numerous baskets or cups of same size and
type as in the sole; tube feet mostly with small end plate and walls
packed with curved squarish plates, a few may be oblong. In some speci-
mens numerous towers are present.
Type.—Probably in Italy.
Type locality —Puntas Arenas, Strait of Magellan, 15-25 fms.
Distribution.—Strait of Magellan and vicinity. Also Panama and
Gulf of Calif.
Depth.—From 6-60 fms.
Specimens examined—The following from the Allan Hancock
Expeditions:
744-37. Near Point Piaxtla, Sinaloa, Gulf of Calif., 6-8 fms., April 1,
1937, 4 specimens (poor).
144 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
863-38. Bahia Honda, Panama, off North Island, 30-50 fms., March 1,
1938, 1 specimen (fair).
Remarks.—The Velero material differs from the material examined
by Ludwig and Perrier in the presence of the numerous towers with
broad smooth base. The cups or baskets agree well with Ludwig’s figure ;
possibly Perrier’s drawing is somewhat inaccurate; the marginal teeth
on the cups are too few and too coarse. It was a satisfaction to discover
that the tube feet actually were packed with the curved squarish plates as
was to be expected, not exclusively lying free in the skin as Perrier
describes it; a few simple curved oblong plates may be found in the wall
of the feet as in the other species.
2. Psolidium eubullatum, new species
Plate 28, Figs. 4-7
Diagnosis—Small form (type measures about 1 cm.). Sole with feet
restricted to the margin and the anterior end of the midventral ambu-
lacrum. Dorsal side with about 14 scales between oral and anal scales;
dorsal feet few.
Spicules in sole heavy buttons or plates with marginal and central
knobs. Feet with end plate and perforated plates. Dorsal side apparently
without any spicules except the scales; even the tube feet seem to lack
spicules entirely.
Type.—Holotype, AHF no. 38, 3 paratypes.
Type locality.—Station 213-34, La Plata Island, Ecuador, 7-10 fms.,
north of Anchorage, rocky, February 10, 1934, 3 specimens.
Distribution —Known from Ecuador.
Depth—From 7 to 55 fms.
Specimens examined.—The type and 3 specimens from the following
Velero stations:
212-34. La Plata Island, Ecuador, dredging 45-55 fms., sand shale,
rock, mud, February 10, 1934, 1 specimen.
213-34. La Plata Island, Ecuador, 7-10 fms., February 10, 1934, type
and 2 paratypes.
Remarks.——The specimens bear some resemblance to P. bullatum
Ohshima, (1915, p. 282) from the Alaskan waters, 54 fms., and may
possibly be found to be identical with that form. The unusually low
depth at which it is taken is noteworthy, and information about the hydro-
graphic conditions at La Plata Island is most desirable.
No.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 145
3. Psolidium ekmani, new species
Plate 30, Fig. 6
Diagnosis.—Small form (length of sole about 1 cm.). Oral and anal
openings closed by indistinct valves; stout radial teeth present inside the
oral plates; about 5 scales between the oral and anal valves. Scales
strongly sculptured with several pearllike projections, firmly attached;
dorsal tube feet few. Sole with rows of marginal feet and a row of few
feet scattered along the midventral ambulacrum. Spicules in sole delicate
4-holed buttons and plates with a larger number of holes; a varying num-
ber of knobs are present on the surface. Feet with end plate and support-
ing rods. Dorsal side with no other spicules than the scales (?).
Type.—Holotype, AHF no. 39, 1 paratype.
Type locality.—Station 213-34, La Plata Island, Ecuador, 7-10 fms.,
dredging north of anchorage, February 10, 1934.
Distribution—From Ecuador to Panama.
Depth.—From 7-10 fms.
Specimens examined.—The type and one other specimen from the
following Hancock stations:
213-34. La Plata Island, Ecuador, 7-10 fms., February 10, 1934, 1
specimen.
863-38. Bahia Honda, Panama, off North Island, rock, mud, sand, 30-
50 fms., March 1, 1938, 1 specimen.
Remarks.—The species was at first assumed to represent Psolus pata-
gonicus Ekman, but comparison with the latter showed clearly that the
spicules were entirely different (plate 30, fig. 8). Neither could it be
referred to Psolus squamatus (Koren), var. segregatus Perrier because
the projections on the scales were not loose but part of the scales. Also the
presence of tube feet in the midventral ambulacrum separates it distinctly
from both species mentioned above. Finally a few tube feet were dis-
covered on the dorsal scales and the generic position thereby settled. The
species is named in honor of Dr. Sven Ekman, the well-known zoogeog-
rapher and student of holothurians, from Upsala, Sweden.
4. Psolidium planum, new species
Plate 30, Figs. 1-5
Diagnosis—Small form (sole up to 1.5 cm. long), body capable of
becoming extremely flattened. Sole distinct; feet marginal and present as
a scattered row of much smaller appendages in the midventral ambu-
146 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
lacrum. Dorsal side with smooth imbricating scales, gradually decreasing
in size around the oral and anal openings; tube feet few, small, easily
overlooked. Entire dorsal surface covered by numerous large grains and
other deposits, easily rubbed off.
Spicules in sole an external layer of delicate smooth 4-holed flattened
cups with marginal teeth. Beneath these a few larger plates may be found
with several holes and smooth to undulating surface. Dorsal side closely
packed with thimble-shaped grains with flattened basal plate (i.e., short,
squat towers), besides numerous small baskets with delicate teeth and 4
apical knobs, sometimes united to a ring, thus approaching the hourglass
shape. Tube feet with delicate end plate and circular or squarish curved
supporting plates. (Tentacles not examined.)
Type.—Holotype, AHF no. 40.
Type locality —Station 845-38, Sechura Bay, Peru, Lat. 5° 39’ 30”
S.; Long. 81° 01’ W., 9.5 fms., February 15, 1938.
Distribution —From Peru to Colombia.
Depth—F rom 9-60 fms.
Specimens examined.—The following specimens from the Hancock
Expeditions:
845-38. Sechura Bay, Peru, coarse sand, worm tubes, red algae, 9.5 fms.,
February 15, 1938, 3 specimens.
854-38. Gorgona Island, Colombia, north of island, mud, rocks, 40-60
fms., February 24, 1938, 1 specimen.
Remarks.—The species has been referred to Psolidium because the
cups in the dorsal integument are mostly simple and the feet on the sole
are few. The dorsal spicules are easily rubbed off, but the spicules in the
sole are so characteristic that it seems impossible to confuse it with any
other species.
Genus 3. PSOLUS Oken, 1815
Diagnosis.—Typical psolids with distinct sole with marginal tube
feet, often also a band of feet along the midventral ambulacrum. Dorsal
side covered by scales, few to many; oral and anal apertures with or with-
out an operculum of triangular scales; sometimes narrow radial teeth
present inside of and between these valves. No appendages on dorsum
except the anal papillae and in some forms a few tube feet on the intro-
vert. Tentacles 10 in number, the 2 ventral may be smaller; in some
deepwater forms the tentacles are unbranched, finger shaped. Inner
No.3. DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 147
anatomy typical of the family; third loop of the intestine seems always
to be attached in the right ventral interambulacrum.
Spicules in sole plates or buttons, smooth or knobbed, in some forms
reduced with age, in others transformed into reticulated masses. Feet
usually with end plate and supporting rods. Dorsal side has besides the
scales often grains, rarely a few cups are present. Introvert and tentacles
with or without rods or plates; sometimes spicules reduced, in older in-
dividuals.
Type species.—Psolus phantapus (Strussenfeldt).
Remarks.—The genus comprises about 30 different species and repre-
sents apparently the most successful and widespread of the genera
included in the family. A number of the species are exceptionally large
(sole more than 10 cm. long). The members of the genus occur in tropical
as well as in Arctic seas.
The Hancock Expeditions have secured only one species, but 2 others
are discussed below to emphasize the differences between the species. One
represents a common form ranging from the Strait of Magellan to the
southern part of Bering Sea; the other, although hitherto known only
from the vicinity of Cape Horn, may possibly be expected in the southern-
most part of the Panamic region although it is not likely.
Key To THE SpEciEs OF Psolus KNOWN FROM THE PANAMIC REGION
AND WEstT CoasT OF SOUTH AMERICA
1. Dorsal scales large but few (2-3 between oral and anal valves).
Oral and anal valves large. Small form. Spicules in sole
knobbed plates. . . . . . 3. Psolus diomedeae Ludwig
1. Dorsal scales more numerous. . . . eo 2
Spicules in sole slightly hollow buttons is plates ete afl
holes and large marginal knobs. Distinct oral valves. . . .
; : . . . . 2. Psolus patagonicus Ekman
2. Seales in on flat, mostly 4-holed buttons reduced with age.
Dorsal side with few large round grains; oral valves indistinct.
. . 1. Psolus squamatus (Koren), var. segregatus Perrier
1. Psolus squamatus (Koren), var. segregatus Perrier
Plate 30, Fig. 7
Psolus squamatus (Koren), var. segregatus Perrier, 1905, p. 59.—
Ekman, 1925, p. 136, text figure 33.
148 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Psolus squamatus H. L. Clark, 1901, p. 165; 1901, p. 491.—Ohshima,
1915, p. 280.
Psolus pauper, Ludwig, 1894, p. 139.
Diagnosis —Large form (sole up to 13 cm. long); scales thin,
smoothly overlapping, about 12 between oral and anal scales. Oral and
anal apertures surrounded by a number of triangular scales but not de-
veloped as regular valves, except in very young individuals. Sole thin,
with marginal feet. Spicules in sole 4-holed delicate buttons (0.07-0.09
mm.) with shorter or longer marginal projections, sometimes with a few
low warts; often the spicules disappear with age. Feet with end plate and
few perforated rods. Dorsal scales covered by a varying number of loose
grains.
Type.—Paris.
Type locality.—Patagonia.
Distribution —From Cape Horn to the southern part of Bering Sea.
Depth.—In the Patagonian waters ranging from 7-207 fms.; in the
northern waters taken between 155-603 fms.
Specimens examined.—About 50 in the U.S.N.M. and Hopkins Ma-
rine Station from various 4/batross stations.
Remarks.—The specimens examined agree with Perrier’s form seg-
regatus from Cape Horn except in minor points (equal development of
the gonads and slightly different position of the paired retractors, prob-
ably because of a different degree of contraction). The material has been
compared with the typical form from Norway and shows the same differ-
ences as those which Ekman has tabulated.
Ludwig’s P. pauper from the west coast of Mexico represents a small
specimen of var. segregatus in which the spicules in the sole are unusually
scarce. Ludwig’s “antarcticus’ from off Peru, which has always been
regarded with suspicion, is most likely a small specimen of segregatus (or
a specimen of Psolus patagonicus Ekman or of Psolidium ekmani). ‘The
young individuals of P. sqguamatus, var. segregatus may superficially re-
semble either of these two forms.
2. Psolus patagonicus Ekman
Plate 30, Fig. 8
Psolus patagonicus Ekman, 1925, p. 140, text figures 35-36.
Diagnosis—Small form (sole up to 2 cm. long) with distinct oral
and anal valves and radial teeth inside the valves. Few scales between
oral and anal valves; often smaller scales intercalated between the pri-
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PARTI, DENDROCHIROTA 149
mary scales. External side of scales with pearllike projections, rare or
totally lacking in young individuals which may have a few loose grains.
Sole distinct with 1-2 marginal rows of feet. Spicules in sole slightly con-
cave 4-holed buttons (0.12 mm.) with relatively small holes and a
varying number of knobs on the surface, sometimes the knobs are united
by crossbars. Feet with end plate and perforated plates (0.18 mm.).
Tentacles with broad perforated plates.
Type.—Hamburg.
Type locality —Patagonian Bank, Lat. 46° S.
Distribution —The type locality and various localities in the Strait
of Magellan.
Specimens examined —Various individuals from Albatross Station
2777, and 2 received from the Museum in Buenos Aires.
Remarks.—The species is mentioned because it resembles Psolidium
ekmani. It was not secured by the Allan Hancock Expeditions and it
seems not likely that it will be found in the Panamic region.
3. Psolus diomedeae Ludwig
Plate 11, Figs. 9-10
Psolus diomedeae Ludwig, 1894, p. 136, pl. 14, figures 1-3.—Fisher,
1907, p. 717 (passim).
Diagnosis—Small form (sole less than 2 cm.) with enormous oral
and anal valves; radial teeth present inside to the valves. Dorsal scales
few and large, 2-3 between oral and anal valves; scales covered by few
but large grains; sole with marginal feet. Spicules in sole strongly
knobbed plates with 4 central holes and smaller marginal ones. (Feet and
tentacles not examined.)
Type.—U.S.N.M.
Type locality —Off Cocos Island, Costa Rica.
Distribution —From Gulf of California to Ecuador including Gala-
pagos.
Depth.—F rom 7-80 fms.
Specimens examined—The type in Washington and the following
specimens from the V’elero stations:
143-34. Wenman Island, Galapagos, 100-150 fms., January 11, 1934,
1 specimen.
213-34. La Plata Island, Ecuador, 7-10 fms., February 10, 1934, 38
specimens.
854-38.
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Octavia Bay, Colombia, north end of channel, coarse sand and
gravel, 35-40 fms., January 27, 1935, 1 specimen, young.
North of Octavia Bay, Colombia, south end of channel, 45
fms., January 27, 1935, 1 specimen.
Off San Francisquito Bay, Gulf of Calif., shale and gray mud,
165 fms., March 1, 1936, 1 specimen.
Chatham Bay, Cocos Island, Costa Rica, 40-47 fms., January
14, 1938, 3 specimens.
Off Daphne Minor Island, Galapagos, 70-80 fms., January 20,
1938, 1 specimen.
North of Gorgona Island, Colombia, 40-60 fms., February 24,
1938, 1 specimen, young.
Remarks.—The species is very striking and cannot be mistaken for
any other Pacific form, except the closely related P. macrolepis Fisher
from Hawaii, 154 fms. depth. The latter has almost smooth plates in the
sole and small basal scales intercalated between the oral valves. The
depth, 7-10 fms., for the specimens from La Plata Island, is unusually low
for this species and may indicate that peculiar hydrographic conditions
prevail in that locality.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 151
Ayres, W.
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EKMAN, S.
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FISHER, W.
1907.
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152 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Fores, E.
1841. A History of British Starfishes and other Animals of the Class Echino-
dermata, London, pp. I-XX, 1-267, with text figures.
Go.prFuss, G. A.
1820. Handbuch der Zoologie, Pt. 1-2, pp. I-XLVI, 1-696, pls. 1-4.
HEpInNG, S. G.
1936. Echinoderms, 6 & 7 Thule Expeditions. Medd. Gronland, vol. 108,
No. 1, pp. 1-34, text figures 1-6.
HEROUARD, E.
1901. Note preliminaire sur les Holothuries rapportées par |’Expedition
Antarctique Belge. Arch. Zool. Exp., notes 3, 9, pp. XXXIX-XLVIII.
JAcrr, G. F.
1833. De Holothuriis. (Dissertation Inauguration). Turin, pp. 1-40, pls. 1-3.
Koren, J.
Beskrivelser over Thyone fusus og Cuviera squamata. Nyt. Mag. f.
Naturvid., vol. 4, 1844-45, pp. 203-225, pls. 1-2.
LAMPERT, K.
1885. Die Seewalzen, Holothurioidea, eine systematische Monographie,
Wiesbaden, pp. 1-310, pl. 1.
Lesson, R. P.
1830. Centurie Zoologique, Paris, pp. 1-244, pls. 1-80.
Lupwice, H.
1875. Beitrage zur Kenntniss der Holothurien mit Nachtrag. Arbeiten aus d.
Zool.-Zoot. Institut in Wiirzburg, vol. 5, pp. 77-120, pls. 6-7.
1883. Verzeichniss der Holothurien des Kieler Museums. Berlin Oberh. Ges.,
vol. 22, pp. 155-176.
1887. Die von G. Chierchia auf der Fahrt der Kgl. Corvette “Vettor Pisani”
gesammelten Holothurien. Zool. Jahrb., vol. 2, pp. 1-36, pls. 1-2.
1887a. Drei Mittheilungen iiber alte und neue Holothurienarten. Sitzungsber.
Akad. Berlin, pp. 1217-1244 (reprint pp. 1-28), pl. 15.
1894. The Holothurioidea. Reports on an Exploration off the West Coasts of
Mexico, Central and South America, etc. Mem. Mus. Comp. Zool., vol.
17, No. 3, pp. 1-183, pls. 1-19.
1898. Holothurien. Hamburger Magelhaenische Sammelreise, pp. 1-98, pls.
1-3.
1904. Brutpflege bei Echinodermen. Zool. Jahrb., suppl. vol. 7, pp. 683-699.
MiiLLeR, J.
1854. Ueber den Bau der Echinodermen. Abhandl. Berlin Akad. Wiss., pl. 9,
fig. 9.
OHSHIMA, H.
1912. On the System of Phyllophorinae with Descriptions of the Species found
in Japan. Annot. Zool. Japan, Tokio, vol. 8, pp. 53-96, pl. 1, text figures
1-7.
1915. Report on the Holothurians collected by the U.S. Fisheries Steamer
Albatross in the Northeastern Pacific during the Summer of 1906. Proc.
U.S. Nat. Mus., vol. 48, pp. 213-291, pls. 8-11.
No.3 DEICHMANN: HOLOTHURIOIDEA; PARTI, DENDROCHIROTA 153
OKeEN, L.
1815. Lehrbuch der Naturgeschichte, vol. 3, Zoologie, Jena, pp. 351-352.
PALLAs, P. S.
1766. Zoophytorum sistens generum adumbrationes, etc., Hagae comitum,
pp. 1-451.
PERRIER, R.
1904. Holothuries du Cap Horn. Bull. Mus. d’Hist. Nat., vol. 10, pp. 11-17.
1905. Holothuries antarctiques du Museum d’Histoire Naturelle de Paris.
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Sars, M.
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SELENKA, E.
1867. Beitrage zur Anatomie und Systematik der Holothurien. Zeit. Wiss.
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SLuITER, C. P.
1880. Ueber einige neue Holothurien von der West-Kuste Javas. Naturkund.
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1886. Report on the Holothurioidea. Report on the Scientific Results of the
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TrRoscHEL, F. H.
1846. Neue Holothurien Gattungen. Arch. f. Naturgesch., vol. 12, pt. 1, pp.
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1866. On the Polyps and Echinoderms of New England. Proc. Boston Soc.
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1867. Supplementary note on the Echinoderms of the West Coast of America.
Ibid., p. 376. DS cesti
Fe.
154 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
EXPLANATIONS OF PLATES
Magnification: Scale divisions indicate 499 mm.
PLATE 10
Cucumaria cax;, new Species... ee p. 83
1. Supporting rod from tube foot.
2-5. Crackerlike button from integument.
Gucumaria ¢aliforntca Semper... 2 ee p. 79
6-7. Knobbed plate from integument.
8. Supporting rod from tube foot.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 10
156 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 11
Cucumaria dubiosa Sempet......-.-- ee ern ee p. 81
1. Knobbed plate with handle from integument.
2. Supporting rod from tube foot.
Cucumaria clilensis Ludwig. p. 80
3. Knobbed plate (medium sized) from integument.
4. Button from integument.
5. Supporting rod from tube foot.
VOL. 8
158 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 12
Pentamera. Geebet Teich asa aan nae nee p- 86
1-6. Supporting tables from feet.
7-9. Tables from integument.
DVep TT IIIA MORES LAL) OX CV 601510 0 renee Por p. 85
10-11. Supporting tables from tube foot.
12-17. Acorn-shaped bodies from integument.
VoL. 8
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA PL. 12
ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 13
Apentamerailepra; New? SpeCies ss. a eee p92
1-2. Supporting tables from tube foot.
3-4. Rosettes (or flat baskets) from outer layer of
spicules.
5-6. Knobbed plates from inner layer of spicules.
7-8. Tables from introvert.
9-10. Rod and plate from tentacles.
Neopentamera anexigua, NEW SPECIES........-....-----------c-n-eeeeeeeeeeeeeee=- p. 90
11. Supporting rods from tube foot.
12-14. Knobbed buttons from integument.
\Pentamera chiloensts (Ludwic). p. 88
15-17. Tables from integument, lateral view and from above
(spire omitted in the latter cases).
18. Supporting table from tube foot.
Penlamera chierchia (Ludwig) 22 =.= ie p. 86
19-20. ‘Tables from integument.
21. Supporting rod from tube foot.
4
tr
162 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 14
Lepiopentacta grisea WH, V.. Clark .22- 2 p- 93
1-2. Baskets from external layer of integument.
3-4. Buttons from external layer of integument.
5. Supporting plate from tube foot.
Leptopentacta pamammica, NEW SPECiES......----.2-------2------ceeeeeeeeneneeee p. 96
6-7. Baskets from external layer of integument.
8-10. Knobbed buttons from external layer of integument.
11. Supporting table from tube foot.
12. Small-holed plate with spinelike projection, from
external layer of integument or wall of tube foot.
Leptopemtacta 2OUG, WCW SPCClCS canon an ac a ones essences p. 95
13-15. Irregularly knobbed button from external layer of
integument.
16. Large swollen button from external layer of integu-
ment.
17-19. Biscuit-shaped plates from external layer of integu-
ment.
20-22. Biscuit-shaped plates with marginal tooth from ex-
ternal layer of integument.
O
eXe)
Oo
—
| OSS
oe
164 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
PLATE 15
Bepiopentacta nina, Qew SPeCles aan asco ceee p. 97
1-2. Hollow plates or buttons from external layer of
integument.
3. Larger plate from external layer of integument.
4-9. Irregular knobbed buttons from inner layer of integu-
ment.
10-12. Supporting plates from tube foot.
= ° : ; ;
eo ee SHS)
ax noe ro |
ne os @SS SRF Qs
\oR09'5
HO a 7 Ge
2 :
~~ c
166 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 16
Pestacta peruana (Sempe!) — p. 97
1-3. Baskets from external layer of integument.
4-6. Knobbed buttons from inner layer of integument.
7. Supporting plate from tube foot.
8. Knobbed plate from inner layer of integument near
base of tube foot.
168 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 17
Thyonacta mexicana, N€W SPECCICS............-.-..0----.c---ce--eeenencneeneneeee p. 101
1-5. Baskets from external layer of integument.
6-9. Smaller knobbed buttons from inner layer of integu-
ment.
10. Large knobbed button from inner layer of integu-
ment.
11-13. Supporting plates, knobbed and smooth, from tube
feet.
Ly
170 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 18
Thyone neosusus, New speCies.2.2..- oe p. 104
1. End plate from tube foot.
2-3. Tables from integument.
4-5. Supporting tables with no trace of spire.
6. Disk of table from introvert.
Thyone parapusus, new Species... 6: i p. 106
7-9. Oblong tables with tapering spire and “handle.”
10-11. Supporting tables with 3 pillars in spire.
12. Disk of table from introvert.
ihyonre pidentata; Rew Species.) ee Oe p. 105
13-14. Tables from integument.
15. Incipient table.
All from type.
16. Supporting table from a small individual less than
1 cm. long.
VOL. 8
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 18
172 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
PLATE 19
Thyone strangert, NEw SPCCles a eae p. 107
1-5. Tables from integument.
6-8. Tables from introvert.
9-10. Supporting tables with conical spire from tube foot.
11. Rosette from tentacle.
No. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 19
174 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 20
Weothyore givber (Selenka) 2. >. = p. 109
1-4. Knobbed to smooth buttons.
5-6. Knobbed button from external layer, lateral view
showing the external cluster of spines and the inner
“handle.”
7. Supporting table from tube foot.
Table from introvert.
9. Supporting rod from tentacle.
All from Selenka’s type in M.C.Z.
se
VOL. 8
E5083
: 0
= ;
176 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 21
Neothyone panamensis (Lud Wwig)....---------------------cscsesesesenenseeeeee p. 112
1-2. Supporting plates from ventral feet.
3-4. Knobbed plates from inner layer of integument.
5-6. Baskets derived from buttons from external layer of
integument.
Neothyone gibbosa, new Species. —...---a--2n00e0ec2sccenceeeeewenenesenaes p. 113
7-8. Knobbed buttons (outer handle often with 3 knobs)
from inner layer of integument.
9. Knobbed button from external layer of integument,
with a few spines (rare).
10-11. Supporting tables from tube foot.
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 21
178 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 22
Neothyone gibbosa, new SPeCies........------------:-------e---eeeeeeneneneeeeeeene p. 113
1-3. Large deformed supporting tables from dorsal feet
of large individual from San Felipe, Gulf of Cali-
fornia.
4. Basketlike buttons from outer layer of integument
(rare).
5-6. Knobbed buttons from inner layer of integument.
7. Small smooth supporting plate from ventral tube foot.
8. Table from introvert (disk obviously somewhat re-
duced through reduction along the margin).
Pachythyone pseudolugubris, new species............---------------------- p. 116
9-10. Supporting tables from tube foot, dorsal and ventral,
respectively, from large individuals from San Felipo,
Gulf of California.
VOL. 8
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 22
180 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 23
Pachythyone pseudolugubris, new SpeCieS.......--------------------------+ p. 116
1-2. Supporting tables from ventral tube feet of specimens
few cm. long.
3-5. Knobbed buttons from inner layer of integument.
6. Knobbed plate from outer layer seen from the inter-
nal side (the reticulated external layer is not drawn).
Pachythyone lugubris (Deichmann)............-.-----a------.0-------------- p. 116
7-8. Elongate knobbed buttons from inner layer of integu-
ment.
9. Supporting table from ventral tube foot.
Pachsihyoreattorg | (tle. Clark) eee ee p. 115
10. Supporting table from ventral tube foot.
11-12. Knobbed buttons from inner layer of integument.
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA PL. 23
182 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 24
Euthyonidium ovulum (Selenka) -.........--.-------------------eeesesnensoeeee p. 124
1-6. Plates with or without trace of spire.
7. Supporting rod from tube foot.
Euthyonidium veleronis, new SPeCis........-----------------0---see--eeneeoe= p. 126
8-10. Plates with few spires, indicating traces of a spire.
11. Plate from introvert.
Phyllophorus zacae Deichmann 23... p. 134
12-15. Disks of tables.
16-17. Tables, lateral view.
18. Disk of supporting table from tube foot.
19-20. Supporting tables, lateral view.
21. Rosette from tentacle.
VOL. 8
NO. 3 DEICHMANN: HOLOTHURIOIDEA; PART I, DENDROCHIROTA PL. 24
184
ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 25
Phyllophorus aculeatus Lud wigss.--< <2 ecenterecrte p. 133
1-3. ‘Tables seen from above.
4. Large supporting table from tube foot, lateral view.
5. Disk of supporting table, seen from above.
6. Disk of table from introvert.
7. Rosette from tentacle.
8. Delicate rods from tentacle.
VOL. 8
NO. 3 DEICHMANN: HOLOTHURIOIDEA ; PART I, DENDROCHIROTA PL. 25
186 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 26
Dagonepsovus nuiriens TH, Vi Clark 2002 oc eee p. 138
1. Curved supporting plate from dorsal tube foot.
2-3. Hourglass-shaped bodies from dorsal side.
4. End plate from dorsal tube foot.
5. Perforated plate from ventral sole.
Thyonepsolus veleronis, NEW SPECIES............00--c20---ce0-n-eeeeeseneeeeenee- p. 140
6-8. Knobbed plates from ventral sole.
VOL. 8
188 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 27
Thyonepsolus hancocki, new SpeCieS.......----------------1---0----eeeeneeoeo p. 140
1-2. Knobbed buttons from ventral sole.
3. Curved supporting plate from dorsal tube foot.
4-5. Hourglass-shaped bodies from dorsal side.
6-7. Perforated plates from stem of tentacle.
Thyonepsolus veleronis, NEW SPCCiCS.........-------------------eeeeneeeenneeee p. 140
8. Curved supporting plate from dorsal tube foot.
9. Hourglass-shaped body from dorsal side.
190 ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 28
Thyonepsolus beebet Deichmann_.._............----.-----0-----0----n-ne0e0---- p. 139
1. Perforated plate from ventral sole.
2. Curved supporting plate from dorsal side.
3. Hourglass-shaped body from dorsal side.
Psolidium eubullatum, new SspeCies.....--.---------------------+--eeseseeeee= p. 144
4-7. Knobbed plates from ventral sole.
VOL. 8
192 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 29
IP Sn eclititn OSSEPDES) TANIA eee p. 143
1-2. Knobbed and smooth plates from ventral sole.
3-4. Strongly knobbed plates from sole.
5. Supporting rod from ventral tube foot.
6. Towerlike body from dorsal side.
7. Curved supporting plate from dorsal tube foot.
8-10. que or baskets from external layer of sole and dorsal
side.
11. Narrow, curved supporting rod from dorsal tube foot
(rare).
12. End plate from dorsal tube foot.
29
194
ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 30
Psolidium planum, new SPpeCics.......--------------------n--neenenee ene p. 145
1-3. Shallow buttons or baskets from external layer in
sole.
4. Smooth plate in deeper layer of sole.
5. Curved plate from tube foot in dorsal.
Psolidium ekmant, new SPeCies.....----------2-s2n-a0--cee-nederecnnnnoeeessonne p. 145
6. Knobbed button from sole.
Psolus squamatus (Koren), var. segregatus Perrier-............. p. 147
7. Button from sole.
Psolus patagonicus Ekman..........--..-- Bo as tee Ne. ced a | p. 148
8. Slightly hollow button with marginal knobs from sole.
Psolwus: diomed eae Wud wits cece p. 149
9-10. Knobbed to almost smooth plates from sole.
VOL. 8
REPORTS ON THE COLLECTIONS OBTAINED BY ALLAN HANCOCK PACIFIC EXPEDITIONS OF
VELERO III OFF THE COAST OF MEXICO, CENTRAL AMERICA, SOUTH AMERICA,
AND GALAPAGOS ISLANDS IN 1932, IN 1933, IN 1934, IN 1935,
IN 1936, IN 1937, IN 1938, IN 1939, IN 1940, AND IN 1941.
NEW EASTERN PACIFIC SEA STARS
(PiaTes 31-34)
By FRED C. ZIESENHENNE
Tue UNIVERSITY OF SOUTHERN CALIFORNIA PUBLICATIONS
ALLAN Hancock PAcIFIC EXPEDITIONS
VOLUME 8, NUMBER 4
IsSUED OCTOBER 14, 1942
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
Los ANGELES, CALIFORNIA
CONTENTS
INTRODUCTORY REMARKS
Famity ASTROPECTINIDAE
Genus Leptychaster Smith
Subgenus Parastropecten Ludwig
Leptychaster (Parastropecten) stellatus, new species . ~. Pp. 197
Famity BENTHOPECTINIDAE
Genus Luidiaster Studer
Luidiaster californicus, new species . . «© «© + «+ Pp 201
Famity LINCKIIDAE
Genus Phataria Gray
Phataria mionactis, new species . . « +» + «+ « PP» 206
Genus Tamaria Gray
Tamaria obstipa, new species . . « + + + + «+ Pp 208
Famity ASTERIIDAE
Genus Coronaster Perrier
Coronaster marchenus, new species . . + + + «+ Pp 210
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NEW EASTERN PACIFIC SEA STARS
(PLATES 31-34)
Frep C. ZIESENHENNE
Research Assistant
Allan Hancock Foundation
During the last ten years the Velero III, research cruiser of the Allan
Hancock Foundation, The University of Southern California, on various
scientific cruises to the eastern tropical Pacific and southern California
waters has dredged an extensive collection of sea stars, including the fol-
lowing new species. In keeping with the policy of making expedition
results immediately available, it has been thought advisable to publish on
the new species rather than to await the final report on the Asteroidea.
The writer acknowledges his indebtedness to Dr. H. L. Clark of the
Museum of Comparative Zoélogy, Harvard College; to Mr. A. H. Clark
of the U. S. National Museum; and to Dr. W. K. Fisher of the Hopkins
Marine Station for valuable and helpful suggestions in the preparation
of the manuscript. Thanks are due Mr. Anker Petersen, staff artist of
the Allan Hancock Foundation, for the drawings and recording of color
notes, and Mr. G. P. Ashcraft of the Allan Hancock Foundation for the
accompanying photographs.
Last, but not least, the author expresses grateful appreciation to Cap-
tain Allan Hancock for the privilege of participating in most of the
Velero III expeditions and for the opportunity of a trip east to study and
compare many of the specimens with type material of the U. S. National
Museum and the Museum of Comparative Zodlogy.
Astropectinidae
Genus LEPT YCHASTER Smith
Subgenus PARASTROPECTEN Ludwig
Leptychaster (Parastropecten) stellatus, new species
Plate 31, Figs. 1-3
Description Rays 5, R=15 mm., r=6 mm., R=2.5 r. Width of
ray at second marginal plate 6 mm. Disk moderate, rays about as long
as width of disk, gradually tapering to a bluntly pointed tip ; general form
depressed. Abactinal paxillar areas not crowded, paxillae arranged in
[197]
198 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
regular, transverse, oblique rows at sides of ray and median interradial
area; no definite arrangement of paxillae in the center of disk. Paxillae
largest at base of ray and interradial areas, decreasing conspicuously in
size toward the center of disk and tip of ray. Column of paxilla slightly
higher than width of base, constricted at the summit. The largest bear 10
short spinelets constricted at the base and greatly enlarged into a finely
serrated sphere at the tip. Each paxilla bears 1 or 2 central spinelets and
up to 9 in the peripheral series ; smaller paxillae carry only 4 or 5 periph-
eral spinelets, not reduced in size.
Abactinal plates are small, compact along the midradial line and cen-
ter of the disk where papulae are lacking. The plates of the papular areas
in general are circular with 4 to 6 lobes, irregular in length, thickness, and
distribution. Plates of the papular areas are arranged in regular transverse
oblique rows, parallel with the interradial line. There are 6, often 5, and
occasionally 4 papulae around each plate, emerging from between the
lobes by which the plates usually touch. Plates along the median area of
the ray are even less regular, often without lobes; they are slightly small-
er, and the papulae are absent from a median strip about three plates
wide. Toward the center of the disk the plates rapidly decrease in size,
they become more crowded, and lobes if present are inconspicuous and of
irregular occurrence.
Superomarginal plates 18 from median interradial line to tip of ray,
forming an arched bevel to border the abactinal area. They are rectangu-
lar, with a convex surface, about three times wider than long, largest
interradially and gradually decreasing in size distally. Marginal plates
of both series are separated transversely by deep, narrow, fasciolar
grooves, and a deep longitudinal groove separates the supero- from the
inferomarginal plates. Superomarginal plates covered with numerous,
close-set, finely serrated capitate spinelets. The marginal spinelets are
slightly smaller and more delicate. The heavier, coarse, median spinelets
are arranged in 3 transverse rows on the interradial marginals, then 2
rows on median plates, and a single row on the most distal plates. They
are about the same size as the actinal, paxillar spinelets.
Terminal plate large, conspicuous, its length equal to the width of
four adjoining superomarginal plates; abactinal surface broadly convex;
proximal margin with a broad concave notch; distal margin blunt, with
a deep furrow groove. Distal half naked, proximal half covered with
capitate spinelets, which diminish gradually in size distally, where they
are only one third the size of those on the proximal margin.
NO. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 199
Inferomarginal plates 20, of the usual form, about as wide as length
of three plates interradially, whence the width rapidly diminishes toward
end of ray. The outer end of the first two plates is narrower than the
inner. The outer end of the plates, forming the rounded ventrolateral
border of the ray, is closely covered with rather truncated capitate spine-
lets similar to those of the superomarginals (but projecting a little farther
outward); but on the actinal surface these become slenderer, clavate,
with the thorny ends sometimes obliquely pressed, or bent, toward mar-
gin of ray. There are no enlarged tubercles on any marginal plate.
Actinal interradial areas small, comprising 2 complete chevrons of
plates. The inner chevron, without an odd interradial, extends to fourth
inferomarginal; the second, with a conspicuous unpaired interradial,
reaches to the second plate, while outside the odd plate are two others
indicating a third and fourth chevron to come. The plates have a low
tabulum, carrying a few capitate spinelets, like those on outer end of
inferomarginals, and also usually a few slender clavate ones.
Adambulacral plates 26, with an angular furrow margin, a convex
surface, and somewhat sunken transverse sutures. The first few plates
are wider than long, but they continue to narrow distally until on outer
third of ray they are longer than wide. Furrow spines proximally 4, but
over most of ray 3, subspatulate, blunt, the median member with edge to
furrow. There are proximally 9 or 10, distally about 6, clavate, thorny
subambulacral spinelets a little smaller than the furrow members, ar-
ranged in about 3 longiseries proximally and 2 distally. The distal mem-
ber of the longiseries often stands on the furrow margin and can reason-
ably be counted as a fourth or fifth furrow spinelet.
Mouth plates relatively large, the suture length about twice width of
combined plates. Furrow margin more than half suture length. Furrow
spines about 6, increasing in size toward apical spine, which is heavy,
clavate, and blunt. There are 10 or 12 suboral spinelets along margin of
median suture; 5 or 6 on margin adjacent to first adambulacral, and 4
or 5 scattered in between.
Madreporite oval, convex, diameter 1.25 mm., raised above disk
plates, located in the median interradial area along the superomarginals.
Striations coarse, irregular, centrifugal, arched domelike. Madreporite
surrounded by 6 large paxillae, which often conceal it.
Color.—Dried, actinal surface is cream buff, abactinal surface and
marginals yellow cream buff.
Holotype—AHF no. 44; 7 paratypes.
200 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Type locality—Holotype and 2 paratypes from Station 701-37, 32
fms., sand and shell, Angeles Bay, Gulf of California, Mexico, March 20,
1937. Four paratypes from Station 650-37, 47 fms., coarse sand, off San
Francisco Island, Gulf of California, Mexico, March 9, 1937. One para-
type from Station 1249-41, 34-44 fms., fine white sand, one mile south
of San Benito Islands, Lower California, Mexico, February 26, 1941.
Remarks.—The youngest specimen measures R=5.2 mm., r==3.2
mm., R=1.6 r, width at base of ray (second superomarginal plate) 2.7
mm. It has 7 very large, rectangular superomarginal plates separated
from one another by deep fasciolar grooves, and 14 adambulacral plates.
The terminal plate is conspicuous, large and convex, raised above the
abactinal surface. Anal aperture is barely visible. Paxillae bear a central
and 6 or 7 peripheral spinelets. There are 3 or 4 ambulacral furrow
spines. Each mouth plate bears 3 large oral spines. Actinal interradial
series consists of 7 plates. The madreporite is small, partly concealed by
crowded paxillae, and located in the median interradial area.
RELATIVE DIMENSIONS OF
Leptychaster (Parastropecten) stellatus
Width at No. of No. of
Station (am) (aun) R:r feed amas a
701-37 15 6 2524 6 18 26
701-37 7.6 3.8 a 3.4 14 18
701-37 10 4.2 Zot 4.2 16 23
1249-41 7.8 3.8 Page| 3.4 11 18
650-37 10 523 1:81 4.6 14 20
650-37 6 Sel 17 al 2.8 14
650-37 5:2 Sez ton 23. 14
650-37 5.4 Sel 1.7:1 ya | 14
This species is closely related to Leptychaster (Parastropecten) in-
ermis (Ludwig) of the tropical Pacific but differs from it in having 6
papulae instead of 4 around each plate; proximally 4, rarely 5, and dis-
tally 3 furrow spines instead of 6 or 7; a more stellate disk, longer arms,
No. 4
ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS
201
and very large, conspicuous terminal plate instead of a pentagonal disk
and small inconspicuous terminal plates.
It differs from the northern Pacific Leptychaster (Parastropecten)
anomalus Fisher by the following characters:
L. (P.) stellatus
Paxillar spinelets coarse, clavate.
Central paxillar spinelets 1 or 2.
Peripheral paxillar spinelets up-
ward of 9.
Ambulacral spines proximally 4,
rarely 5, distally 3.
Furrow margin of adambulacral
plate angular.
Actinal spinelets clavate, less nu-
merous.
Terminal plate large and conspicu-
ous.
Disk stellate.
Adambulacral plates shorter.
First adambulacral plate com-
pressed.
Adults have 18 superomarginals,
26 adambulacrals.
L. (P.) anomalus
Paxillar spinelets slender, terete.
Central paxillar spinelets 3 or 4.
Peripheral paxillar spinelets up-
ward of 10.
Ambulacral spines proximally 5 or
6, distally 4 or 5.
Furrow margin of adambulacral
plate convex.
Actinal spinelets slender, tapering,
numerous.
Terminal plate small and incon-
spicuous.
Disk pentagonal.
Adambulacral plates longer.
First adambulacral plate not com-
pressed.
Adults have 16 superomarginals,
29 adambulacrals.
The name ste//atus has reference to the long, narrow rays in contrast
to the usual short-rayed, pentagonal forms.
Benthopectinidae
Genus LUIDIASTER Studer
Luidiaster californicus, new species
Plate 31, Figs. 4-5; Plate 32, Figs. 1-7
Description.—Rays 5, R=87 mm., r=16 mm., R=5.43 r. Breadth
of ray at base 19 mm. Abactinal surface of the disk and basal portions of
the rays slightly swollen. Abactinal integument composed of large circu-
lar plates interspaced with numerous small, irregular, round and elon-
gated plates. A section of integument cleared in xylene revealed the plates
as irregularly arranged and close together, but not in actual contact. The
smaller plates serve as connectives between the larger ones, and the papu-
202 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
lae pass through the interspaces. As seen from the coelomic surface, on
the basal part of arm and adjacent disk integument, are 3 longitudinal
series of large plates, a median and 2 lateral rows, where the large plates
are compacted with fewer small connectives and no intervening papulae.
The plates are low and form 3 simple kinds of intergrading paxillae. The
tabulum is only slightly elevated, and the smallest plates bear from 1 to 8
short, bluntly pointed, granulelike spinules. The intermediate plates bear
one long, stout, tapering, movable spine and a surrounding ring of 1 to 5
sharp spinules of varying length and size, and on the base of the tabulum
a circle of short granulelike spinules. The largest of the intermediate
plates bear 1 to 3 equally long central spines and 1 to 6 slightly shorter
secondary spines. The largest paxillar plates confined to the disk and basal
ray usually bear a long, sharp, movable central spine and 4 to 9 equal or
subequal spines surrounding it. There is considerable variation as to the
length and density of these spines. These paxillae are free of secondary
spinelets and bear comparatively few granulelike spinules on the edge of
the tabulum. Dorsal spines are largest on the center of the disk, where
they are about 3 mm. long and fairly stout. The paxillar armature de-
creases rapidly in size distally and beyond the papular areas only the small
single-spinuled and granulelike spinule forms are present. The differences
between the large, small, and intermediate paxillae are more distinct on
the disk and papular areas of the rays than on nonpapular areas. The in-
termediate paxillar plates are in the process of forming large paxillar
plates bearing the long primary spines.
Marginal plates of two alternate series. There is no encroaching of
the superomarginals upon the abactinal area except far along the rays.
Each plate is strongly convex, almost tabulate, bearing 1 or 2, often 3,
movable tapering spines, the uppermost usually the largest, equal in length
to the combined length of three superomarginal plates. These are sur-
rounded by 2 to 7 secondary spines of varying length, less than one third
the length of the largest spine. On the remainder of the plate, except for
the naked border, are numerous, well-spaced, short, blunt, granulate
spinules. The plates of both series have their raised portion separated by
shallow, naked channels. The spines of the first and second superomar-
ginal plates are shorter than the succeeding ones.
Inferomarginal plates are slightly wider than superomarginals. The
central portion of each plate is elevated to form a transverse spine-bearing
ridge. Each ridge bears 2 or 3, often 4, rigid but movable, tapering,
pointed subequal, outstanding spines, the central or uppermost the largest.
No. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 203
They are stouter and somewhat longer than the corresponding supero-
marginal spines. There are fewer secondary spines and spinules compared
with the superomarginals. The terminal plate is small, wider than long,
densely covered with spinules.
Adambulacral plates more than twice as wide as long, widely sepa-
rated by deep, broad, naked grooves. The center of the plate is raised in a
prominent, transverse ridge bearing 3 or 4 subequal, stout, tapering spines,
the longest more than the combined length of two adambulacral plates.
The outermost spine slightly the longest. In addition, there may be from
3 to 9 sharply tapering spinelets on the outer edge of ridge. Furrow mar-
gin ridge prominent, bearing 2 to 4 subequal, heavy, blunt, slightly
curved, capitate furrow spines, the outer two shortest, middle one or pair
equal to the length of one adambulacral plate. ube feet in 2 rows; suck-
ing disks well developed, ampullae double.
Mouth plates oval shaped, more than twice as long as wide; actino-
stomial margin widest, broadly rounded; distal margin narrow, convex;
actinal surface highly convex. Marginal spines, 5 or 6, broad and flat-
tened, tapering and bluntly tipped. Innermost spine longest, about two
thirds the length of oral plate, next shorter, and the remaining diminish-
ing rapidly in size to the short, delicate outer spine. On the actinal sur-
face are 3 to 5 large, stout, tapering, blunt spines, half the length of oral
plate and 2 to 4 smaller secondary spines. Distally on each plate is a
longitudinal row of 3 to 8 short, sharp spinules. Actinal intermediate
plates 4 to 8, confined to the disk, oval shaped, wider than long, each bear-
ing 3 to 5 transverse rows of 3 to 4 short, sharp, granulelike spinules.
Papulae very numerous, absent in a narrow interradial line of the
disk and the distal two thirds of the ray. They are present on the disk and
basal portion of ray, where there is some semblance of 4 longitudinal rows,
each with an irregular transverse series of 3 to 6 papulae. Longitudinal
rows are separated by 3 longitudinal rows of loosely compacted, large
paxillar plates; the two central rows consist of 2 or 3 transverse papulae,
extending to the third or fourth superomarginal. Outer rows consist of
3 to 5 transverse papulae, extending to sixth, seventh, or eighth supero-
marginal; and in larger specimens there is a continuation of a single row
of papulae, close to the superomarginals extending to the thirteenth su-
peromarginal. In a specimen of R90 mm. each papular area contained
about 260 papulae. The youngest specimen of R32 mm. contained 62
papulae in each papular area.
204 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Pectinate pedicellariae very inconspicuous, flat, confined to the actinal
surface. One or 2 pedicellariae in each actinal interradial area, and up-
ward of 13 have been found on a ray between the adambulacral plates.
The margins of 2 adjacent plates each bear a comb of 5 to 7, straight,
broad, short, bluntly tipped, interlocking spinules. Pectinate pedicellariae
could not be found on all of the specimens.
Madreporite body convex, striae radiate out irregularly. It is located
in the interradius about its own diameter (1 or 2 plates) from the supero-
marginal plates; surrounded and protected by 4 or 5 large paxillae.
On the coelomic surface of the dorsal integument are 2 long, narrow,
strong muscle bands, extending from the basal portion to the tip of the
rays. Proximally they are separated, but on the distal third of ray they
join, continuing as a single band to the tip of ray. The proximal tendon is
from 8 to 10 mm. long. In the larger specimens the main portion of the
tendon is attached fanwise to the proximal side of the crest of the promi-
nent ninth or tenth ambulacral ossicle. The attachment is not noticeably
fanwise in the smaller specimens. Small lateral tendons branch off and are
attached to the third, fourth, fifth, and sometimes the sixth superomargi-
nal plate.
Color.—In alcohol, the ground color is dull cream buff, lightly over-
cast with gray, papular areas slightly darker gray; spines almost white
with tinges of cream buff.
Holotype—AHF no. 45; 7 paratypes.
Type locality—Holotype and 6 paratypes from Station 1306-41,
228-267 fms., loose rock, submarine bank 6 miles east of Long Point,
Santa Catalina Island, California, May 3, 1941. An additional paratype
from Station 1425-41, 280 fms., loose rock, 11.5 miles S.E. of Church
Rock, Santa Catalina Island, California, September 28, 1941.
Remarks.—There is some variation in the length and number of
spines on the paxillae; some bear secondary spines, some have just spi-
nules surrounding the large central spines, and others, mainly on the larg-
est specimens, have a central spine surrounded by equally large spines.
There are other variations already mentioned. The youngest specimen of
R=32 mm., r=5 mm., had no papulae in the center of the disk, the nar-
row interradial area, longitudinal center of the proximal part of ray, and
distal two thirds of ray. In each area there were about 62 papulae, which
extend out in 2 double rows to the fifth and a single row to the ninth su-
peromarginal. The armature of the plates and of the paxillae was es-
sentially the same as in the larger specimens. No pectinate pedicellariae
were observed.
NO. 4
ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS
205
From Luidiaster dawsoni (Verrill) of Alaskan waters this species
differs as follows:
Luidiaster dawsonti
Abactinal paxillae with short spines
surrounding the central spine.
Paxillar spinules long, sharp, dis-
tinct.
Large, prominent, circular pecti-
nate pedicellariae on actinal and
abactinal surfaces, composed of
2 or 3 combs of 12 or less long,
heavy, strongly recurved spin-
ules, almost L-shaped.
Adambulacral furrow spines 5-7.
Adambulacral actinal spines 2-4.
Actinostomial oral spines 6-8.
Suboral spines 7-10.
Muscle tendon attached to 7th or
8th ambulacral ossicle and 3rd
superomarginal.
Madreporite in young nearly
touching superomarginal.
Papulae absent from center of disk.
Luidiaster californicus
Abactinal paxillae with long spines,
surrounding the central spine
and often equal in length.
Paxillar spinules short, blunt,
granulelike.
Small, inconspicuous, flat pectinate
pedicellariae confined to actinal
surface, composed of 2 combs of
7 or less short, broad, blunt,
straight spinules.
Adambulacral furrow spines 3 or
4,
Adambulacral actinal spines 2 or 3,
rarely 4.
Actinostomial oral spines 5 or 6.
Suboral spines 3-5.
Muscle tendon attached fanwise to
9th or 10th ambulacral ossicle
and 2 or 3 lateral tendons to
3rd, 4th, 5th, or 6th superomar-
ginal.
Madreporite more than its diame-
ter from superomarginals.
Papulae present on center of disk,
except young.
This species is distinct from Lwidiaster horridus (Fisher) of the Ha-
waiian Islands as follows:
Luidiaster horridus
No pectinate pedicellariae.
Oral plates with 3 enlarged spin-
ules.
Luidiaster californicus
Pectinate pedicellariae present ac-
tinally.
Oral plates with 3-5 actinal promi-
nent spines.
206 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Inferomarginal spines 5-8. Inferomarginal spines 2 or 3, rare-
ly 4.
Paxillae with accessory spines two Paxillae with accessory spines often
thirds the length of central equal the length of central spine.
spine.
Actinostomial marginal spines 7. Actinostomial marginal spines 5 or
6.
Actinal oral spines 10 or 11. Actinal oral spines 3-5.
Papulae absent from center of disk. Papulae present on center of disk.
The name californicus has reference to the known distribution of this
species.
Linckiidae
Genus PHATARIA Gray
Phataria mionactis, new species
Plate 33, Figs. 3-4
Description.—Rays 5, R=50 mm., r=10 mm., R=5 r. Rays short,
heavy, and bluntly pointed, more or less cylindrical and tapering slightly
toward the tips. Height of ray at base 9.5 mm., at ray tip 6 mm. Width
of ray at base 11 mm., midray 10 mm., ray tip 8 mm. Skeleton composed
of numerous, irregular, swollen, compacted, granulated plates. The disk
is covered with large and small, irregular plates; the larger plates are ar-
ranged in an irregular circle within the-ray bases. Within the circle and
surrounding the anal opening are about 27 irregularly shaped plates of
various sizes. The plates are compactly covered with coarse, mosaiclike
granulations, which are round or rectangular. The granules surrounding
the anal aperture are heavier and longer.
The broad abactinal surface of the ray is covered with large and com-
pact plates of irregular shapes and sizes, between which are often wedged
smaller, irregular plates. Transversely there are about 5 plates across the
arm, irregularly placed and overlapping without a definite serial arrange-
ment. On the sides of the rays are 2 regular series of papular areas of
about 20 pores each. The upper series, immediately above the supero-
marginal plates, averages 17 areas and extends to within 8 mm. of the
ray tip. The lower or intermarginal series averages 22 areas and extends
to the tip of the ray. The papular areas, which are separated one from
another by a single transversely oriented plate, are reinforced by a reticu-
lation of tiny ossicles which serves to separate the papulae and support
the granules.
No. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 207
Marginals regularly arranged, lateral; the plates of the two series of
the same size, roughly rectangular, and united in dorsoventral pairs by
the secondary ossicle which separates the papular areas. The plates of
each series are strongly imbricated, free edge adoral, and they also overlap
the interpapular secondary plates; superomarginals about 18, inferomar-
ginals 20.
Actinal plates abruptly much smaller than inferomarginals, in 3 per-
fectly regular series, with the beginning of a fourth at interradius. ‘They
also form regular transverse series of 3, two of these corresponding to an
inferomarginal. Actinal granules become gradually coarser toward fur-
row.
Adambulacral plates compressed, somewhat crescent shaped, concave
distally, the actinal aspect about as broad as the furrow length of two suc-
cessive plates. Armature consists of a spatulate subtruncate furrow spine
with a shorter, much slenderer aboral companion; immediately behind
these (without intervening granules) stands a compressed, generally
abovate spine, about as long as the major furrow spine, and a secondary
aboral spinelet smaller than the corresponding one of the furrow series.
Back of these the plates are granulated as the adjacent actinals. Along
the margin of the extremely narrow furrow there are thus 2 series of
short, compressed spines (each with an inconspicuous companion).
Madreporite subcircular, slightly concave, 2 mm. in diameter, with
numerous small pores but no obvious carinae. It is in contact with 2 inter-
radial superomarginals.
Color.—Dried, light vinaceous buff. Color in life, abactinal surface
maroon with 2 purple longitudinal bands covering the papular areas, the
upper band terminating short of the ray tip and the lower band extend-
ing to the ray tip. Actinal surface creamish buff.
Type —AHF no. 41.
Type locality—Station 212-34, 45-50 fms., La Plata Island, Ecua-
dor, February 10, 1934.
Remarks.—The rays of this specimen are unusually short. This con-
dition might be caused by a pathological disease or mechanical injury.
Many coral-dwelling fishes, especially the parrot fishes, are known to bite
off sea star rays and even stout sea urchin spines. Upon examination ter-
minal constrictions were noted suggesting regenerating ray tips. Com-
pared with Pharia and Phataria, specimens in which ray tips were regen-
erating, the type differed by having short, blunt, and less tapering ray tips,
while the other specimens had longer, decidedly tapering ray tips termi-
208 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
nating in blunt points, and the regenerating ray was distinct from the
older portion.
The short, robust, nontapering blunt arms, 2 regular series of margi-
nal plates, 2 regular series of lateral papular areas, and the very broad
abactinal area distinguish this species from other members of the family
Linckiidae in the eastern tropical Pacific.
The type locality was again visited by the Velero III in 1935 and
1938 in the hope of collecting additional specimens of this unique species,
but none were obtained.
The name mionactis (less-ray) refers to the rays that are shorter
than usual.
Genus TAMARIA Gray
Tamaria obstipa, new species
Plate 33, Figs. 1-2; Plate 34, Fig. 1.
Description Rays 5, R=36 mm., r==7 mm., R=5 r. Rays slender
and tapering; width at base 8 mm., midray 6 mm., ray tip 3.5 mm. Disk
slightly convex, plates swollen, to one side of the central plate is the anal
aperture. The plates of ray are disposed in the following series: at base of
ray, 3 abactinal, 4 marginal, 6 actinal in addition to a series of secondary
intermarginals; at middle of ray, 2 actinal series less; near tip of ray, 1
abactinal (carinal), 4 marginal, 0 actinal. All of the plates are covered
with closely compacted, rounded or polygonal granules; those on the con-
vex surface of the plates are larger than those in the depressed areas be-
tween the plates. There is no definite arrangement of the granules, except
in the depressions between the plates where single, transverse rows often
mark the margins of the plates, leaving between them a noticeable trans-
verse furrow, especially distinct in the abactinal and marginal series.
The papular areas are arranged in 6 rows; 4 abactinal series extend
two thirds of the ray; they are not continuous; often there are 2 or 3
plates in succession without papular areas. Intermarginal series regular,
reaching within a few plates of ray tips. Papulae in each area vary from
6 to 24, distally some areas had as few as 3. There are no papular areas
on the actinal surface.
The adambulacral plates about three times as broad as long, about 3
plates opposing 2 of the adjoining series. Each plate bears a large sub-
ambulacral and 2 smaller furrow spines in addition to numerous enlarged
granules. The adambulacral armature consists of 2 distinct rows of spines,
set close together; those on the furrow are small, flat, blunt, chisellike,
NO. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 209
about as wide as long; the width is variable, often a broad spine opposing
a narrow spine. They are convex on the outer side and concave on the
furrow side, often with a narrow, distinct groove. The subambulacral
row is composed of large, robust, spatulate, blunt-tipped spines, longer
than wide, about four times as long and twice as wide as the furrow
spines. Between the subambulacral spines are slightly enlarged granules,
arranged in 1 or 2 series, transversely to the furrow. The mouth plates
bear a prominent, enlarged, triagonal, apical oral spine, 2 oral marginal
spines, and a prominent, spatulate suboral spine. Granulations on the
proximal half of mouth plates small; distal of the suboral spine they be-
come enlarged and spatulate.
Alveolate pedicellariae on the abactinal surfaces; although lacking on
the actinal surface of the holotype, they are abundant on the paratype.
The distribution of pedicellariae, however, is of no specific importance.
Abactinally the pedicellariae are well distributed ; actinally they are con-
fined to a single series of 7 to 14 along the subambulacral spines; in sev-
eral cases the pedicellariae have replaced the subambulacral spine on the
adambulacral plate. The shape of the jaws varies from slightly bent to
bifid ; they are thin, flaring widely at the tips and constricted at the base—
some flare out widely and are notched to form a letter Y, while others
flare asymmetrically to form a letter L. The majority resemble small
socks. The sockets in which the pedicellariae lie are straight ; the concavity
and broad ridge of the socket face the ambulacra.
The madreporite is 2.3 mm. in diameter and is located interradially
about its own diameter from the center of the disk. ‘The surface is covered
with fine, close convolutions and striations that radiate from the center.
The anal aperture in the center of the disk is closed by 6 long, narrow
spinelets, longer than wide and a number of elongated granules.
Color.—Dried from alcohol, light buff with a faint lavender hue.
Types.——Holotype AHF no. 42; one paratype.
Type locality —Station 780-38, 40-47 fms., Chatham Bay, Cocos Is-
land, Costa Rica, January 14, 1938. One paratype from Station 183-34,
50-70 fms., James Bay, James Island, Galapagos, January 24, 1934.
Remarks.—The paratype measures R=54 mm., r=10 mm., R=5 r.
It has fewer papular areas, which are not continuous but alternate with
the plates. The alveolate pedicellariae of the actinal surface are more
abundant, 7 to 14; abactinal surface, 12 to 15. The anal aperture is closed
by 8 small, triangular granules of different sizes. The groove on the fur-
row spines is deeper and conspicuous. 0 :
210 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
This species is closely related to J. floridae (Perrier), the pedicel-
lariae of which have small racquet-shaped jaws with a slender, long pedi-
cel and a broad distal expansion with smooth, convex margin. T. sclero-
derma (Fisher) is more distantly related. It has exceedingly tiny gran-
ules, difficult to detect until the specimen is dried. The subambulacral
spines are spaced from the furrow series, and the interval is thickly cov-
ered with minute granules. The proximal furrow spinelets of JT. sclero-
derma have a rudimentary groove or concavity on the furrow face.
The name odstipa has reference to the pedicellariae, which are bent
out of a straight line.
Asteriidae
Genus CORONASTER Perrier
Coronaster marchenus, new species
Plate 33, Figs. 5-6; Plate 34, Figs. 2-6
Description.—Rays 11 to 17, deciduous; R—80 mm., R=8 r.
Skeleton weak, reticulate, composed of numerous 3- to 5-lobed pri-
mary plates connected by 1 to 4 slender, imbricated ossicles. Reticulum of
disk irregular with much smaller meshes than that of the ray. These are
very small in a zone at periphery of disk, where the plates are irregular
and form a more solid framework. Between this peripheral frame and
the first transverse arch of abactinal ray plates is a line of weakness where
autotomy occurs. Primary plates of disk with a single acicular spine about
1.5 mm. long.
Skeleton of ray is composed of 5 radial rows of slender, 3- to 5-lobed
plates connected radially and transversely by narrow, elongated ossicles,
forming 4 rows of subrectangular papular areas. The carinal row consists
of 9 to 12 spine-bearing plates that extend to middle of ray, beyond which
degeneration takes place, leaving only the marginals. On the distal half
of ray gradual degeneration of longitudinally united secondaries has oc-
curred. The larger specimens have proximally a dorsolateral series con-
sisting of 5 to 8 spine-bearing plates confined to the basal third of ray.
The superomarginal and inferomarginal series consists of 20 to 25 spinif-
erous plates extending to ray tips. Each plate bears a large, acicular spine
about 3 mm. long.
The adambulacral plates are long, narrow, and set at a 45° angle to
the furrow; the distal half of one plate overlaps the proximal half of the
next plate on the furrow side. Each plate bears an oblique series of 2
NO. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 211
blunt, heavy, tapering spines with an enlarged bulbous base, inner spine
about 1.3 mm. long, outer spine 1.7 mm. long. Alternate adambulacral
plates carry on the furrow face a straight pedicellaria. ‘ube feet are
large, in 2 rows.
Mouth plates rectangular, longer than wide; each bears a large apical
oral spine and 2, sometimes 3, and often even a fourth oral spine. Inner
of furrow spines shortest, 1.3 mm. long; other spines larger, about 1.8
mm. long. These spines are flattened and widely flared out at the base,
then taper distally to a cylindrical blunt tip. Under high power all fur-
row spines are longitudinally ribbed and form a serrate edge on the spine
tip.
Papular areas of the disk are small and irregular, bearing a few pap-
ulae and often free rudimentary ossicles. There are from 2 to 15 papulae
in the large, rectangular-ray papular areas.
Straight pedicellariae lanceolate, slender, 0.4 to 1 mm. long, scattered
rather sparsely over plates, papular areas, and along the furrow face of
adambulacrals. The largest have 3 short, curved teeth at distal end, while
the smaller terminate in a short hook only. There are no hand-shaped
unguiculate pedicellariae such as are present in C. volsellatus, C. briareus,
and C. halicepus.
Crossed pedicellariae of the spine wreaths vary from 0.27 to 0.45 mm.
long. Each jaw is armed with 2 large terminal teeth between which are
6 small, sharp, triangular teeth about one half the length of the termi-
nals. These teeth are curved downward and form an acute angle with
the shank. The shank of the jaw is armed with a single series of 6 (vary-
ing from 5 to 7) long, sharp, downward-curved hyaline teeth. The ter-
minal teeth of the jaw give the pedicellariae the appearance of a minia-
ture saber-toothed tiger skull. The pedicellariae of the furrow are 0.4 mm.
long; distal four fifths of jaw is serrated and terminates in one or more
long, curved claws. They are found on alternate adambulacral plates.
Madreporites, 1 to 5, a large and 4 small accessory madreporites. ‘The
largest is 4 mm. long and 2 mm. wide; it is composed of 2 small
madreporites fused together. The 4 smaller madreporites are 1.5 mm. in
diameter. All five are located about their diameter from the edge of the
disk. The striations radiate irregularly from the center, except the large
madreporite, where they run parallel transversely to the long axis.
Color.—In life, abactinal surface creamish white with carmine stria-
tions running radially and transversely, more noticeable distally. The
radial and transverse rows of skeletal plates light gray, papulae and papu-
212 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
lar areas darker gray, wreaths of pedicellariae on spines commonly buff,
often varying from cream to a rich orange. Regenerating arms somewhat
lighter, salmon to orange in color. Actinal surface light gray, tube feet
pale yellow. When dried, ground color on abactinal surface of the disk
deep walnut brown; color extends on basal half of arm and gradually
fades distally to olive buff at tips; network of plates and spines olive-buff
with a tinge of brown. Actinal side similar in color but lighter in tone.
In alcohol, very pale purplish gray.
Holotype -—AHF no. 43; 33 paratypes.
Type locality Holotype and 10 paratypes from Station 310-35, 15
fms., tangles, Marchena Island, Galapagos, December 3, 1934. T'wenty-
three additional paratypes as follows: 2 from Station 307-35, 20 fms.,
tangles, Marchena Island, Galapagos, December 3, 1934; 3 from Station
308-35, 3-5 fms., sand, north bay, Marchena Island, Galapagos, Decem-
ber 3, 1934; 14 from Station 773-38, 31-50 fms., coralline bottom, off
Nuez Island, Cocos Island, Costa Rica, January 13, 1938; 3 from Station
779-38, 30-50 fms., rock, coral, and coralline, off Nuez Island, Cocos
Island, Costa Rica, January 14, 1938; and one from Station 795-38, 35-
40 fms., rock and sand, Sulivan Bay, James Island, Galapagos, January
21, 1938.
Remarks.—That this species is fissiparous is well borne out by a study
of the 34 specimens examined, in which 11 was the minimum and 17 the
maximum number of rays, 13 the average per individual. Specimens with
a ray length less than 50 mm. averaged 12.7 rays, while those exceeding
50 mm. ray length averaged 13.6 rays. Exclusive of the holotype, 49.3
per cent of the rays were young or regenerating rays. Madreporites ranged
from 1 to 5 per individual, 3 being the average ; 24 per cent of the madre-
porites were on young or regenerating portions of disk. In fissiparous re-
production a portion of the disk is cast away with the rays. The small
madreporites and regenerating rays grow from this section of regenerating
disk, which in some specimens is very distinct. There is no definite order
or arrangement in the manner the new rays are added to the disk. In gen-
eral, 5 or more rays of a first set grow from the regenerating disk; then
at a later period a few scattered rays of a second set appear between the
adult rays. In some specimens a third set of rays appears between the re-
generating first set just opposite the old adult rays. The average ray
length to the disk radius for 20 of the largest specimens is R=9.5 r.
These are the first Coronasters to be recorded from the eastern tropi-
cal Pacific Ocean and increase the number of Pacific species to 4: Coron-
NO. 4 ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS 213
aster volsellatus (Sladen) and Coronaster halicepus Fisher, Philippine
Islands, and Coronaster eclipes Fisher, Hawaiian Islands.
Marchenus, having 2 adambulacral spines, cannot be confused with
volsellatus. It differs from halicepus as well as briareus of the West Indies
in lacking broad, unguiculate, hand-shaped, straight pedicellariae. In C.
eclipes (described from a single ray) these unguiculate pedicellariae are
also lacking; and the lanceolate type, so characteristic of marchenus, 1s
found only on the furrow face of the adambulacrals. But the furrow
pedicellariae of marchenus are similar to those of volsel/atus and quite un-
like the smaller more triangular ones of eclipes (see Fisher, Sea Stars
Tropical Central Pacific, Bishop Museum Bulletin No. 27, 1925, p. 78,
fig. 9 d, e, £). The crossed pedicellariae are essentially the same as those
of volsellatus, with a similar wide range in size. Characteristic of mar-
chenus are the fissiparous habit and the presence of upward of 5 madre-
porites, features which have not been reported for the other species.
The name marchenus has reference to Marchena Island, on whose
rough lava reefs this species was taken in abundance.
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216 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 31
Leptychaster (Parastropecten) stellatus, new species
Fic. 1. Abactinal view of paxillae and mar-
ginal plates x 10.
Fic. 2. Abactinal view, scale in mm.
Fic. 3. Actinal view, scale in mm.
Luidiaster californicus, new species
Fic. 4. Abactinal view.
Fic. 5.
t
aS
3? ,
res
Fic.
Fic.
Fic.
Fic.
FIG.
Fic.
ALLAN HANCOCK PACIFIC EXPEDITIONS
Ww bo
6.
PAP ATE E32
Luidiaster californicus, new species
Abactinal view of dissection showing the muscle
attachments of the dorsal integument to the 9th
and 10th ambulacral ossicles and the side wall.
Side view of the adambulacral armature.
Side view of the most advanced and largest type
of abactinal paxilla.
Actinal view of the dorsal integument, upper
part cut from the center of the disk and lower
part from the proximal portion of the arm,
showing large and small skeletal plates and
papulae distribution.
Pectinate pedicellaria from the actinal inter-
radia area.
Side-actinal view of upper and lower marginal
plates.
Actinal view of mouth plates showing arma-
ture.
VOL. 8
RN PACIFIC SEA STARS
EASTE
NEW
ZIESENHENNE:
No. +
220 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLEAD E 33
Tamaria obstipa, new species
Fic. 1. Abactinal view.
Fic. 2. Actinal view.
Phataria mionactis, new species
Fic. 3. Abactinal view.
Fic. 4. Actinal view.
Coronaster marchenus, new species
Fic. 5. Abactinal view.
Fic. 6. Actinal view.
NO. + ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS
3 J 3
Tim TT TTT
HG:
FIG.
Fic.
FIG.
Fic.
FIG.
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
RIVAGE 35,
Tamartia obstipa, new species
Actinal view showing adambulacral armature
and three pedicellariae.
Coronaster marchenus, new species
Large, straight abactinal pedicellaria.
Small, straight abactinal pedicellaria.
Ambulacral furrow pedicellaria.
End view of a crossed pedicellaria showing ar-
rangement of teeth.
Side view of a crossed pedicellaria.
No. + ZIESENHENNE: NEW EASTERN PACIFIC SEA STARS pL. 34
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HUBERT LYMAN CLARK: TEACHER AND FRIEND
January 9, 1870 - July 31, 1947
Probably the two forms of activity which Dr. Hubert Lyman Clark
enjoyed above all others were teaching and dredging for marine animals.
It is perhaps appropriate, therefore, that this brief note of appreciation
should be offered by one of the very few students Dr. Clark was destined
to have during his long tenure at Harvard University and published
through the generosity of Captain Allan Hancock, who made possible Dr.
Clark’s many happy days of dredging aboard the Velero III. No attempt
is made to incorporate in this tribute an evaluation of his important contri-
il
butions to the study of echinoderms, indicated by the bibliography which
follows; that appraisal should be reserved for other workers in his chosen
field.
My association with Dr. Clark began in 1930 when he agreed to
supervise my graduate studies on Crustacea. At the time, my knowledge
of Crustacea was extremely fragmentary, and he readily admitted his
ignorance of some of the more involved aspects of crustacean classification.
Thanks to his insistence that the best way to learn the relationships of any
group was by direct examination of as many of the representative forms
as possible, both of us eventually gained an adequate working knowledge
of this diverse assemblage of animals. As this form of training entailed the
drudgery of checking identifications, relabeling specimens, and_ re-
arranging the crustacean collections in the far reaches of the catacomb-like
basement of the Museum of Comparative Zodlogy, the assignment was
not consistently welcomed or understood by a young taxonomist who
associated systematic work with somewhat more significant tasks in a less
obscure environment. Not until several years later did the value of this
fundamental curatorial training become apparent to the recipient. I am
sure that Dr. Clark appreciated his student’s mild discomfort, and I will
always be grateful that he had the courage of conviction to see to it that
this apprenticeship was fully served.
In subsequent years, as a member of the staff at the Museum of
Comparative Zodlogy, I became better acquainted with Dr. Clark and
came to recognize his loyal friendship as one of the most important and
valued associations of my life. Although he was unusually reticent about
offering unsought criticism of any kind, he was always eager to be of
assistance whenever his advice was solicited. His almost emotional grati-
tude for all favors returned in kind was a unique characteristic familiar
to all who came in close contact with him.
Except for an attack of yellow fever suffered during a collecting
trip to Jamaica in 1896, from which he was the only one of six victims
to survive, Dr. Clark enjoyed remarkably good health until his last illness.
Always one to minimize his infrequent minor indispositions, he sometimes
found it difficult to appreciate the reaction of others to similar disorders.
The bout with yellow fever had a profound effect upon his later life,
however, for it left him with impaired hearing which showed progressive
deterioraion with the passage of time. This affliction seemed to affect in
no way his optimistic outlook toward life, but it did virtually force his
retirement from classroom teaching, which he so enjoyed, and prevented
iii
intimate contact with his fellow men; this, to a man of Dr. Clark’s
friendly disposition, must have been a heavy cross to bear, but I cannot
recall that he ever referred to it with the least trace of bitterness.
His unusual robust health was reflected in his general mental outlook.
Whether the activity involved was his scientific work, caring for his
valuable stamp collection, playing tennis, or climbing his beloved New
Hampshire mountains, all of which were indulged in until the last year
of his life, Dr. Clark believed in giving the best that was in him. The only
times that I have seen him violently incensed about anything were on
those infrequent occasions when he discovered a typographical error he
had overlooked in one of his publications or lost a point at tennis which he
felt should have been his with a little additional skill or effort, and at
these times his anger was always directed at himself. In his scientific work,
he was prone to set goals and deadlines for himself and he did his best to
adhere to these even when it meant driving himself far into the night.
This is a characteristic which may seem questionable to others of us
acquainted with the unpredictability of taxonomic research, but it was a
fundamental part of Dr. Clark’s nature and one which apparently served
him well.
No insight into Hubert Lyman Clark’s character would be complete
without reference to his religious activities. An ardent churchman, he
thoroughly believed that the Sabbath was a day to be set aside for worship
and human companionship. All who accompanied him on field trips will
recall how determined he was that Sunday, as far as he was concerned,
was not a day for collecting, no matter how propitious the weather or the
tide.
This religious nature was at the root of his entire philosophy of life.
As he once expressed himself in print, he put his faith in a “confident
belief in oneself, in a loving God and in our fellow men as children of one
Father.” Only on some such foundation as this did he feel that one could
build a satisfying personal philosophy which, in his words, “‘must give full
scope to the mind, .. . set no arbitrary limits to human knowledge, .. .
encourage and stimulate the quest for truth and have no fear thereof
wherever found, . .. allow full play for the emotions, ... [and]... provide
a fundamental motive, a driving power for an achieving life.”
FENNER A. CHACE, JR.
U.S. National Museum
July 16, 1948
1896.,
1897.
1898.
1899.
BIBLIOGRAPHY OF Dr. H. L. CLark’s ECHINODERM PAPERS
Compiled by Marjorie Pattee, Research Assistant, M.C.Z.
Notes on the life history of Syxapta vivipara Oerstedt. Jour. Jamaica Inst.
July, vol. 2, pp. 278-282.
The viviparous Synapta of the West Indies. Zool. Anz., September 14, vol.
19, no. 512, pp. 398-400.
The viviparous Syzapta of the West Indies. Johns Hopkins Univ. Circulars,
November, vol. 17, pp. 2-3.
Notes on the echinoderms of Bermuda. Ann. N. Y. Acad. Sci., September,
vol. 11, pp. 407-413.
Synapta vivipara: a contribution to the morphology of echinoderms. Mem.
Boston Soc. Nat. Hist., vol. 5, no. 3, pp. 53-88, pls. 11-15.
Synapta vivipara: a contribution to the morphology of echinoderms. Mem.
Biol. Lab. Johns Hopkins Univ., vol. 4, no. 2, pp. 53-88.
The echinoids and asteroids of Jamaica. Johns Hopkins Univ. Circulars,
November, no. 137, pp. 4-6.
Further notes on the echinoderms of Bermuda. Ann. N. Y. Acad. Sci., vol.
12, pp. 117-138, pl. 4.
The Synaptas of the New England coast. U.S. Fish. Comm. Bull., December
~ 13, pp. 21-31, pls. 10, 11.
1901.,
Bermudan echinoderms. A report on observations and collections made in
1899. Proc. Boston Soc. Nat. Hist., vol. 29, no. 16, pp. 339-345.
The echinoderms of Porto Rico. U.S. Fish Comm. Bull., vol. 2, pt. 2, pp.
231-263, pls. 14-17.
_ The holothurians of the Pacific Coast of North America. Zool. Anz., March
25, vol. 24, no. 639, pp. 162-171.
Synopses of North American invertebrates: The holothurioidea. Am. Nat.,
June, vol. 35, pp. 479-496.
The breeding habits of holothurians. Mich. Acad. Sci. Third Rep., pp. 83-85.
_ Echinoderms from Puget Sound. Proc. Boston Soc. Nat. Hist., vol. 29, no.
1902.
1904.
1905.
1906.
1907.,
15, pp. 323-337, 4 pls.
Notes on some North Pacific holothurians. Zool. Anz., July, vol. 25, pp.
562-564.
Echinodermata papers from the Hopkins Stanford Galapagos Expedition,
1898-1899. Proc. Wash. Acad. Sci., September 30, vol. 4, pp. 521-531.
Echinoderms of Woods Hole region. U.S. Fish Comm. Bull. for 1902, pp.
545-576, pls. 1-14.
Fauna of New England: List of the Echinodermata. Occ. Papers, Boston
Soc. Nat. Hist., February, vol. 7, pp. 1-13.
List of echinoderms available for exchange in the M.C.Z. September, pp. 1-8.
With A. Agassiz. Hawaiian and other Pacific Echini. The Cidaridae. Mem.
M.C.Z., February, vol. 34, no. 1, 50 pp., 44 pls.
With A. Agassiz. Preliminary report on the Echini collected, in 1902,
among the Hawaiian Islands, by the U.S. Fish Commission Steamer
“Albatross.” Bull. M.C.Z., March, vol. 50, no. 8, pp. 231-259.
__The starfishes of the genus Heliaster. Bull. M.C.Z., vol. 51, no. 2, pp. 25-76,
8 pls.
Recent literature on echinoderms. Science, July, n. s. vol. 26, pp. 12-17.
With A. Agassiz. Preliminary report on the Echini collected in 1906, from
May to December, among the Aleutian Islands, in Bering Sea, and along
the coasts of Kamtchatka, Sakhalin, Korea, and Japan, by U.S. Fish Com-
mission Steamer “Albatross.” Bull. M.C.Z., October, vol. 51, no. 5, pp.
109-139.
As to holothuria. Science, October, n. s. vol. 26, pp. 549-550.
The Cidaridae. Bull. M.C.Z., December, vol. 51, no. 7, pp. 165-230, 11 pls.
The apodous holothurians. Washington, January 1908, Smithsonian Cont.
to Knowl., vol. 35, 231 pp., 13 pls.
Vv
1908.
1909.
1910.
1911.
1912,
1913.
1914.
A brittle star new to the Woods Hole region. Science, January, n. s. vol. 27,
. 147.
Seite Japanese and East Indian echinoderms. Bull. M.C.Z., April, vol. 51,
no. 11, pp. 279-311.
Renewed interest in recent crinoids. Am. Nat., May, vol. 42, pp. 350-355.
The type of Cidaris. Ann. Mag. Nat. Hist., June, ser. 8, vol. 1, pp. 532-533.
With A. Agassiz. Hawaiian and other Pacific Echini: The Salenidae,
Arbaciadae, Aspidodiadematidae, and Diadematidae. Mem. M.C.Z.,
September, vol. 34, no. 2, pp. 43-132, 17 pls.
Les Cidaridae. Bull. Soc. Sci. Nat. Haute-Marne, 5¢ Ann., no. 22, pp.
165-228.
Abstract of unpublished paper: Brood-protection and sexual dimorphism
among echinoderms. Science, n. s. vol. 29, p. 711.
With F. A. Bather. The type of Cidaris. Ann. Mag. Nat. Hist., January,
ser. 8, vol. 3, p. 88.
Notes on some Australian and Indo-Pacific echinoderms. Bull. M.C.Z.,
March, vol. 52, no. 7, pp. 109-135.
Scientific results of the trawling expedition of H.M.C.S. “Thetis” off the
coast of New South Wales, in February and March 1898. Echinodermata.
Australian Museum Mem. 4, pt. 11, pp. 519-564, 11 pls.
With A. Agassiz. Hawaiian and other Pacific Echini. The Echinothuridae.
Mem. M.C.Z., November, vol. 34, no. 3, pp. 135-206, pls. 60-89.
A new ophiuran of the West Indies. Proc. U.S. Nat. Mus., August 19, vol.
37, pp. 665-666.
The echinoderms of Peru. Bull. M.C.Z., October 10, vol. 52, no. 17, pp.
321-358, 14 pls.
The development of an apodous holothurian. (Chiridota rotifera). Jour.
Exp. Zool., November, vol. 9, pp. 497-516, pls. 1, 2.
North Pacific ophiurans in the collection of the U.S. National Museum. U.S.
Nat. Mus. Bull. 75, pp. i-xvi, 1-302, figs. 1-144.
The genera of recent Clypeastroids. Ann. Mag. Nat. Hist., ser. 8, vol. 7, pp.
593-605.
The purpose and some principles of systematic zoology. Pop. Sci. Monthly,
September, vol. 79, pp. 261-271.
Fossil holothurians. Science, February 16, n. s. vol. 35, pp. 274-278.
_Biotypes and phylogeny. Am. Nat., March, vol. 46, pp. 139-150.
Los equinodermos del Peru. Anales de la Direccion de Fomento, no. 5, May,
pp. 22-72. Traduccion del Boletin del Museo de Zoologia comparado del
Harvard College, vol. 52, no. 17, pp. 321-358, 14 pls.
Review of Jackson’s “Phylogeny of the Echini.” Science, June 28, n.s.
vol. 35, pp. 986-993.
Hawaiian and other Pacific Echini. The Pedinidae, Phymosomatidae,
Stomopneustidae, Echinidae, Temnopleuridae, Strongylocentrotidae, and
Echinometridae. Mem. M.C.Z., June, vol, 34, no. 4, pp. 205-383, pls. 90-121.
Investigations on echinoderms at Montego Bay. Carnegie Inst., Washington,
Yearbook 11, pp. 134-135.
Autotomy in Linckia. Zool. Anz., June, vol. 42, pp. 156-159.
Echinoderms from Lower California, with descriptions of new species. Bull.
Am. Mus. Nat. Hist., July, vol. 32, pp. 185-236.
Asterozoa (and) Holothurioidea. Zittel’s textbook of paleontology. C. R.
Eastman, editor, second edition, December, vol. 1, pp. 244-257, 312-313.
Growth-changes in brittle stars. Carnegie Inst., Washington, Publ. 182
(March), pp. 91-126, 3 pls.
Hawaiian and other Pacific Echini. The Clypeastridae, Arachnoididae,
Laganidae, Fibulariidae, and Scutellidae. Mem. M.C.Z., June, vol. 46, no. 1,
80 pp., 22 pls.
The echinoderms of the Western Australian Museum. Rec. W. Austr. Mus.
and Art Gall., vol. 1, pp. 132-173, pls. 17-26.
vi
1915.
Review of Verrill’s “Shallow-water star fishes of the North Pacific coast.”
Science, October 9, n. s. vol. 40, pp. 523-525.
The classification of the regular Echini. Zool. Anz., December 4, vol. 45,
pp. 171-172.
Report on the work at the Torres Strait Laboratory. Carnegie Inst., Wash-
ington, February, Yearbook 13 (for 1914), pp. 200-201.
A remarkable new brittle star. Jour. Ent. and Zool., March, vol. 7, pp. 64-
66.
The comatulids of Torres Strait: with special reference to their habits and
~ reactions. Carnegie Inst., Washington, Publ. 212, May, vol. 8, pp. 97-125.
The echinoderms of Ceylon other than holothurians. Spolia Zeylanica,
~ October, vol. 10, pp. 83-102.
_-Catalogue of recent ophiurans: based on the collection of the Museum of
1916.
19U7.
1918.
1919.
Comparative Zoology. Mem. M.C.Z., December, vol. 25, no. 4, 214 pp., 20
pls.
Report on the sea lilies, starfishes, brittle stars, and sea-urchins obtained by
the F. I. S. “Endeavour” on the coasts of Queensland, New South Wales,
Tasmania, Victoria, South Australia, and Western Australia. Biol. Results
“Endeavour,” 1909-14, June 2, vol. 4, pp. 1-123, pls. 1-44.
Report on studies at Tobago, British West Indies. Carnegie Inst., Washing-
ton, Yearbook 15, February 15, pp. 192-193.
Hawaiian and other Pacific Echini. The Echinoneidae, Nucleolitidae,
Urechinidae, Echinocorythidae, Calymnidae, Pourtalesiidae, Palaeostomati-
dae, Aeropsidae, Palaeopneustidae, Hemiasteridae, and Spatangidae. Mem.
M.C.Z., vol. 46, March, no. 2, 203 pp., 18 pls.
The habits and reactions of a comatulid, Trofiometra carinata. Carnegie
Inst., Washington, Publ. 251 August, pp. 111-119.
Reports on the scientific results of the expedition to the Tropical Pacific,
1899-1900. XVIII. Reports on the scientific results of the expedition to the
Eastern Tropical Pacific 1904-1905. XXX. Ophiuroidea. Bull. M.C.Z.,
October, vol. 61, no. 12, 28 pp., 5 pls.
Report on the Crinoidea and Echinoidea collected by the Bahama
Expedition from the University of Iowa in 1893. Bull. Lab. Nat. Hist. Univ.
Iowa, April, vol. 7, no. 5, pp. 3-37, pls. 1-5.
Brittle stars, new and old. Bull. M.C.Z., October, vol. 62, no. 265-338, 8 pls.
Review of Walter Fisher’s “Starfishes of the Philippine seas and adjacent
waters.” Science, October 10, n. s. vol. 50, pp. 348-350.
__The distribution of the littoral echinoderms of the West Indies. Carnegie
1920.
1921.,
Inst., Washington, Publ. 281, November, pp. 49-74, pls. 1-3.
Reports on the scientific results of the expedition to the Eastern Tropical
Pacific, in charge of Alexander Agassiz, by the U.S. Fish Commission
Steamer “Albatross” from October 1904 to March 1905, Lieut. Commander
L. M. Garrett, U.S.N., commanding. XXXII. Asteroidea. Mem. M.C.Z.,
February, vol. 39, no. 3, 46 pp., 6 pls.
Echinoderms in birds’ stomachs. Science, June 11, n. s. vol. 51, pp. 594-595.
Reports on the scientific results of the expedition to the Tropical Pacific, in
charge of Alexander Agassiz, by the U.S. Fish Commission Steamer
“Albatross” from August 1899 to March 1900, Commander Jefferson F.
Moser, U.S.N., commanding. XXIJ. Reports on the scientific results of the
expedition to the Eastern Tropical Pacific, in charge of Alexander Agassiz,
by U.S. Fish Commission Steamer “Albatross,” from October 1904 to March
1905, Lieut. Commander L. M. Garrett, U.S.N., commanding. XXXIII.
Holothuroidea. Mem. M.C.Z., September, vol. 39, no. 4, 40 pp., 4 pls.
The echinoderm fauna of Torres Strait: its composition and its origin.
Carnegie Inst., Washington, Publ. 214, January, vol. 10, 223 pp., 40 pls.
Report on the Echinoidea collected by the Barbados-Antigua Expedition
- from the University of Iowa in 1918. Univ. Iowa Studies, Nat. Hist., 15,
March, vol. 9, pp. 103-121, pls. 1, 2.
Vii
1922.
1923.
1924.
1925. ,
1926.
1927.
1928.
1929.
1930.
1932.
With Austin Clark. Results of the Hudson Bay Expedition, 1920. III. The
echinoderms. Contr. to Canad. Biol. n. s. 1, pp. 21-25.
The holothurians of the genus Stichopus. Bull. M.C.Z., February, vol. 65,
no. 3, 38 pp., 2 pls.
Review of Mortensen’s “Studies of the development and larvel forms of
echinoderms.” Science, April 21, n. s. vol. 55, pp. 431-433.
The echinoderms of the Challenger Bank, Bermuda. Proc. Am. Acad. Arts
and Sci., June, vol. 57, no. 13, pp. 353-361, pl. 1.
The echinoderm fauna of South Africa. Ann. So. African Mus., May, vol.
13, pp. 221-435, pls. 8-23.
The distribution and derivation of some New England echinoderms. Am.
Nat., May-June, vol. 57, pp. 229-237.
Some echinoderms from West Australia. Jour. Linn. Soc. London, June 5,
vol. 35, pp. 229-251, pl. 13.
Two new genera of sea-urchins. Bull. M.C.Z., July, vol. 65, no. 10, 10 pp.
Echinoderms from Lower California, with descriptions of new species:
supplementary report. Bull. Am. Mus. Nat. Hist., October 5, vol. 48, pp.
147-163.
Some holothurians from British Columbia. Canad. Field Nat., March, vol.
38, pp. 54-57.
The holothurians of the Museum of Comparative Zoology. The Synaptinae.
Bull. M.C.Z., June, vol. 65, no. 13, 46 pp., 12 pls.
Echinoderms from the South African fisheries and marine biological survey.
Pt. 1. Sea-urchins (Echinoidea). Fish. and mar. biol. surv. Union South
Africa, September 10, 1924. Rep. 4, Spec. Rep. 1, pp. 1-16, pls. 1-4.
A catalogue of the recent sea-urchins (Echinoidea) in the collection of the
British Museum (Natural History). London, 1925, 28, 250 pp., 12 pls.
Some sea stars from the Riksmuseum, Stockholm. Arkiv for zoologi, Novem-
ber 16, vol. 18A, no. 8, pp. 1-8.
A new Clypeaster from Angola. Ann. So. African Mus., December,
vol. 20, pp. 317-318, pl. 33.
Echinoderms from the South African fisheries and marine biological survey.
Pt. 2. Sea Stars (Asteroidea). Rep. 4, Spec. Rep. 7, Fish and mar. biol.
sury. Union So. Africa, May 1, pp. 1-34, pls. 1-7.
Echinoderms other than sea stars of the Tropical Central Pacific. Bull. 27,
Bernice P. Bishop Mus., July, pp. 89-112, pls. 9-11.
Notes on a collection of echinoderms from the Australian Museum. Rec.
Austr. Mus., November 18, vol. 15, pp. 183-192.
With B. W. Arnold. Jamaican fossil Echini. Mem. M.C.Z., October, vol. 50,
no. 1, pp. 1-84, (errata sheets), pls. 1-22.
Review of Mortensen’s “Handbook of the echinoderms of the British Isles.”
Science, December 23, n. s. vol. 66, pp. 625-626.
The sea lilies, sea stars, brittle stars, and sea-urchins of the South Australian
Museum. Rec. So. Austr. Mus., May 9, vol. 3, no. 4, pp. 361-482, text figs.
108-142.
A new Miocene echinoid from California. Trans. San Diego Soc. Nat. Hist.,
August 5, vol. 5, no. 17, pp. 259-262.
The Carnegie-Australia Harvard Expedition to Northwestern Australia.
Science, February 21, n.s. vol. 71, p. 180.
Echinodermata (other than Asteroids). British Museum (Natural History)
Great Barrier Reef Expedition, 1928-1929. Scientific Reports, vol. 4, no. 7,
pp. 197-239, 9 text figs., 1 pl.
The ancestry of the Echini. Science, December 23, vol. 76, no. 1982, pp.
591-593.
Viii
1933.
1934.
1935.
1936.
1937.
1938.
1939.,
1945.
1946.
1948.
A handbook of the littoral echinoderms of Porto Rico and the other West
Indian Islands. Sci. Survey, Porto Rico and Virgin Islands, N. Y. Acad.
Sci., vol. 16, no. 1, pp. 1-147, 7 pls.
Marine collecting in Australia. Harvard Alumni Bull., June 2, vol. 35, no.
32, pp. 901-906.
With B. W. Arnold. Some additional fossil Echini from Jamaica. Mem.
M.C.Z., December, vol. 54, no. 2, pp. 139-156, pls. 1-5.
Some new echinoderms from California. Ann. Mag. Nat. Hist., January,
ser. 10, vol. 15, no. 85, pp. 120-129.
The holothurian genus Caudina. Ann. Mag. Nat. Hist., February, ser. 10,
vol. 15, no. 86, pp. 267-284.
With Elisabeth Deichmann. On Psolicucumis Heding and its allies. Ann.
Mag. Nat. Hist., May, ser. 10, vol. 17, no. 101, pp. 564-568.
A new Eocene sea-urchin from Alabama. Jour. of Paleontology, April, vol.
11, no. 3, pp. 248-249, 3 figs.
The echinoid name Diadema. Ann. Mag. Nat. Hist., December, vol. 20, ser.
10, no. 120, pp. 635-636.
A new sea-urchin from the “Oligocene” of Oregon. Trans. San Diego Soc.
Nat. Hist., December 15, vol. 8, no. 28, pp. 367-374, pl. 24.
Echinoderms from Australia, an account of collections made in 1929 and
1932. Mem. M.C.Z., vol. 55, 596 pp., 28 pls., 64 text figs.
List of Bahama echinoderms. (In W. J. Clench’s report) Bull. M.C.Z., June,
vol. 80, no. 14, pp. 536-537.
Ewo new ophiurans from the Smithsonian-Hartford Expedition, 1937. Proc.
U.S. Nat. Mus., May, vol. 86, no. 3054, pp. 415-418, pl. 52.
A remarkable new genus of sea-urchin (Spatangidae). Allan Hancock
Pacific Expeditions, vol. 2, no. 11, pp. 173-176, 2 text figs.
Ophiuroidea. The John Murray Expedition, 1933-1934. British Museum
(Natural History) Scientific Reports, October 28, vol. 6, no. 2, pp. 29-136,
62 text figs.
A new Astroconus from South Australia. Rec. So. Austr. Mus., December 16,
vol. 6, no. 3, pp. 207-208, pl. 18.
Notes on echinoderms from the west coast of Central America. Zoologica,
November 14, vol. 25, pt. 3, pp. 331-352, figs. 1-4, pls. 1-2.
Revision of the keyhole urchins (Mellita). Proc. U.S. Nat. Mus., December,
vol. 89, no. 3099, pp. 435-444; pls. 60-62.
The echinoderms (other than holothurians). Reports on the scientific results
of the Atlantis expeditions to the West Indies. Mem. Soc. Cubana Hist.
Nat., May, vol. 15, no. 1, pp. 1-154, pls. 1-10.
The echinoderm fauna of Bermuda. Bull. M.C.Z., April, vol. 89, no. 8, pp.
367-391, 1 pl.
Echinoidea in “Geology of Lau,” Fiji, by Ladd and Hoffmeister. Bull. 181,
Bernice P. Bishop Mus., pp. 312-328, pls. 41-43.
‘The echinoderm fauna of Australia, its composition and its origin. Carnegie
Inst., Washington, December, Publ. 566, 567 pp.
A report on the Echini of the warmer Eastern Pacific, based on the collections
of the Velero II. Allan Hancock Pacific Expeditions, vol. 8, no. 5, pp.
i-xii, 225-351, pls. 35-71.
ix
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PREFACE
Mrs. H. L. Clark and her family have done me the great honour of
asking me to edit my late friend H. L. Clark’s Report on the Echini of
the Warmer Eastern Pacific collected by the Velero III. In so doing I
have thought it a holy duty towards my dear colleague and friend through-
out nearly half a century to let his report appear in the main as it was
intended by him, adding only some few remarks here and there, which I
have deemed necessary. Only in his dealing with the genus Encope a more
serious correction has been introduced. It was the idea of Clark that the
genus ought to be divided in two separate genera, the species grandis to
form a genus of its own in contradistinction to all the other species. This
is in my opinion quite unacceptable. The only difference is that grandis is
thicker and somewhat heavier than the other species ; but whether the edge
of the test is 5 mm or only 2-3 mm thick is certainly not a morphological
difference of generic value, and there are no other differences ; especially it
should be emphasized that the internal structure of the test is exactly the
same as in the other species. Moreover, separating grandis as a distinct
genus from all the other species would necessitate most regrettable nomen-
clatorial changes. His idea that the species emmarginata would be the geno-
type of Encope is decidedly erroneous. When establishing the genus
Encope L. Agassiz (1840) named only a single species, grandis ; accord-
ingly grandis is the genotype of Encope. In one of his last letters to me
Clark said that he would abide at the decision which I would make in this
question. I wrote him at once that there could be no question at all but
that grandis is the genotype of Encope, and he would certainly have
changed his manuscript in accordance herewith; but he did not get the
time to do so. I have thus had to change that part of his manuscript.
As for the numerous new species of Encope which he establishes I think
he is going much too far, small individual or local variations of this pro-
lific genus being taken to represent distinct species. A few specimens (one
of each species, and some of them too young to be of any use) sent me by
Professor McCulloch do not at all convince me of the value of all his
species; but without having access to the rich material on which Clark
based his various species it is impossible for me to decide which of his
Encope species are of real value and which are not. I deem it therefore
our duty to publish them all, as Clark meant to have it done. It must then
be left to posterity to solve definitely the question of the species to be
distinguished within the forms belonging to the circle of—especially—
Encope micropora. I cannot help also thinking how many species Clark,
on the base of the study of a similar rich material of the Caribbean Encope
emarginata might have made out of that polymorphic species.
I hope that by the few corrections and additions introduced in this
manuscript I am doing justice to my late, lamented friend, the eminent
echinologist, Hubert Lyman Clark.
Copenhagen, in April 1948.
TH. MortENSEN
xii
REPORTS ON THE COLLECTIONS OBTAINED BY ALLAN HANCOCK PACIFIC EXPEDITIONS OF
VELERO III OFF THE COAST OF MEXICO, CENTRAL AMERICA, SOUTH AMERICA, AND
GALAPAGOS ISLANDS IN 1932, IN 1933, IN 1935, IN 1936, IN 1937, IN 1938, IN 1939,
IN 1940, AND IN 1941,
A REPORT ON* THE ECHINI
OF THE WARMER EASTERN PACIFIC,
BASED ON THE COLLECTIONS OF THE
VELERO III
(Piates 35-71, Text Ficures 1-3)
By HUBERT LYMAN CLARK
THE UNIVERSITY OF SOUTHERN CALIFORNIA PUBLICATIONS
ALLAN Hancock PAciFic ExpEDITIONS
VOLUME 8, NUMBER 5
IssUED DECEMBER 29, 1948
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
Los ANGELES, CALIFORNIA
A REPORT ON THE ECHINI OF THE WARMER EASTERN
PACIFIC, BASED ON THE COLLECTIONS OF THE
VELERO III
(PLaTes 35-71, TeExT Ficures 1-3)
HuBeErt LYMAN CLARK
The great collections of Echini (28,835 specimens, including over
11,000 young Dendrasters) made by the Velero III under the direction
of Captain Allan Hancock, during the years 1931-1941, have proved a
field for research as varied and interesting as it is large and illuminating.
It was just eighty years ago that Verrill published his first papers on the
Echinoderms of Panama and the western coast of America and for four
years he continued his studies of that fauna. In 1871 there appeared
(Trans. Conn. Acad. vol. 1, pt. 2, art. 5, no. 8, pp. 593-595) his list of
the Echini of the Gulf of California which included 22 species. Subsequent
additions and emendations enabled A. Agassiz in 1904 to list 28 species
from the Panamic region and 21 more from very deep water (300-2200
fms). In recent years additional species have now and then been added or
corrections made, so that Grant and Hertlein (1938) include some 47
recent species in their list. Several, however, are from very deep water
and are hardly to be counted in the present fauna of the warmer Eastern
Pacific.
The area thus designated may be defined as extending from the coast
of Oregon at Ocean Park (Lat. 44° 50’ 45” N), southward along the
American coast to San Juan Bay, Peru (Lat. 15° 20’ S), and westward
to include the islands, Socorro, Clarion, Cocos, and the Galapagos, down
to depths of 80 fms or less, very rarely to 300-400 fms. In depths exceeding
100 fms, the water is no longer warm and the bottom fauna can not be
properly included as belonging to the “warmer Eastern Pacific.” The
great gulf of California has an interesting fauna including 22 species of
Echini. Only one of these, however, is restricted to the Gulf, 8 extend
their range to the north and 19 to the south. Due to certain local restric-
tions, collecting along the coast of Mexico and Central America did not
yield so large a number of Echini as did the Gulf, and even in the Bay of
Panama only 18 species were taken. Colombia yielded but 11 species while
Ecuador has 14 to her credit and only 6 were recorded from Peru. North
of the international boundary (United States-Mexico) some 14 species
have been taken by the Velero. The outlying islands have proved good
collecting grounds, for while Cocos yielded only 11 species, Clarion had
10, Socorro 12, and the Galapagos no few than 23, of which 4 were not
found elsewhere.
[225]
226 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
The five visits of the Velero to the Galapagos, occupying 245 stations,
have undoubtedly served the purpose of making the Echinoderm fauna
of that group sufficiently well known to warrant the belief that it is
essentially American and has not received any considerable influx of
Indo-Pacific forms. Several of the species are, so far as we now know,
endemic and warrant a belief in the long-sustained isolation of the marine
fauna. In view of this it is worth while to list here the Galapagos Echini
with a few words as to their occurrence:
Eucidaris Thouarsit
Hesperocidaris panamensis
Centrocidaris Doederleini
Centrechinus mexicanus
Centrostephanus coronatus
Lytechinus semituberculatus
Toxopneustes roseus
Tripneustes depressus
Caenocentrotus gibbosus
Echinometra oblonga
VanBrunti
Clypeaster elongatus
europacificus
ochrus
rotundus
Encope galapagensis
Cassidulus pacificus
A gassizia scrobiculata
Brissopsis pacifica
generally Panamic and western
Mexico
Cocos Island, Galapagos Islands and
Ecuador
Cocos and Galapagos Islands
generally Panamic, Mexico and
Galapagos Islands
generally Panamic, Gulf of Cali-
fornia, southern California
Galapagos Islands
Panamic, Socorro Island
Panamic, Mexican Islands
Ecuador, Peru and Galapagos
Islands
Indo-Pacific to Northern Galapagos
Islands and Mexican Islands
Panamic, Western Mexico
Galapagos Islands
Panamic, Western Mexico
Panamic
Panamic, Western Mexico
Galapagos Islands
Panamic, Mexican Islands
Panamic, Gulf of California
Panamic, Western Mexico,
southern California
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 227
Idiobryssus coelus Galapagos Islands
Meoma grandis Panamic, Western Mexico
Gonimaretia laevis southern California and Gulf of
California
Lovenia cordiformis Panamic, Western Mexico,
southern California
Aside from the light thrown on the distribution of the known Echini
in the Eastern Pacific, the Yelero has collected an abundance of material
useful in delimiting the known species, and revealing the existence of 11
forms which appear to require names as they do not fit too well into the
series of species now known. One of these is a Cidarid and one a Clype-
aster, but others are Scutellids of the well-known genera Dendraster and
Encope. Never before have such large numbers of specimens been avail-
able for comparative study, and it is not strange that new and perplexing
forms occur which require designation. Quite likely more material from
the coasts of southern Mexico, Costa Rica, Nicaragua, and Panama will
indicate different and more correct grouping, but it is believed that the
specific limits here suggested indicate a step forward in our knowledge of
these multiform genera.
The naming and describing of new species, however, is only one prod-
uct of the Velero’s extraordinary collecting. Equally important is the
accumulation of immature and very young individuals showing stages in
the development of the adult which throw light on evolutionary processes.
The most striking of these is exhibited in the growth changes of young
Spatangus, a genus represented by but one species in the Eastern Pacific,
widely separated from its congeners in Europe, Asia and Africa. The
V elero first met with Spatangus in 1936 in the Gulf of California but later
collections showed that the vicinity of the Channel Islands in depths of 35-
225 fms is the real home of this interesting sea-urchin. Considerably more
than two hundred specimens are in the collection and quite a number are
young, including some very early stages of development, only about 3 mm
long. In these very small individuals there is no indication of either petals
or pores. When the individuals are 8 or 9 mm long the petals are indi-
cated by more definite outlines and slight depressions, but there are no
pores. After the length exceeds 10 mm, the petals can be distinguished
more or less definitely and pores are visible at the apical end of the ambu-
lacra. But the pores are single and the petals evidently are not sunken at
all. There is thus a Palaeotropus stage of development shown which is
usually passed through befere the Spatangus is 12 mm long. In some
228 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
cases, however, the growth changes are retarded and one individual
caused much perplexity as it is 20 mm long and still reveals only single
pores in the petals and hence was regarded as a new species of Palaeotropus.
But the discovery of intermediate stages, showing all steps from poreless
plates to normal twin-pored petals has been convincing in the demonstra-
tion of this striking example of stages in development, so ably presented
by Jackson (1899).
For the opportunity to study this extraordinary collection, the writer
is greatly indebted to Chancellor R. B. von KleinSmid of The University
of Southern California and to Captain Allan Hancock and Professor
Irene McCulloch of the Allan Hancock Foundation. Lieutenant Com-
mander Fred C. Ziesenhenne, who played a major part in the collecting,
preserving and arranging of the material, deserves high praise for the
quality of his work. For the pleasure of being his co-worker in the 1938
cruise of the Velero III, I am deeply indebted to Captain Hancock. That
was one of life’s great experiences and words are quite inadequate to
express my appreciation. Finally I take pleasure in acknowledging the
very great debt I owe to Mrs. Ora Willett for her patient assistance in
the study of the collection and in the preparation of the manuscript—
assistance for which these few words of appreciation are quite inadequate.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 229
Order CIDAROIDA
Family Cidaridae
Eucidaris Thouarsii (Valenciennes)
Plate 35, Fig. 1
Cidaris Thouarsii L. Agassiz and Desor, 1846, p. 326.
Cidaris (Eucidaris) T houarsii Déderlein, 1887, p. 20.
Eucidaris Thouarsii Mortensen, 1928a, pp. 393-400, pl. 42, figs. 5-13.
There are, in the Hancock collections, more than 1,200 specimens from
175 stations. They range in size from 3.5 to 67 mm in diameter. The
largest and finest specimens are from the Galapagos Islands, particularly
Darwin Bay, Tower Island, and Cartago Bay, Albemarle Island. The
character of the spines, both primary and secondary, shows extraordinary
diversity. The secondaries range from relatively short, truncate spines to
rather long ones tapering to a chisel-like tip. The primaries show even
greater diversity ranging from short, stout spines with diameter .25-.30
of length to slender terete spines with greatest diameter less than a tenth
of the length. There is great diversity also as to the extent to which the
spines are covered with sponges, bryozoa or calcareous algae. Occasionally
these foreign growths increase the diameter of the spine (at least at its
base) to one-third of its length. In other individuals of the same size, from
the same station, the spines may be entirely free from any foreign growth.
The greater part of the Velero collection of Thouarsii is from the
Galapagos Islands and a notable proportion of these individuals are larger
and stouter than the material from the mainland coast. The specimens
from Colombia, Panama, Costa Rica, and Mexico are on the whole smaller
and have more slender spines than those from the islands. Occasional,
sometimes conspicuous, exceptions prevent the drawing of any clear line
ot distinction however. Mortensen’s emphasis on the “nearly black”
secondary spines in galapagensis is misplaced as most specimens from either
mainland or islands have the secondaries a deep purplish brown, the shade
apparently deepening with age. On the whole, it seems to me futile to try
to maintain any well-defined form as a variety or subspecies galapagensis.
Mortensen (1928, p. 399) has called attention to certain peculiarities
in a specimen from Clarion Island which, if constant, would warrant
recognition as at least a variety of Thouarsii. Unfortunately the present
collection contains only a very few specimens from Clarion Island. While
the largest of these seems to resemble the one studied by Mortensen, the
others are not distinguishable satisfactorily from specimens from the main-
land coast. Further collecting at Clarion Island is greatly to be desired.
Distribution.—This is one of the most characteristic sea-urchins of
the western coast of tropical and subtropical America. E. Thouarsii
occurs also at the outlying islands (Clarion, Socorro, Cocos and the
230 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Galapagos), but not at great depths. Mortensen reports it in 45 fms in
the Gulf of Panama, while the Velero took specimens at depths of 62-76
fms off Angel de la Guardia Island in the Gulf of California. The most
northerly point from which it is reported is near Consag Rock in the Gulf
of California. It is noteworthy that it has not been taken along the western
coast of Lower California. Its southern limit, on the continental coast, is
La Plata Island, Ecuador. It is very common among the Galapagos
Islands where the /’elero collected specimens at no fewer than 70 stations.
Type——Paris Museum?
Type locality —“Californie (Neboux.) Galapagos.”
Depth.—Shore to 76 fms.
Specimens examined.—1,201 specimens from 175 stations.
Hesperocidaris panamensis (A. Agassiz)
Plate 35, Fig. 2
Dorocidaris panamensis A. Agassiz, 1898, p. 73; 1904, p. 20, pls. 1-4.
Hesperocidaris panamensis Mortensen, 1928, pp. 73-74; 1928a, Mon.
Ech. p. 416.
The material hitherto known of this sea-urchin consists of a few
specimens taken by the 4/batross in 1891 and now distributed in the U.S.
National Museum, the Museum of Comparative Zoology and the British
Museum. The Velero, however, has secured no fewer than 84 specimens,
ranging from 4 to 49 mm in diameter; the height of the test is about half
as much or a little more; both the upper and lower surfaces are definitely
flattened. The very young individuals are white orally but more or less
light coral red above; the primary spines are whitish with a tinge of red
(at least basally) and have one or two ill-defined dusky bands and orange-
red or flesh-red collar. In one specimen from the Galapagos Islands
(Station 183-34) in 50-70 fms, the primaries are white except for more
or less of the tip which is to some extent dusky or reddish; the contrast
between such primaries and the dull red secondaries is striking. With
increasing size, however, the primaries become unicolor, gray or brown-
ish, save for the collar, and rapidly increase in darkness until in adults,
they are deep brown or brownish purple, except where more or less in-
crusted with bryozoa, or other symbiotic forms. As a rule incrusta-
tion is not extensive and the primaries are slightly flattened at the tip.
In young specimens the collar tends to be conspicuously orange red, but
with age the color deepens and in adults is dark red or even reddish
purple of a very dark shade. The secondary spines are brownish red in
small specimens but become dark, almost a deep blood red in adults.
A full grown specimen is thus a very striking and rather handsome urchin.
In some individuals the young primaries at the upper end of each series
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 231
are uniformly more or less coral red in rather noticeable contrast to the
older spines; this is particularly striking if the spine is entirely free from
bryozoa or other incrustations. On the whole the coloration and general
appearance of panamensis is remarkably uniform and makes the species
easy to recognize.
Distribution —The most striking fact about the distribution of this
Cidarid is that it has not yet been taken at Panama or anywhere else on
the coast of North or Central America. The original specimens were from
the vicinity of Cocos Island, and from 85-112 fms off Galera Point,
Ecuador. The Velero material is all from Cocos Island and vicinity, where
this handsome urchin is notably common in 30-50 fms, or from the
Galapagos Islands where it was taken at eight stations in depths of 30-150
fms, chiefly in the vicinity of Albemarle Island. The station numbers are:
143-34, 147-34, 155-34, 171-34, 183-34, 190-34, 324-35, 325-35, 772-38,
773-38, 780-38 and 810-38.
Type.—M.C.Z. no. 327 (Cotype).
Type locality —“Cocos Island and Galera Point, Ecuador.”
Depth.— 30-150 fms.
Specimens examined.—84 specimens from 12 stations.
Hesperocidaris perplexa (H. L. Clark)
Plate 36, Fig. 3
Tretocidaris perplexa H. L. Clark, 1907, p. 205; pl. 6, figs. 1, 2; pl. 7,
figs. 1-4.
Hesperocidaris perplexa Mortensen, 1928, p. 421.
Only five specimens of this near relative to panamensis were known
hitherto, so that the large series of nearly 500 specimens in the Velero
collections contribute all the information needed to verify the authenticity
of the species and reveal such diversity as it shows. The specimens range
from 5 mm in test diameter, with spines scarcely 5 mm long to large adults
with the test 43 mm in diameter and 25 mm high, with primary spines
35-43 mm long. The flattening of the distal part of the large primary
spines, on which Mortensen lays great stress, is not evident until the test
is 30 mm or more in diameter and even then it is not conspicuous in most
specimens. In the most extreme case at hand, the test is about 37 mm in
diameter and the longest primaries are about the same; the tips of ten or
a dozen of these, at or above ambitus, are 3 to 5 mm wide; the distal half
of each of these widened spines is very rough with crowded, more or less
sharp tubercles. In most specimens the primary spines are nearly cylindri-
252 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
cal even at tip or somewhat terete, and are seldom noticeably rough.
Their color is grayish or dull brownish but is frequently altered by bryo-
zoa, sponges or other foreign growths. ‘The striking and very constant
feature of the coloration in perplexa is the conspicuous broad, brown
longitudinal stripe which occurs on all the spinelets. The test itself is more
or less yellowish or greenish but this shade is well shown chiefly, if not
only, in the ambulacral mid-zones. The color of the basal part of the
primary spines is commonly pale brown, usually greenish or yellowish
just above the collar. The collar itself may have a more or less similar
tint or may be more green, or yellow; in many specimens the general color
of the spine is more or less dark brown, with the primary color flesh red
or yellow brown. There is a great deal of variety in the shades of color
in perplexa, some individuals being notably dark while many specimens,
especially when young are definitely light. But no matter what the shades
may be one feature of the coloration is remarkably constant and that is
the longitudinal dark stripe found on all the secondary, and smaller, spines.
Distribution—Judging from the Velero collections, this Cidarid is
particularly characteristic of the Gulf of California, having been taken at
16 stations, as far north as 29° 39’. Off the west coast of Lower Cali-
fornia, there are no records, save one half grown specimen in poor condi-
tion supposed to be from Station 1119-40, south of San Benito Islands,
and a very young individual also in poor condition labeled as from
Station 1153-40, off Santa Catalina Island, California. It is probable that
these two specimens have in some way become mislabeled.t South of
Lower California, no specimens were taken save at Bahia Honda, Panama,
(Stations 244-34, 863-38, and 948-39) in 30-50 fms, north of Gorgona
Island, Colombia (Stations 854-38 and 855-38) in 10-60 fms, and off
La Plata Island, Ecuador, (Stations 212-34, 213-34) in 7-55 fms. The
entire absence of perplexa from the Galapagos area is worthy of note and
even more remarkable is its absence from the vicinity of Cocos Island,
where panamensis is common.
Type.—M.C.Z. no. 188.
Type locality—Gulf of California.
Depth.—36-39 fms.
Specimens examined.—487 from 25 stations.
1 There are evidences in the studies of other groups that unexpected distribu-
tional records are not uncommon. The work of the Velero III must be considered as
the preliminary exploration of a coast involving many miles with little information
concerning the fauna of the intervening areas. Ed.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 233
Hesperocidaris asteriscus”, new species
Plate 36, Fig. 4; Plate 37, Figs. 5-6
Test somewhat flattened; vertical diameter about .60 of horizontal;
coronal plates 6; areolae large, occupying most of plate surface; median
interambulacral area narrow and fully covered with tubercles, smallest
next to vertical suture which is quite indistinct ; ambulacra about one-third
of interambulacra in width; poriferous zones rather wide and hardly at all
sunken; median ambulacral area narrow with a single row of relatively
large tubercles on each margin and between these an irregular double series
of much smaller tubercles which become more and more scattered and
disappear entirely before the ocular plate is reached; pores rather large,
horizontal or slightly oblique, nearly circular. Abactinal system about .50
h. d., rounded pentagonal, clearly defined, the anal area somewhat ele-
vated; each of the component plates and the anal area covered (save on
the margins) with flat spinelets, the larger ones on the anal system with
truncate or rounded tips, the others quite acute; the genital plates carry
30-35 spine-bearing tubercles, the ocular plates about 20; genital plates
distinctly longer than wide and much wider at the inner end than at the
slightly convex outer end; ocular plates triangular, twice as wide as long,
the margins slightly convex; both genital and ocular plates are bare,
smooth and cinnamon color? on the margins (excepting the distal margins
of the oculars) so that the genital plates form a conspicuous star-shaped
figure; ocular plates all markedly excluded from the relatively large pen-
tagonal anal system. Actinostome rather large, more than .40 h. d., not
noticeably sunken, well covered with rather stout plates, which are hidden
by the numerous spines they carry ; these spines are long and narrow, with
the distal end widened, rounded and curved inward very slightly. Primary
spines longer than horizontal diameter of test, nearly cylindrical, fairly
stout and blunt; they are densely covered with low, blunt tubercles in
about a dozen more or less regular longitudinal series ; collar very narrow;
secondary spines flat, only a little wider near the base than at the truncate
slightly rounded tip; miliary spines, few, very small, acute. No globiferous
pedicellariae were found on any one of the specimens; tridentate of two
kinds, one with long narrow jaws, are abundant. In color, the test is
pinkish cinnamon with the median ambulacral area distinctly light yellow-
ish olive; secondary and miliary spines brownish olive of diverse shades ;
young primaries Japan rose or slightly darker, in some cases with one or
2 asteriscus—a little star, in reference to the star-like figure formed by the
genital plates.
3 Color names are from Ridgway Color Chart as determined at the time of this
study.
234 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
two light bands near tip; mature primaries blackish brown except where
covered by encrusting bryozoa or other organisms ; collar very low, verona
brown; primary spines of oral surface light colored, those around the
actinostome may be nearly white. The holotype is 26 mm h. d., 13 mm
v. d.; longest primaries 35 mm with a diameter near base of nearly 3 mm;
at tip, not quite two.
Besides the holotype, there are two other Cidarids that must be re-
ferred to this species, as they agree in all essentials. One is nearly as large
as the holotype but is badly damaged on one side. The primaries of the oral
surface show more red in their coloration and the secondaries are a lighter
shade of brown. This specimen was taken with the holotype at Velero
Station 948-39, March 28, 1939, off Medidor Island, Bahia Honda,
Panama, in 30-35 fms on a bottom of mud, rocks and corallines, in com-
pany with large numbers of perplexa. A third specimen, 15 mm in diameter
with primaries 20-26 mm long, is supposed to be from Station 575-36, in
the Gulf of California, but there is some doubt as to the reliability of the
label. It is unquestionably conspecific with the specimens from Bahia
Honda but shows much more red in the coloration and the primaries are
not so black as in the holotype. The relationship of this species to perplexa
is obvious, but to refer these specimens to that species would evidently be
quite wrong.
Type.——Holotype, AHF no. 46, 1 paratype.
Type locality.—Station 948-39, off Medidor Island, Bahia Honda,
Panama, 30-35 fms, March 28, 1939.
Distribution.—Type locality and the Gulf of California (?).
Depth.—30-100 fms (?).
Specimens examined—The type, paratype and one other specimen,
from 2 stations.
Centrocidaris Doederleini (A. Agassiz)
Plate 38, Fig. 7
Goniocidaris Doederleini A. Agassiz, 1898, p. 73.
Centrocidaris Doederleini A. Agassiz, 1904, p. 33, pl. 14, figs. 1, 2.
Apparently this is a very rare urchin for, in all her collecting, the
V elero has met with but one specimen—a small adult taken in the Galapa-
gos Islands, near Barrington Island, in 48-73 fms. It is 24 mm in diameter
and 13 mm high. The primaries are 20-28 mm long, exceedingly slender,
and nearly cylindrical, either smooth or with low, well separated ridges.
The abactinal system shows a star-like pattern, as in Hesperocidaris asteris-
cus but in Centrocidaris the lines are dark brown with light brown plates
instead of the reverse.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 235
Distribution —Cocos Island, Galapagos.
Type.—U.S.N.M. No. 27348.
Type locality:—Albatross Station 3369, off Cocos Island, 52 fms.
Depth.—48-300 fms.
Specimens examined.—1 specimen.
Order CENTRECHINOIDA
Family Centrechinidae
Centrechinus mexicanus (A. Agassiz) *
Plate 38, Fig. 8
Diadema mexicanum A. Agassiz, 1863, p. 20.
Centrechinus mexicanus Ziesenhenne, 1937, p. 231.
Diadema mexicanum Mortensen, 1940, p. 275, pl. 60, figs. 7-11.
This big black sea-urchin, with long slender, acutely pointed and very
poisonous spines, is a nuisance to bathers, beachcombers and fishermen
throughout the tropical Eastern Pacific. Adult specimens are 70-80 mm
in diameter with the height of the test about half as much. The primary
spines may exceed twice the diameter of the test but they are so very fragile
that it is extremely difficult to preserve specimens with the spines unbroken.
The secondary spines are very slender and acicular but not very long;
when they dry they are so fragile they are easily broken and lost. The
absence of spines on the peristome is a constant and striking feature. Full
grown individuals are virtually unicolor, black or very dark brown; occa-
sionally definitely brown specimens occur. As in all the species of Centre-
chinus (or Diadema) very young specimens have the long spines prettily
banded with white and some dark shade, usually purple, but the white is
sometimes tinged with brown and the purple is nearly black. With increas-
ing size the white becomes more and more dingy and half grown specimens
may have it entirely suppressed and the spines all uniformly dark. Banded
spines in specimens half grown or more are common but usually only a few
of the primaries show it. The poisonous character of the spines is well
known and those who have once suffered the pain which even a single
spine can cause, are forever after extremely careful in the vicinity of
Centrechinus.
Distribution.—This species was taken by the Velero at 38 stations,
of which the most northern was at Consag Rock in the Gulf of California,
the most southern at Gorgona Island, Colombia. The absence of specimens
from the western coast of Lower California is notable, since the urchin
occurs at both Socorro and Clarion Islands. It is also found at Cocos
Island and throughout the Galapagos Islands. It is definitely a shallow
water echinoid, for, although it was taken in 40-57 fms at the island, along
*Please see explanation on p. 243.
236 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the mainland coast nearly all the specimens at hand were taken along
shore, very rarely in more than 2 fms.
Type.—M.C.Z. no. 635 (Cotype).
Type locality Acapulco, Mexico.
Depth.—Shore to 57 fms.
Specimens examined.—191 specimens from 38 stations.
Astropyga pulvinata (Lamarck)
Plate 39, Fig. 9
Cidarites pulvinata Lamarck, 1816, p. 59.
Astropyga pulvinata L. Agassiz and Desor, 1846, p. 345.
Mortensen, 1940, p. 197, pls. 20-23.
One of the characteristic Echini of the warmer Eastern Pacific, this
handsome sea-urchin is well represented in the Velero collections. There
are altogether 165 specimens, ranging in size from 5 mm h. d. with
primary spines 7 mm long, to adults 100-117 mm in diameter. The
diversity of color is very striking but is brightest in the young which are
often very handsome and dullest in the adults. The largest specimen in the
present collection is dry and measures 117 mm h. d. and about 40 mm v. d.
It is uniformly dark (almost black) above save for the usual large triangu-
lar interradial spots which are light buff but were probably white in life.
Apparently this represents the form collected by Mortensen at Contadora
Island, Panama, which he designates as variety venusta Verrill. The
present specimen was taken in 15-20 fms in Tangola Tangola Bay,
Mexico, in company with 3 specimens 70-90 mm h. d. which are all
typical, handsomely variegated pu/vinata, as are many other Mexican spec-
imens. A very fine specimen, however, from Tiburon Island, Gulf of
California, 10 fms, is almost completely dark violet above, the interradial
spots being scarcely distinguishable and the lower surface being very
markedly suffused with violet. This is the most heavily pigmented speci-
men in the collection though only about 90 mm h.d. Young specimens
(17-35 mm) from the Gulf of California, Panama, and Ecuador are
very handsome in their variegated liveries of bright red, violet and yellow.
It does not seem to me desirable to give varietal names to any of the varied
forms, even though such extremes as venusta may be recognizable.
Distribution —This sea-urchin was taken by the Velero at some 16
stations along the Mexican and Central American coasts in depths down
to 20 fms though many collections were made along shore. The northern-
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC PAY |
most stations were at Tepoca Bay, Sonora, where only two very young
specimens were taken and Tiburon Island, in the Gulf of California, in
16 fms, where 74 specimens were secured. ‘he southernmost station was
at Santa Elena Bay, Ecuador, in 8-10 fms, but no specimens were taken
at either Cocos or the Galapagos Islands.
Type.—Paris Museum?
Type locality—“ Habite—probablement les mers de 1’ Asie.”
Depth.—Shore to 20 fms.
Specimens examined.—165 specimens from 16 stations.
Centrostephanus coronatus (Verrill)
Plate 39, Fig. 10
Echinodiadema coronata Verrill, 1867, p. 295.
Centrostephanus coronatus A. Agassiz, 1872a, p. 97.
Mortensen, 1940, p. 314, pl. 36, figs. 7-10.
Compared with the preceding two species, this is a relatively small
sea-urchin, the largest known specimens being only 63 mm in diameter.
The largest of the 131 specimens collected by the Velero are only 45-50
mm with the primary spines about twice as much. As the spines are very
brittle, especially when dry, few specimens (even in the Velero collec-
tions, notable for the fine condition of typical specimens in nearly every
species) show the handsome appearance of this sea-urchin in life. As in
Centrechinus the spines are finely verticillate and very acute but there is
no evidence that they carry poison at the tip as do those of Centrechinus.
A very interesting character of this Centrostephanus is the presence on
the uppermost interambulacral plates of short claviform spinelets the tips
of which are bright red purple. This color persists in most preserved
specimens but may be much duller and hence less noticeable than in life.
Mortensen discovered that these spinelets are never resorbed or replaced
by the long primary spines but the latter grow up around them and en-
close them without resorption. Young individuals of this Centrostephanus
are so similar to young examples of Centrechinus mexicanus that they are
often confused with them but there are three characters by which the two
species may be readily distinguished. The most important of these is the
presence of these brightly tipped claviform spinelets in Centrostephanus
which are never present in Centrechinus. A second distinctive character is
the presence of slender, blunt, light-colored spinelets on the five pairs of
oral plates around the mouth (in Centrechinus these plates may have
pedicellariae but never spinelets). The third difference is in the color of
238 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 35
Fig. 1. Eucidaris Thourasii (Valenciennes), aboral view, x34, p. 229
Fig. 2. Hesperocidaris panamensis (A. Agassiz), aboral view, x44, p. 230
PLATE 36
Fig. 3. Hesperocidaris perplexa (H. L. Clark), aboral view, x%, p. 231
Fig. 4. Hesperocidaris asteriscus, new species, lateral view, x¥, p. 232
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC PL. 35
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PL. 36
ay
4a)
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC PL. 37
“ss
ALLAN HANCOCK PACIFIC EXPEDITIONS
VOL. 8
no. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 243
PLATE 37
Fig. 5. Hesperocidaris asteriscus, new species, aboral view, x4, p. 232
Fig. 6. Hesperocidaris asteriscus, new species, oral view, x44
PLATE 38
Fig. 7. Centrocidaris Doederleini (A. Agassiz), aboral view, x1, p. 234
Fig. 8. Centrechinus mexicanus (A. Agassiz) ,* oral view, x4, p. 235
*In a recent letter from Th. Mortensen he states that The International Com-
mission on Zoological Nomenclature meeting in Paris this summer (1948) rendered
an Opinion which codified the name Diadema as a nomen conservandum.
This species is, therefore, Diadema mexicanum A. Agassiz, order Diadema-
toidea, family Diadematidae. These corrections replace corresponding names
used on p. 235 of the text.
244 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the primary spines; in young Centrechinus the spines are banded bluish
or purplish and white, in Centrostephanus dull purplish brown and light
yellow. With increasing age and size, the colors become duller until the
mature primaries are uniformly black in Centrechinus, dark brown in
Centrostephanus. It is unusual for banded spines to be evident in adults of
either genus but carefully preserved alcoholic specimens of Centro-
stephanus, even when full grown, commonly show the banding on some
primaries at least, though faintly. Most dry specimens, however, appear
to be more or less uniformly dark brown or quite black. On the contrary,
very young specimens (less than 10 mm in diameter) are unusually pretty
objects, the test and small spines deep brown, the dull yellow primaries
more or less evidently banded with purplish brown, and five or more
claviform spinelets, with bright rose-purple tips around the apical system.
Distribution —This species of Centrostephanus is definitely a shallow
water Mexican sea-urchin but the Velero took it at five stations in the
Galapagos Islands. At Albemarle Island it was found in shallow water
but off James Island it occurred in 36-60 fms. It was not met with at
either Cocos or the more northern islands (Socorro and Clarion), or on
the west coast of Lower California. Yet it is frequently found at Newport
and Corona del Mar, California, near low water mark. It has yet to be
taken on the mainland coast south of the Gulf of California but in the
Gulf it has been found as far north as Rocky Point, Sonora, and Consag
Rock, in depths down to 45 fms.
Type.—Peabody Museum, Yale University, no. 964.
Type locality—Cape San Lucas, Lower California.
Depth.—Shore to 60 fms.
Specimens examined.—131 specimens from 37 stations.
Family Arbaciidae
Arbacia incisa (Blainville;? Gmelin)
Plate 40, Fig. 11
Echinocidaris incisa A. Agassiz, 1863, p. 20.
Arbacia incisa H. L. Clark, 1913, p. 220.
Arbacia stellata Mortensen, 1935, p. 575, pl. 70, figs. 6-9.
This is one of the commonest sea-urchins of the Gulf of California
and was taken by the Velero at many stations there. Along the coast
southward it seems to be less common and relatively few specimens were
secured but unmistakable examples were collected as far south as Peru.
Indeed one of the largest specimens in the collection, 47 mm h.d., was
taken nine miles south of Zorritos Light, in shore collecting; from tip to
NO. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 245
tip of the extended spines it measures 120 mm. Mortensen refers to the
“largest size recorded for this species is 60 mm diameter” but he does
not give his authority for this giant. Only a very few of the Velero’s 1,121
specimens exceed 35 mm h. d. (the largest is 56) and a very large number
are less than 20 mm across. Very young specimens (2-8 mm h. d.) are as
a rule light colored more or less flesh red, becoming red brown, or violet
or deep purple, until finally they are nearly or quite black. A few dry
specimens are more or less definitely gray. One curious variety has some
of the primaries, or at least portions, quite pink or white in striking con-
trast to the black of the remaining spines and test. Mortensen (p. 566)
makes the “conspicuous red spots in the interambulacra on the aboral
side” the distinguishing mark of the species and the Velero collection con-
firms his judgment. Even in the most nearly black adults, careful examin-
ation in good light will show at least faint indications of the tell-tale red
blotches. Young individuals with the test flesh color or red and the red
spines with one or more whitish bands are so different from the more
mature specimens, it is hard to believe they can ever become as dark
colored as they do, but the present large series compels the acceptance of
the fact. In one or two of the very black specimens there is real difficulty
in seeing any red, but such individuals cannot be distinguished satis-
factorily as anything other than excessively pigmented incisa.
Mortensen’s (1935, p. 577) plea for the specific name stellatus is
inadequate, and if we once let such arguments control our decisions we
shall have more disagreements and resulting confusion than we have had
hitherto. Nobody knows to what sea-urchin the name stellatus was first
given. Everyone knows what Echinocidaris incisa was at the start and
there has never been any confusion about it, so I have no hesitation in
continuing the use of the name here.
Distribution——The Velero collections show that this is essentially a
species of the Gulf of California. It ranges north to the southern United
States as shown by Ziesenhenne (1941, pp. 117-120). There are 6 very
fine adult specimens in the Velero collection from Newport Harbor,
California, but the Velero has taken no other specimens north of Lower
California, nor along its western coast. South of the Gulf, incisa does not
appear to be very common though there are specimens at hand from Bahia
Honda, Panama; Gorgona Island, Colombia; Santa Elena Bay and La
Plata Island, Ecuador; Zorritos Light, South Bay, Lobos de Afuera,
4 The authority for this giant is Clark himself, who says in his Echinoderms of
Peru, p. 345, that “large specimens may be 60 mm in diameter.” TH. MorTENSEN
246 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
and Middle Chincha Island, Peru. The last is the farthest south for
incisa, 13° 39’ 15” S. All stations are in shallow water, less than 50 fms
and almost always less than 20. A very small specimen is labeled as from
Station 1105-40 in the Gulf of California, with a depth of 113-127 fms.
It is probable that this individual was actually dredged at Station 1101-
40, but was lost in the meshes of the dredge until the careful search after
a deep water haul. It is notable that no specimens of incisa were taken at
the Galapagos Islands, Cocos, Clarion or Socorro Islands, nor along the
Costa Rican coast, though it has been previously recorded from the
Galapagos.
Type—M.C.Z. no. 467 (Cotype).
Type locality —Guaymas, Lower California, Mexico.
Depth.—Shore to 50 fms.
Specimens examined.—1,121 specimens from 82 stations.
Arbacia spatuligera (Valenciennes)
Plate 40, Fig. 12
Echinus (Agarites) spatuliger Valenciennes, 1846, pl. 5, fig. 2.
Arbacia spatuligera A. Agassiz, 1872a, p. 93.
H. L. Clark, 1910, p. 346, pl. 10, fig. 2.
Mortensen, 1935, p. 577, pl. 70, figs. 1-5.
Adult specimens of this rather handsome sea-urchin are said to reach
a test diameter of 70 mm but the finest of the 15 specimens secured by the
Velero are barely 50 mm h. d. and the longest spines scarcely exceed 40
mm. These large specimens (30-50 mm h. d.) are brown, very dark on
the test but with the spines much lighter, a real fawn brown. Small speci-
mens (15-25 mm h. d.) are dark; the smaller has the interambulacra
definitely tinged with green and the spines are dull pink or rose, brightest
on the lower surface; the larger (25 mm h.d.) has the dark interambula-
cra scarcely tinged with green and the primary spines dark violet. A series
of 5 very small Arbacias from Sechura and Independencia Bays, Peru,
are pale gray or whitish or reddish brown, with the primary spines either
colorless or with one or two light red bands. While it is not certain that
these little Arbacias are young spatuligera, there is little reason to doubt it.
Distribution.—Even including the very small Arbacias just men-
tioned, the Velero material throws no new light on the distribution of
spatuligera. The largest specimens are from San Lorenzo Island, Peru in
5 fms. The remaining material is from Sechura and Independencia Bays
in water 10 fms deep or less. So far as the V’elero collections are concerned
this Arbacia is found only in Peru.
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 247
Other records, however, extend its range to Ecuador and Chile.
Type.—Unknown.
Type locality—‘Carthagena on the west side of the Isthmus of
Darien.” (Panama)
Depth.—Shore to 10 fms.
Specimens examined.—16 specimens from 6 stations.
Tetrapygus niger (Molina)
Plate 40, Fig. 13
Echinus niger Molina, 1782, p. 200.
Tetrapygus niger A. Agassiz and H. L. Clark, p. 73.
H. L. Clark, 1910, p. 345, pl. 10, fig. 1.
Mortensen, 1935, p. 582, pl. 70, figs. 10-12.
This is one of the most characteristic of the sea-urchins of the South
American Pacific coast for it is primarily a Peruvian species and has not
yet been found north of the equator nor at the Galapagos Islands. The
Velero collection contains 274 specimens. Over 250 of these were found
along shore and are for the most part adult; few are less than 15 mm h. d.
Only at Fronton Island, near Callao, in 5 fms, were specimens of niger
dredged. The color of the adults is dull black; small specimens (under 25
mm) tend to be more brownish but not markedly so. South of Peru, niger
ranges far down on the Chilean coast but its northern limit is near Payta,
Peru. At the Lobos de Afuera Islands it is very common. Other authors
include Chile and Patagonia in the range.
T ype.-—Unknown.
Type locality —Chile.
Depth.—Shore to 5 fms.
Specimens examined.—274 specimens from 9 stations.
Family Kchinidae
Lytechinus anamesus H. L. Clark
Plate 41, Fig. 14
Lytechinus anamesus H. L. Clark, 1912, p. 254, pl. 107, figs. 7-11.
Mortensen, 1943, p. 452, pl. 23, figs. 33-40.
Apparently the commonest sea-urchin in moderate depths off the
southern California coast, this rather attractive small species was taken by
the Velero at no fewer than 155 stations at depths of 8 to 160 fms. Alto-
gether 5,090 specimens were secured, the largest 37 mm in diameter.
When very young it is light colored, often quite white, more often cream
248 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
color, but by the time it is 8-10 mm across, spots and blotches of gray or
dull green have begun to appear and in mature specimens, 20-30 mm
across, the upper surface may be more or less dull colored with spots or
blotches of cream color. The slender, more or less acicular, spines are
unicolor, generally cream color or even white, but the smaller ones are
often gray. In many specimens, however, the primary spines are yellowish
brown or even rust color, and in others they are dull gray. From some
stations the specimens are so dark as to be easily confused with pictus.
This is most apt to be the case with material from the deeper water sta-
tions, much of the test and most of the spines being dull greenish gray;
in such specimens the spines are apt to be shorter and blunter than usual,
and the resemblance to the following species may be confusing. The dif-
ferences will be emphasized under pictus. Very young specimens of
anamesus are almost pure white or very pale yellow and the primary
spines are often conspicuously long and acicular. Before the test is 10 mm
in diameter, a dull colored spot begins to appear at the upper end of each
interradial area and these are soon followed by others until in adult
specimens the whole upper surface of the test is conspicuously blotched
with the darker shade. At the same time primary spines also take on the
dull color, in more or less marked contrast with their fellows. As already
stated some individuals, and at some stations all, are so dark as to be
difficult to recognize as anamesus. Occasionally individuals are met with
of so dark a brown as to make identification difficult but the relatively
long acicular spines are distinctive in most of such cases. Not rarely
specimens 12-15 mm in diameter are met with which have developed no
pigment but are uniformly white or cream color. As a rule, however,
pigmentation begins very soon after the test is fully formed.
Distribution—The remarkably limited range of this sea-urchin is
notable for the thousands of specimens taken by the Velero are convinc-
ing proof that it is a very common species off the coast of southern Cali-
fornia and the adjoining islands, as far south as Station 283-34 off
Thurloe Head, Lower California (27° 37’ 30” N). It was not taken
north of Station 1410-41 (34° 53’ 35’ N), 3 miles east of the southern
point of Santa Rosa Island, in 17-20 fms. It was seldom taken in less
than 20 fms, the least depth being in 8-10 fms, at the southernmost sta-
tion, off Thurloe Head. The greatest depth was at 160 fms at Station
1182-40, near Catalina Island. This distribution is strikingly similar to
that of the 414 specimens on which the species was based. The Albatross
no. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 249
took them at 25 stations between 28° 12’ N and 34° 24’ 30” N at depths
of 20-113 fms, including Guadeloupe Island.
Type.—U.S.N.M. No. 32660.
Type locality Albatross Station 2930, off San Diego, 60 fms.
Depth.—8-160 fms.
Specimens examined.—5,090 specimens from 155 stations.
Lytechinus pictus (Verrill)
Plate 41, Fig. 15
Psammechinus pictus Verrill, 1867, p. 301.
Lytechinus pictus H. L. Clark, 1912, p. 258, pl. 107, figs. 12-14.
This species has undoubtedly been confused with the preceding and
the suggestion has been made that the two are identical but show great
diversity. The thousands of specimens of anamesus taken by the Velero
are supplemented by some 488 specimens of pictus, ranging from 2 to 38
mm in diameter. When young the two species are so different they could
not be confused but mature specimens may look very much alike. Young
anamesus are pure white with long, acicular white spines. As they mature
the color becomes more yellowish, and blotches of gray or dull purple
appear on the upper side of the test, but there is no hint of red or violet
on either test or spines. When adult, the slender, acute, unbanded spines
are quite different from the relatively short, rather thick, blunt spines of
pictus. In specimens of pictus less than 8 mm in diameter the primary
spines of the oral side at least are definitely reddish, ranging from dull
pink to a clear rose purple, with one or more definitely white bands. In
some cases the bands are not evident but there is more or less red or rose
purple evident on at least the distal part of the spines, but after the test
is more than 10 mm in diameter the spines are unicolor, gray, brown or
red purple. Often the basal half is nearly white, the distal half more or
less dull rose or dull yellow. In young specimens there is usually rose
purple or violet evident on the abactinal system but this soon disappears
and the whole system becomes a dull gray. Young pictus with banded
rose purple spines may be easily confused with young Strongylocentrotus
purpuratus, but in Lytechinus the color is much more pink than in pur-
puratus. In anamesus there is never any pink or rose purple shade in the
coloring of the spines but yellow, yellow brown or dull orange may be
marked; usually the shades are dull gray or olive when not yellowish or
brown; on the oral surface the spines are commonly white. In pictus there
is usually no white on the oral side but some individuals are as white as any
anamesus.
250 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 39
Fig. 9. Astropyga pulvinata (Lamarck), aboral view, x approx. 34,
p. 236
Fig. 10. Centrostephanus coronatus (Verrill), aboral view, x approx.
IZ, p. 237
PLATE 40
Fig.11. Arbacia incisa (Blainville, ? Gmelin), aboral view, x,
p. 244
Fig. 12. Arbacia spatuligera (Valenciennes), aboral view, x4, p. 246
Fig. 13. Tetrapygus niger (Molina), aboral view, x34, p. 247
PLATE 41
Fig.14. Lytechinus anamesus H. L. Clark, aboral view, x1, p. 247
Fig. 15. Lytechinus pictus (Verrill), aboral view, x1, p. 249
Fig. 16. Lytechinus semituberculatus (Valenciennes), aboral view,
xi} ip: 261
PLATE 42
Fig.17. Toxopneustes roseus (A. Agassiz), aboral view, x1, p. 262
Fig. 18. Tripneustes depressus A. Agassiz, aboral view, x%, p. 263
No. 5
CLARK: ECHINI OF WARMER EASTERN PACIFIC
PL. 39
PL. 40 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
CLARK: ECHINI OF WARMER EASTERN PACIFIC
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No. 5
PL. 44 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
in
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC PL. 4
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PL. 46
No. 5
Fig. 19.
Fig. 20.
Fig. 21.
Fig. 22.
Fig. 23.
Fig. 24.
Fig. 25.
Fig. 26.
CLARK: ECHINI OF WARMER EASTERN PACIFIC 259
PLATE 43
Loxechinus albus (Molina), aboral view, x1, p. 265
Caenocentrotus gibbosus (L. Agassiz and Desor), aboral
view, x1, p. 265
PLATE 44
Allocentrotus fragilis (Jackson), aboral view, x1, p. 276
Strongylocentrotus purpuratus (Stimpson), aboral view, x1,
D279
PLATE 45
Strongylocentrotus franciscanus (A. Agassiz), aboral view,
x¥, p. 278
Echinometra oblonga (Blainville), aboral view, x1, p. 281
PLATE 46
Echinometra VanBrunti A. Agassiz, aboral view, x%, p. 293
Clypeaster europacificus H. L. Clark, aboral view, x%, p. 294
260 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Aside from color and character of the spines, it is hard to find any
very tangible differences between the two species. On the whole the
number of coronal plates and of large tubercles is greater in pictus than
in anamesus. Specimens 25 to 30 mm h. d. having about 20 interambula-
cral plates in a column, have in pictus, 27 or 28 ambulacrals, in anamesus
about 20 or 21. In the same specimens, pictus has 4 large primary tubercles
on each coronal plate just below the ambitus, while in anamesus there are
but 3 tubercles. Furthermore, the interambulacral areas above the ambitus
obviously carry fewer tubercles in anamesus than in pictus. Mortensen
(1943, pp. 437, 450) had no specimens of pictus available for critical study
but saw Verrill’s type and cotype at New Haven. He describes them as
“light purple with radiating whitish bands, the spines bright purple, more
or less distinctly banded.” In all the hundreds of Lytechinus from the
southern California coast, no specimen with any definitely purple tint has
been seen. Neither has any individual with “‘spines bright purple, more or
less distinctly banded” been detected. The difficulty is probably due to mis-
understanding in the use of color terms. Mortensen’s “purple” is prob-
ably what to me is “rose violet,”’ but none of the Lytechinus at hand could
by any possibility be called “bright” colored.
Distribution —The 42 stations at which pictus was taken by the
Velero are widely scattered and show a perplexing distribution. Some of
the best material has been secured at Newport, Balboa, and Corona del
Mar, California, at or near low water mark and a few specimens have been
taken at or near the Channel Islands in moderately deep water, 3-39 fms.
But most of the specimens are from stations in the Gulf of California
usually in rather deep water, 40-95 fms. Much further south some very
small Echini have been taken which are best treated as pictus. Half a dozen
were secured at La Plata, Ecuador, 7-10 fms, which may possibly be
Mortensen’s panamensis but their very small size prevented satisfactory
determination. A single specimen from Gorgona Island, Colombia, in 20
fms and one from Bahia Honda, Panama, 5-8 fms offer the same difficulty.
They may possibly be panamensis but it is not wise to assert that they are.
Type.—Peabody Museum, Yale University, no. 961.
Type locality —“Cape San Lucas, California” (Grant and Hertlein,
1938, p. 25).
Depth.—Shore to 95 fms.
Specimens examined.—488 specimens from 42 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 261
Lytechinus semituberculatus (Valenciennes)
Plate 41, Fig. 16
Echinus (Psammechinus) semituberculatus L. Agassiz and Desor, 1846,
p. 368.
Lytechinus semituberculatus Verrill, 1867, p. 301.
Mortensen, 1943, pp. 456-459, pl. 26, figs.
ae G:
This well marked species is represented in the Velero collection by 863
specimens, but it should be stated at once that 570 of them are bare tests
from a sand beach adjacent to lava rock at Bindloe Island, Galapagos
Islands. The largest specimen at hand is 43 mm in diameter but most of
them are considerably less than that. The yellow green color of the
primary spines is very characteristic of well grown specimens, but very
young individuals, 5 mm or less in diameter are bright vermilion red or
paler, even a simple cream color with the median interambulacra apically
deep green. Individuals 8-15 mm h. d. are often very handsome with the
test deep green above and more or less pure white beneath; the dorsal half
of each interambulacrum may be deep red in sharp contrast with the ver-
milion red and white of the ambulacra. The apical system in such speci-
mens is greenish with a few vermilion red spots. In some young individuals
the red may be very pale except orally, and in others it may be wholly
wanting, the resulting green and white livery being soon replaced by the
brownish test and long green spines of the adult. In youth the green shades
are rich and rather dark and more or less mixed with white but with in-
creasing size the white as well as the red tends to disappear. It is hard to
believe that the really beautiful red, green, and white young urchins can
so soon and so completely lose their beauty and become the uniformly du!!
brown or blackish adult covered with uniformly yellow-green spines.
Some adults have the spines tinged at the tip with yellow brown and
rarely the color of a dried specimen may be definitely brown with little
indication of green. In the great majority of museum specimens, however,
the characteristic yellow green or light green shade of the primaries is
unmistakable.
Distribution This urchin is one of the most typical of Galapagos
echinoderms. The Velero took it at 43 stations in the archipelago ; 10 times
near Charles Island, 8 near James, 6 at Albemarle, 5 at Hood, 4 at Inde-
fatigable, 3 near South Seymour, 3 at Barrington and one each at Bar-
tholomew, Bindloe and Tower. Not a specimen has been secured at Cocos
262 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Island or anywhere along the American coast. Mortensen reports it “with
full certainty” from Clarion Island® but the Velero visited that isolated
island twice and made 17 hauls of the dredge, besides two collecting trips
along shore, and failed to find a single specimen. Certainly it should be
found on the rich collecting ground at Cocos Island, if it occurs north of
the Galapagos. If it is found on the American coast anywhere it is strange
that the V elero has not met with it. It may be added that Dr. Coker did not
find it on the Peruvian coast during his stay there (1907-1908) nor did
the Templeton-Crocker Expedition secure any specimens at Clarion
Island in 1936. There seems no adequate reason for doubting that this
sea-urchin occurs only at the Galapagos Islands.
Type.—Paris Museum.
Type locality Galapagos Islands.
Depth—Shore to 73 fms.
Specimens examined.—863 specimens from 43 stations.
Toxopneustes roseus (A. Agassiz)
Plate 42, Fig. 17
Boletia rosea A. Agassiz, 1863, p. 24.
Toxopneustes roseus Mortensen, 1903, p. 111.
Grant and Hertlein, 1938, p. 26, pl. 17, figs. 1, 2.
Mortensen, 1943, p. 483, pl. 31, figs. 1-5.
Not a very common sea-urchin of the Panamic region, this large and
striking species was not often secured by the Velero; only in 1933-34,
1934-35, 1938 and 1939 were specimens taken. Of the 22 specimens at
hand, 14 are from Panama, near Secas Islands, in 3 fms. Of the remaining
8, 3 are very small (15-23 mm in diameter) and only one is more than
half grown. Even the smallest is easily distinguished by the relatively
huge and commonly widely open globiferous pedicellariae. The extra-
ordinary appearance of the living sea-urchin is admirably shown in
Mortensen’s figures of the closely related species, pileolus (1943, pl. 26,
fig. 3 and pl. 27, fig. 2). No other sea-urchin gives any such display of
pedicellarian power as does Toxopneustes. The western Pacific species of
5 The expression “with full certainty” quoted from my Monograph (p. 459)
refers to the identification of the specimens of L. semituberculatus from Clarion
Island mentioned by Clark in his catalogue of Recent Sea-Urchins in the Coll. of the
British Museum, p. 120, the identification of other specimens being uncertain, as
pointed out both by Clark and myself. The locality Clarion Island of the specimens
in the British Museum may be erroneous. TH. MorTENSEN
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 263
the genus are also interesting for their coloration which is notably varied
and inconstant, but the Panamic species is unicolor, more or less rose or,
according to Mortensen “light purple.” Preserved specimens are dull
yellowish brown or the test may show a definitely purplish tint. Very
young specimens are yellowish, with spines and the big widely opened
pedicellariae, white. So far as preserved material is concerned the name
roseus is most inappropriate.
Distribution —The Velero took Toxopneustes at only 7 stations. The
northernmost was at Port Culebra, Costa Rica, in 3-10 fms. The southern-
most was at La Plata Isle, Ecuador, in 4 fms. The young specimens refer-
red to above, 15-23 mm in diameter, were found at Port Utria, Colombia.
A small adult was found at Braithwaite Bay, Socorro Island, in 20 fms
and a somewhat larger individual was collected at Sulivan Bay, James
Island, Galapagos Islands, at the same depth. This is the only record
from the Galapagos Islands but there is apparently no doubt about either
the locality or the identification. It is remarkable that Toxopneustes has
not been taken on the Mexican coast or at Cocos Island. It is difficult to
see why it does not occur north of Costa Rica but except for the Velero’s
specimen from Socorro Island and Mortensen’s statement of its occurrence
at La Paz, there are no records from the Mexican coast or islands. Neither
the Velero nor the Templeton-Crocker Expedition found even one speci-
men in the Gulf of California.
Type.—wNo type.
Type locality—Acapulco, Mexico.
Depth.—2-30 fms.
Specimens examined.—22 specimens from 7 stations.
Tripneustes depressus A. Agassiz
Plate 42, Fig. 18
Tripneustes depressus A. Agassiz, 1863, p. 24.
Mortensen, 1943, p. 498, pl. 35, figs. 1, 2; pl.
38, fig. 9.
The largest regular sea-urchin of the western coast of tropical America
this species reaches a horizontal diameter of 150 mm. There is great
diversity in form, for while the height is usually a little more than half the
diameter, the specimen 150 mm horizontally is only 72 mm vertically,
and a specimen 138 mm across is only 65 mm high. On the other hand a
specimen 127 mm across is 83 mm high. The height of the test thus ranges
264 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
from 47 to 65 per cent of the diameter. Growth changes from 30 mm h. d.
to 150 mm are chiefly in the number of secondary and miliary tubercles
(and spines, of course). In the young individuals, there are two columns
of primary tubercles in the ambulacra and two in the interambulacra, with
secondary and miliary tubercles present between them in the interambula-
cra in not very well-marked series. On the ambulacra there are few
tubercles of any importance in the interporiferous area save for one column
on each side. In an adult, 100 mm or more in diameter, there are 10-12
well-spaced columns of primary tubercles and many well-scattered smaller
tubercles in the interambulacra. The ambulacra have undergone a more
marked change; 4-6 imperfect columns of primary tubercles occupy the
interporiferous area, and the poriferous areas have themselves widened
markedly and carry 3 or 4 columns of tubercles mixed with irregularly
scattered tube-feet between an inner and outer column of tube-feet which
sharply delimit the area. The scattered tube-feet are much more numerous
on the inner side of the area than on the outer. As for color, the dry speci-
mens are very unattractive, as they were obviously prepared in haste. The
test is almost black but with purple tinge more or less evident, and the
spines are pale brown or brownish-white. One small adult in alcohol gives
a better idea of the color in life. The test is purple, lightest on the porifer-
ous areas, darker and duller on the interambulacra and on the interporif-
erous areas of the ambulacra. The spines are white in marked contrast to
the test.
Distribution—The 63 specimens of depressus brought in by the
Velero are, with only two exceptions, from outlying islands. The excep-
tions are from the shores of Secas Islands, Panama. Of the other 61 speci-
mens, 7 are from the Galapagos Islands (Albemarle, Tower, Chatham,
Charles, and Gardner), all from along shore except the half grown speci-
men from near Gardner Island which was dredged in 12 fms. Of the
remaining specimens 43 are from Socorro and 11 from Clarion, all secured
by collecting along shore. The type locality is said to be Guaymas, Mexico,
but it is notable that neither the Velero nor the Templeton-Crocker
Expedition found any Tripneustes in the Gulf of California nor on the
Mexican coast.
Type.—No type.
Type locality —Guaymas, Mexico.
Depth.—Shore to 12 fms.
Specimens examined.—63 specimens from 10 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 265
Family Strongylocentrotidae
Loxechinus albus (Molina)
Plate 43, Fig. 19
Echinus albus Molina, 1782, p. 200.
Loxechinus albus Desor, 1858, p. 136.
Strongylocentrotus albus H. L. Clark, 1910, p. 347, pl. 12, fig. 1.
This large sea-urchin of the western coast of South America is exten-
sively used for food along the Chilean coast as Tripneustes is in the West
Indies. In both cases it is the voluminous gonads which are eaten, com-
monly roasted in the “half-shell.” The largest specimen taken by the
Velero is somewhat less than 100 mm h. d. but specimens 120 mm in
diameter are reported. Altogether only 23 specimens were taken. The
smallest is 16 mm in diameter, the largest 84. The parasitic crab,
(Pinnaxoides), which so often lives in the periproct of this sea-urchin and
the following, Caenocentrotus gibbosus, is evidently present in at least 9
cases. The color is pretty uniformly green in spite of being often more or
less bronzed over with red-brown. In specimens less than 20 mm h. d. the
tips of the spines may be quite red. But in none of the specimens that I have
ever seen is there any obvious reason why the specific name albus should
have been attached to this sea-urchin.
Distribution —The Velero met with this urchin only three times—
first in 1935 along the shore in Independencia Bay, again in 1938 in San
Juan Bay and Independencia Bay, and thirdly, a few days later further
north, among the Lobos de Afuera Islands. This extends the range from
the Lobos de Afuera Islands, northern Peru (6° 53’ 50’), southward to
the Straits of Magellan.
Type.—Unknown.
Type locality —Chile.
Depth—Shore to 40 fms.
Specimens examined.—23 specimens from 6 stations.
Caenocentrotus gibbosus (L. Agassiz and Desor)
Plate 43, Fig. 20
Echinus (Toxopneustes) gibbosus L. Agassiz and Desor, 1846, p. 367.
Caenocentrotus gibbosus H. L. Clark, 1912, p. 348.
Grant and Hertlein, 1938, p. 29, pl. 10, figs. 3, 4.
Mortensen, 1943, p. 321, pl. 34, figs. 3, 4; pl.
G3ishess. 7, o. Ly US, to.
This is another South American urchin which has not followed the
coast north of the equator but unlike the preceding species it has followed
266 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 47
Fifi. 27. Clypeaster ochrus H. L. Clark, aboral view, x%, p. 297
Fig. 28. Clypeaster rotundus (A. Agassiz), aboral view, x34, p. 296
PLATE 48
Fig. 29. Clypeaster speciosus (Verrill), aboral view, x24, p. 308
Fig. 30. Clypeaster elongatus, new species, aboral view, x34, p. 308
PLATE 49
Fig. 31. Clypeaster elongatus, new species, oral view, x34, p. 308
Fig. 32. Dendraster excentricus Eschscholtz, aboral view, x¥%, p. 310
PLATE 50
Fig. 33. Clypeaster elongatus, new species, lateral view, x34, p. 308
Fig. 34. Dendraster laevis, new species, aboral view, x%, p. 312
Fig. 35. Dendraster laevis, new species, oral view, x%
Fig. 36. Dendraster laevis, new species, aboral view, smooth surface,
x%
PL. 47
ECHINI OF WARMER EASTERN PACIFIC
CLARK
NO. D
PL. 48 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PL. 49
ECHINI OF WARMER EASTERN PACIFIC
CLARK
No. 5
ALLAN HANCOCK PACIFIC EXPEDITIONS
ECHINI OF WARMER EASTERN PACIFIC
CLARK
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ECHINI OF WARMER EASTERN PACIFIC
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PL. 54 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
No. 5
Fig. 37.
Fig. 38.
Fig. 39.
Fig. 40.
Fig. 41.
Fig. 42.
Fig. 43.
Fig. 44.
Fig. 45.
CLARK : ECHINI OF WARMER EASTERN PACIFIC 215
PLATE 51
Dendraster mexicanus, new species, aboral view, rough
surface, x1, p. 313
Dendraster mexicanus, new species, aboral view, smooth
surface, x1
Dendraster mexicanus, new species, oral view, x1
PLATE 52
Dendraster rugosus, new species, aboral view, x4, p. 318
Dendraster rugosus, new species, oral view, x4
PLATE 53
Encope grandis L. Agassiz, aboral view, x%, p. 325
Encope perspectiva L. Agassiz, aboral view, x1, p. 326
PLATE 54
Encope wetmorei A. H. Clark, aboral view, x94, p. 326
Encope laevis, new species, aboral view, x%, p. 327
276 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
the equator to the Galapagos Islands. Just how it reached the islands
cannot be said but judging from the material at hand conditions are not
as favorable for it there, as the Peruvian specimens are conspicuously larger
and more robust than those from the islands. Of the 82 specimens in the
Velero collection, 69 are from the Lobos de Afuera Islands, Peru. They
range from 30-68 mm in diameter. The tests are usually brown with the
spines a deep bronze-green, their tips more or less reddish, but some indi-
viduals are distinctly greenish rather than brown. All are more or less
deformed at the abactinal system by the presence of the parasitic crab,
Pinnaxoides. Of the remaining 13 specimens, 2 small individuals are from
the northernmost station known for Caenocentrotus, the shore of La
Plata Isle, Ecuador. They are about 25 mm h. d. and have notably long
primary spines. Both are infested with the parasitic crab. Another pair are
from Fronton Island, near Callao, Peru, one a mere fragment, the other a
good adult, like those from Lobos de Afuera. The other 9 specimens are
from the Galapagos Islands, 6 from the reef north of Tagus Cove, Albe-
marle Island, one from Tagus Cove, one from Academy Bay, Indefati-
gable Island, and one from Sulivan Bay, James Island. They range from
24 to 41 mm in diameter, the primary spines are relatively stout, and ap-
parently none are infested with the crab, Pinnaxoides. It should be added,
however, that other specimens of Caenocentrotus from the Galapagos have
shown the presence of the crab. Except for the smaller size and shorter
spines, the Galapagos Caenocentrotus is very similar to the Peruvian.
Distribution —Guayaquil, Ecuador to northern Chile; Galapagos
Islands.
Type.—Paris Museum.
Type locality Galapagos Islands.
Depth.—Littoral.
Specimens examined.—82 specimens from §8 stations.
Allocentrotus fragilis (Jackson)
Plate 44, Fig. 21
Strongylocentrotus fragilis R.’T. Jackson, 1912, p. 128.
H. L. Clark, 1912, p. 354, pl. 113, figs. 3-6.
Allocentrotus fragilis Mortensen, 1942, p. 232.
Mortensen, 1943a, pp. 254-255, pl. 30, figs. 10-17.
If a sea-urchin could ever be called a “beautiful” creature, there is no
doubt that this species would be a first choice for the title. The delicacy of
its test and spines is remarkable and the beauty of their coloring is indis-
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 277
putable. Although large and strikingly colored, no specimens seem to have
been collected until well into the twentieth century. It is probable that
specimens secured accidentally by dredge or trawl were so badly damaged
that they were not brought ashore, and the few that escaped that fate were
not attractive enough to lead anyone, not a specialist in sea-urchins, to give
them any attention. The urchin is so common in the limited area it inhabits
that the Velero has brought in altogether 240 specimens from 59 stations,
ranging in horizontal diameter from 2 mm to just over 100. The height of
the low test is usually less than half its diameter, a specimen 80 mm h. d.
having a height of about 35 mm. Young individuals are quite flat, a speci-
men 30 mm across being only 11 mm high. The test is remarkably fragile,
only a millimeter thick in a well-grown adult. As a rule the colors are
light and often bright. A normal specimen has the test rose red or pink with
a more or less violet cast on the upper side, becoming more and more violet
orally. The small spines are much the same color but the larger ones are
orange or vermilion at base becoming very light, almost white on the distal
part. The conspicuous pedicellariae are nearly white. Occasional speci-
mens are much darker, with test dusky purple and dull orange, and the
spines dull gray or violet or even brown. When freshly caught or well pre-
served, either in alcohol or dry, the slender, delicate spines varied, more or
less bright colors compel admiration.
Distribution —The V elero has taken this fine sea-urchin at 59 stations
in depths of 39-380 fms off the coast of southern California. The material
collected by the Albatross on which the species was based, was taken
between northern Washington (47° 46’ N) and the Mexican boundary
(32° 32’ 30”) in depths of 48 to 417 fms. The Velero took no specimens
at any station north of 34° 06’ 30” or south of 32° 34’ 50’, in depths of
40 to 380 fms, except one half grown specimen labeled as from Station
1246-41, which is off Cedros Island, Lower California, in 81-83 fms.
(There is no good reason for doubting the label with this specimen but it
is odd that no other specimens of fragilis have been taken in waters south
of the United States.) It may properly be considered a definitely North
American sea-urchin. According to other authors the range is from Van-
couver Island to Lower California.
Type.—R. T. Jackson Collection No. 838.
Depth.—39-640 fms.
Type locality —Catalina Island.
Specimens examined.—240 specimens from 59 stations.
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
bo
“I
ioe)
Strongylocentrotus franciscanus (A. Agassiz)
Plate 45, Fig. 23
Toxocidaris franciscana A. Agassiz, 1863, p. 22.
Strongylocentrotus franciscanus A. Agassiz, 1872a, p. 163.
Mortensen, 1943a, p. 242, pl. 28, figs.
1-7; pl. 29, figs. 1-4.
There are in the Velero collection 343 specimens of Strongylocentrotus
from 24 stations which are here referred to this conspicuous sea-urchin.
In point of size it rivals T'ripneustes. The latter reaches perhaps a greater
diameter of the test, 150 mm as against 146 for the present species but the
latter has such long, heavy spines, 50 mm or even more in some individuals,
it is evidently the largest of the west coast Echini. Young individuals with
a test diameter of 10-15 mm have the relatively slender spines about two-
thirds as much, while adults with a diameter of 100 mm or more are satis-
fied with spines only a third of that. In two specimens at hand from Point
Vicente, California, which are rather more than 100 mm in diameter the
number of coronal plates (and hence the number of primary spines) is
considerably more than normal and the length of the primaries is markedly
less, only 20-30 mm. The appearance of these specimens is thus strikingly
different from normal specimens but the differences are probably associated
with some unusual feature of their habitat, such as excessively strong surf
or tidal currents. Further collecting at Point Vicente and study of local
conditions might explain this curious anomaly.
The color of normal franciscanus when young is a light fawn brown
but with growth the color darkens and in most individuals it becomes a
very deep purple or even almost black. Curiously enough many individuals
instead of becoming very dark remain light colored until half grown and
are then a dull and pale “old rose” or pale violet. With increased age and
size the dark color becomes predominant and the big adults are nearly if
not quite black. Other individuals never become black at all but even
when full-grown are light-colored, a peculiar shade of brown, lightest at
the base of the spines, darkest near the tips. Curiously enough such speci-
mens if placed in a dilute solution of formalin (4 per cent) to which a
pinch of corrosive sublimate has been added turn bright green, strikingly
different from any normal shade of the living urchins. On being dried, the
green disappears more or less and the urchin becomes brown once more but
darker, especially on the distal portion of the spines. It is conceivable that
the colors are an indication or a sex dimorphism and an investigation into
the matter of a possible correlation of color and sex might prove rewarding.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 279
Another remarkable thing affecting the color is undeniable hybridiza-
tion of franciscanus and purpuratus. ‘The hybrids when young have the
primaries quite purple or bright violet and shades of these colors persist as
the animal grows. The hybrids are distinguishable from purpuratus by the
smaller number and much greater size of the primary spines, while their
unusual color distinguishes them from franciscanus. Of course, many speci-
mens are so nearly intermediate, their identification becomes a matter of
opinion. A careful study of this hybridization would surely be rewarding.
Distribution —There is no doubt that franciscanus is characteristic of
the coastal region of the western United States and Canada, and Morten-
sen reports its occurrence on the Alaskan coast and even in northern Japan.
How far south on the American coast its range extends is still somewhat
uncertain. Mortensen says Cedros Island, Lower California, is the south-
ern limit and two small specimens in the V elero collection from that island
and two from Thurloe Bay, near the island, confirm that opinion. There
is, however, in the present collection a very good adult specimen from
“Shore collecting at Petatlan Bay, Mexico,” Lat. 17° 31’ 45”, but this
seems quite improbable and needs confirmation. In the vicinity of Newport
and Balboa, California, franciscanus is common and grows to a large size,
but the largest specimens in the Velero collection are from shore collecting
between Cape Arago State Park and Yaquina Head Light on the Oregon
coast.
Type.—M.C.Z. no. 1686 (Cotype).
Type locality—San Francisco, California.
Depth.—Shore to 50 fms.
Specimens examined.—343 specimens from 24 stations.
Strongylocentrotus purpuratus (Stimpson)
Plate 44, Fig. 22
Echinus purpuratus Stimpson, 1857, p. 86; 1857a, p. 526.
Strongylocentrotus purpuratus A. Agassiz, 1872a, p. 165.
Mortensen, 1943a, p. 236, pl. 26, figs. 1-8.
Like franciscanus, this species is characteristic of the western coast of
North America. It is even more common, the present collection contain-
ing over 500 specimens. They show a diversity in spinulation which is more
extraordinary than anything shown by any other American sea-urchin.
The normal condition is the presence of a double series of primary spines
in each ambulacrum and interambulacrum; these spines are about 10 mm
long, rather stout and quite acute. Each ambulacral plate carries an addi-
tional primary spine, not so large at each end of the plate. Each inter-
280 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
ambulacral plate carries a similar but slightly larger spine at each end.
Miliary spines are fairly numerous among these primary and secondary
spines so the test is normally very completely but not densely covered. The
abactinal system of plates is similarly well covered but the peristome is
quite free from spines. Many, perhaps most adult specimens differ from
this typical spinulation in one of two ways. On the one hand the primary
spines are disproportionately bigger, 15 mm or more in length and more
than a millimeter thick basally. In extreme cases the primaries are over 20
mm long and 2 mm thick at base. On the other hand, the primary spines
increase markedly in number but decrease in thickness and length; they are
6 or 7 mm long and scarcely more than half a millimeter in thickness. This
diminution of the primaries occurs for the most part above the ambitus
and may continue to such an extent that there are no normal primaries on
the upper half of the test, but the whole abactinal surface is covered by a
fairly dense coat of small secondary and miliary spines. Whether this
condition is due to some feature of the environment such as constant heavy
surf or to a diseased condition resulting from parasites or some deficiency
of the environment, we do not know, but the result is a sea-urchin whose
identity would be difficult to determine without a good series of connecting
links.
Aside from environmental conditions modifying structure and appear-
ance, there is no doubt that hybridization with franciscanus is a frequent
if not a constant factor in the life history of purpuratus. As the two species
often occupy the same regions and even the same environmental areas, it is
not strange that cross fertilization often occurs and as a result many indi-
viduals are found whose correct identification is too much for even an
experienced taxonomist. Color is the most useful factor in determining the
species for typical purpuratus ; it is at first green and then becomes purple
and when large enough to be called adult it is practically entirely purple
as the name indicates. The larger species, franciscanus, is never purple nor
does it have any trace of such purple as characterizes purpuratus. Young
individuals are light brown, almost or quite a fawn brown, and usually
this becomes increasingly dark, until very large adults are a very dark
violet brown, sometimes almost if not quite black. But there is never any
indication of the purple of purpuratus. Hybrids usually show some indi-
cation of purple on the primary spines. A second indication of hybridi-
zation is seen in the stoutness of the primary spines. While very large
individuals of purpuratus may have unusually large primary spines they
are never equal to the normal spines of franciscanus. Oftentimes specimens
occur which have the general form and appearance of purpuratus but the
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 281
primaries are disproportionately stout yet have a purplish tint. Such indi-
viduals are probable hybrids. A critical study, in a marine laboratory, of
cross fertilization between the two species would yield interesting, perhaps
important results.
Distribution—The range of purpuratus is given by Mortensen as
from Cedros Island, Lower California to Vancouver Island, with a
dubious southern limit at La Paz and an equally dubious northern limit at
Sitka. He considers it strictly littoral with 20 meters apparently as the
bathymetric limit. The /elero has secured specimens at 46 stations, of
which the northernmost was at Boiler Bay, Oregon, the southernmost at
Petatlan Bay, Mexico, Lat. 17° 32’ N. As the only other stations south of
the United States at which purpuratus was secured are at or near Cedros
Island, the Petatlan Bay record needs confirmation. Neither the 4 /batross
nor the Templeton-Crocker Expedition took purpuratus in the Gulf of
California, or near La Paz. The specimens labeled Petatlan are undoubt-
edly purpuratus but, as in the case of franciscanus, the locality is hard to
credit. As for the bathymetrical range of the species, there can be no doubt
that it is greater than supposed for the Velero took it at several stations in
water ranging from 15 to 88 fms. These were young individuals, however,
and it is probable the normal habitat of the species is above 15 fms.
Type.—U.S.N.M. No. 2495.
Type locality—San Francisco, California.
Depth.—Littoral to 88 fms.
Specimens examined.—498 specimens from 46 stations.
Family Echinometridae
Echinometra oblonga (Blainville)
Plate 45, Fig. 24
Echinus oblongus Blainville, 1825, p. 95.
Echinometra oblonga Blainville, 1834, p. 225.
Echinometra mathaei oblonga Mortensen, 1943a, p. 393, pl. 48, figs. 1-20.
The status of this species or form of Echinometra is so perplexing that
no two writers seem to have just the same opinion on the matter. Morten-
sen’s treatment of the problem is entitled to the greatest respect, for his
wide experience, extraordinary industry and unusual skill in wrestling
with such a situation are beyond compare. The only reason for not accept-
ing his conclusion outright is that mathaei does not occur in the Eastern
Pacific whereas oblonga does, and there it intergrades with VanBrunti
in a perplexing way. The Velero collection contains Echinometras which
seem to be oblonga in its most typical form, high, elongated, well-rounded
282
Fig. 46.
Fig. 47.
Fig. 48.
Fig. 49.
Fig. 50.
Fig. 51.
Fig. 52.
Fig. 53.
ALLAN HANCOCK PACIFIC EXPEDITIONS
PLATE 55
Encope laevis, new species, oral view, x94, p. 327
Encope micropora L. Agassiz, aboral view, x%, p. 329
PLATE 56
Encope cocosi, new species, aboral view, x%, p. 330
Encope cocosi, new species, oral view, x34
PLATE 57
Encope galapagensis A. H. Clark, aboral view, x94, p. 331
Encope galapagensis A. H. Clark, oral view, x%
PLATE 58
Encope irregularis, new species, aboral view, x%, p. 332
Encope irregularis, new species, oral view, x%
VOL. 8
ECHINI OF WARMER EASTERN PACIFIC
CLARK
wy
NO.
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PL. 56
ECHINI OF WARMER EASTERN PACIFIC
CLARK
7 de
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ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PL. 58
PL. 59
ACIFIC
ECHINI OF WARMER EASTERN P
CLARK
NO. 5
VOL. 8
ALLAN HANCOCK PACIFIC EXPEDITIONS
PL. 60
PL. 61
R EASTERN PACIFIC
NI OF WARME
ECHI
CLARK
ALLAN HANCOCK PACIFIC PEDITIONS
No. 5
Fig. 54.
Fig. 55.
Fig. 56.
Fig. 57.
Fig. 58.
Fig. 59.
Fig. 60.
Fig. 61.
CLARK : ECHINI OF WARMER EASTERN PACIFIC
PLATE 59
Encope ecuadorensis, new species, aboral view, x4, p. 333
Encope ecuadorensis, new species, oral view, x%
PLATE 60
Encope fragilis, new species, aboral view, x34, p. 335
Encope fragilis, new species, oral view, x%%
PLATE 61
Encope insularis, new species, aboral view, x4, p. 336
Encope insularis, new species, oral view, x4
PLATE 62
Mellita longifissa Michelin, aboral view, x1, p. 337
Cassidulus pacificus A. Agassiz, aboral view, x3, p. 338
291
292 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
test with 4 or 5 rather large ambulacral pores in each arc, and short, stout,
pointed primary spines of an almost black color. These spines tend to point
upward or definitely forward or backward and give the impression that
the urchins live in crevices or fissures in or between rocks. They do not
bore holes (as Echinostrephus does) but it is possible that they enlarge
crevices into which they fit themselves to some extent. Nearly all speci-
mens have the primary spines pointing more or less forward or backward,
parallel to the longer axis of the test. The largest specimen is 45 mm long
(not including spines), 39 mm wide and 29 mm high. If the spines are
included the corresponding measurements are roughly 80 x 55 x 40 mm.
The color of all the Velero specimens is uniformly very dark but not
black as one is inclined to think at first sight. It is rather a very deep purple,
as dark in the very young specimens as in the adults. Specimens of the
following species (VanBrunti) are often nearly as dark but the purple
tinge is more evident especially on the oral side. In the shape of the test,
this species is clearly more elongated and higher than the Panamic, and
the pore-pairs are definitely fewer, only 4 or 5, as against 6 to 8. Hence
there is no serious difficulty in identifying oblonga as it exists in the
Eastern Pacific. The trouble comes in the Western Pacific where the
ubiquitous mathaei occurs so abundantly that it is dificult to draw any
definite line between some of its very dark forms and the similarly dark
oblonga. I think it must be agreed that oblonga is a variety gradually
differentiating into a quite distinct species and in the Eastern Pacific it
has attained that distinction.
Distribution —The Velero has found oblonga common at Sulphur
Bay, Clarion Island, and at Braithwaite Bay, Socorro Island. A fair-sized
adult was taken along shore at Bindloe Island, in the northern Galapagos
Islands. Along the western side of South Seymour Island, 7 very young
but quite typical oblonga were collected and at Cartago Bay, Albemarle
Island, an adult specimen was found. But not a single specimen has been
secured at Cocos Island or along the Mexican or Costa Rican coast. The
indications are that oblonga has come from the west and established itself
firmly at Clarion Island. From there it has spread to its near neighbor
Socorro, and from these two islands it is extending its range southward
into the Galapagos Islands but is not yet common there and has appar-
ently reached a southern limit at Albemarle Island.
T ype.—Unknown.
Type locality—Unknown.
Depth.—Shore and very shallow water.
Specimens examined.—85 specimens from 12 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 293
Echinometra VanBrunti A. Agassiz
Plate 46, Fig. 25
Echinometra VanBrunti A. Agassiz, 1863, p. 21.
Heliocidaris stenopora H. L. Clark, 1912, p. 351, pl. 95, figs. 18-22; pl.
104, figs. 1-3; pl. 110, figs. 4, 5.
Echinometra VanBrunti Mortensen, 1943a, p. 373, pl. 45, figs. 1-3.
This is apparently the commonest littoral sea-urchin of the tropical
Eastern Pacific. The Velero collection contains 658 specimens taken at 92
stations. They range in size from young ones, nearly or quite circular, 5 to
10 mm in diameter, to large adults with a long axis exceeding 70 mm.
Normal large adults have the lesser diameter nine-tenths of the longer,
and the height is very generally about one-half the length. But there is
considerable diversity and individuals having the normally lesser axis
nearly or quite equalling the longitudinal axis are not very rare. There
seems to be no doubt that Heliocidaris stenopora H. L. Clark is based
on a large Echinometra Van Brunti as A. H. Clark, Ziesenhenne and
Mortensen have recently pointed out. In the Velero collection there are
some specimens in which the ambitus is nearly circular but they are com-
monly less than 40 mm in diameter and not one would be identified as
stenopora, as the tuberculation of the test is not coarse enough and the
primary spines are not sufficiently stout. Mortensen inclines to recognize
a variety rupicola although it occurs in the same areas as typical VanBrunti
but the hundreds of specimens in the Velero collection do not, in my
opinion, warrant the recognition of such a form.
It was hoped and rather expected that among the Echinometras in the
present collection, there would be from the Galapagos and Socorro Islands,
some specimens of E. insularis but not one typical example has been found.
Some specimens from Costa Rica have only 5 pore-pairs in the ambulacra
at the ambitus but they have 6 pairs abactinally and the general appearance
and spinulation is like that of YanBrunti from the same region. Unfor-
tunately there is but a single young individual from Clarion Island, and
only two lots from Socorro. This series includes one very large adult and
32 small ones and they are typical VanBrunti. There are 78 specimens
from some 20 stations in the Galapagos but none of them represent
insularis, which justifies the suspicion that that supposed species is not
really valid.
The color of VanBrunti is remarkably uniform although the shade
shows no little diversity. The basic color may be designated as violet
especially when seen from below. There is a tendency on the one hand
294 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
towards gray and on the cther towards black. As a rule specimens from
any one locality are much alike in color though large specimens may be
darker than small ones. There is never any white nor any hint of green,
but suggestions of a red violet are sometimes evident, and very young speci-
mens (under 15 mm h. d.) may have a very definite red orange or a dull
yellow tint around the peristome.
Distribution—According to Mortensen, this sea-urchin occurs as far
north as “off Central California,” but the Velero has not found it north
of the Mexican boundary. Indeed she has not taken it anywhere on the
western side of Lower California, although she has found it at various
points in the Gulf as far north as Guaymas and Angel de la Guardia
Island. It is common on the Mexican and Costa Rican coasts and in
Panamic waters. Further south it has been taken by the Velero at various
points on the coasts of Colombia, Ecuador, and Peru as far south as the
Lobos de Afuera Islands. At the Galapagos Islands, it is quite common,
being taken at more than 20 stations though never in large numbers. Some
typical specimens were secured at Cocos Island and one good series was
collected at Braithwaite Bay on Socorro. A single young specimen was
collected at Clarion Island, but the Echinometras of Clarion Island need
further collecting and study.
Type—M.C.Z. no. 1851 (Cotype).
Type locality Acapulco, Mexico.
Depth.—Shore to 29 fms.
Specimens examined.—658 specimens from 92 stations.
Order EXOCYCLOIDA
Family Clypeastridae
Clypeaster europacificus H. L. Clark
Plate 46, Fig. 26
Clypeaster europacificus H. L. Clark, 1914, p. 27, pls. 129, 130, 131, 136,
fete hs
The 102 specimens of this fine Clypeaster taken by the Velero form a
very complete series from the babies 16 x 15 mm to the full grown adults
just exceeding 200 mm in length. The largest specimen is 202 mm long
by 200 mm wide and 30 mm high; the test margin is about 6 mm thick.
In most specimens the width of the test is not so great proportionately, a
normal adult is 184 by 174 mm with the height 27 mm. The young are
nearly or quite circular and many individuals continue so throughout life.
Some, however, elongate appreciably; the longest of the present series
have the width nine-tenths of the length. As a rule the height is about 15
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 295
per cent of the length but occasionally it is appreciably more, in extreme
cases it may be 30 per cent. The general texture of the test is not very
stout and hence injuries to the margin are rather frequent resulting in
more or less conspicuous deformities. One specimen about 113 mm in
length, with the width somewhat less, has the central portion of the
petaloid area abruptly raised and flattened, resulting in a curious hat-
shaped form as seen from the side.
Color undergoes very considerable change with the growth. The
youngest specimens are pale red violet, almost pink, above and cream
color or nearly white below. With increasing size the color deepens. The
upper surface becomes a deeper and more purple pink, violet, deep violet,
brown violet, brown either yellowish or violet tinted, until in old speci-
mens the very dark brown of the adult is reached. The ambulacral
areas are often of a different shade so that the petals stand out more or
less clearly. The cream-colored lower surface of the very young, becomes
brownish yellow, yellow brown and ultimately a deep brown. In very old
specimens there is a tendency for the ambulacral areas to darken and they
may become more or less black at least along the midline. The dark shades
of dried adults show considerable diversity, some specimens being quite
gray. Young specimens, 25 mm or so in diameter may become deep violet
above quite early, but occasionally remain light pink until sometime after
that size is reached.
Distribution —The original material on which the species was based
came from the Gulf of California, Clarion Island, Hood Island in the
Galapagos, and Panama. The Velero has extended that range very little,
except to the south; the present collection contains a nice set of 12 small
specimens, 25-70 mm long, from off La Plata Island, Ecuador, in 45-55
fms. In the Galapagos area only once was europacificus met with—a single
specimen in Academy Bay, Indefatigable Island in 10-25 fms. It was not
found at Clarion or Socorro Islands. From Panama, Costa Rica and
Mexico a number of specimens were secured but the bulk of the material,
more than 60 specimens, was taken in the Gulf of California, at various
points up to Angel de la Guardia Island. The bathymetric range of this
Clypeaster seems to be from 10 to 95 fms.
Type—U.S.N.M. No. 34226.
Type locality —Bay of Panama, 7° 57’ N, 78° 55’ W, 33 fms.
Depth.—10-95 fms.
Specimens examined.—102 specimens from 24 stations.
(Very likely this species is identical with Pfeffer’s Alexandria magnt-
fica. Th. Mortensen)
296 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Clypeaster rotundus (A. Agassiz)
Plate 47, Fig. 28
Stolonoclypus rotundus A. Agassiz, 1863, p. 25.
Clypeaster rotundus A. Agassiz, 1872a, p. 100.
H. L. Clark, 1914, p. 38, pls. 132, 133.
Although adult specimens of this Clypeaster are easily distinguished
from the preceding, the young, before reaching a length of 25 mm are very
difficult to determine. They are nearly or quite circular, or a little longer
than wide, and very flat. The lower surface, at first nearly white becomes
yellowish, and as a rule is soon yellowish-brown, as in europacificus. The
subsequent color changes are similar in the two species but the body form
is usually characteristic, for ewropacificus has the interradial margins more
or less evidently concave or at least straight, resulting in a pentagonal
form with the length only a little, if at all, greater than the breadth. In
rotundus the interradial margins are more or less convex, and the outline
may be nearly circular, but as a rule the length is somewhat greater than
the width. The largest specimen in the present collection is 175 mm by
160; the height is 27 mm, but a second specimen with the same length and
breadth is only 20 mm high. The most nearly circular adult at hand is 146
mm long by 137 mm wide, but a specimen 37 mm long is 36 mm wide.
Distribution —There are 96 specimens in the Velero collection from
35 stations, well scattered along the coast from Santa Elena Bay, Ecuador
to Georgia Island at the upper end of the Gulf of California. There are
no records from the western coast of Lower California nor from anywhere
above the Mexican boundary. There are 9 specimens from Clarion Island
but none from Socorro. From Cocos Island there are two small specimens,
dredged in relatively deep water, 30-50 fms. There are 20 specimens from
9 stations in the Galapagos Islands, but chiefly from Tagus Cove, Albe-
marle Island. Some were taken in shallow water along shore but they
were mostly dredged in 8-50 fms.
Type-—M.C.Z. no. 2188.
Type locality —Acapulco, Mexico.
Depth.—Shore to 50 fms.
Specimens examined.—96 specimens from 35 stations.
no. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 297
Clypeaster ochrus H. L. Clark
Plate 47, Fig. 27
Clypeaster ochrus H. L. Clark, 1914, p. 30, pl. 141, figs. 1-3.
There can be no doubt of the close relationship between this species
and speciosus but after a prolonged study of all the Velero Clypeasters, it
seems best to maintain ochrus as at least an incipient species. Typical speci-
mens in good condition are easily recognized, but immature or badly pre-
served individuals cause trouble. Moreover well-preserved specimens of
the two species may be so much alike in color that it is very hard to deter-
mine to which species they belong but the greater height of the test with
its somewhat greater concavity can usually be relied upon to distinguish
ochrus; in speciosus, the height is about one-fifth of the length, in ochrus
it is one-fourth. In the Velero collection, there are 47 Clypeasters which
may be referred to ochrus. They range from 5 to 114 mm in length but
the most typical is 104x 96 mm with a height of 28 mm, the mouth is
sunken 12 mm below the test margin. The characteristic color is a yellow
brown with the petaloid areas and the lower surface dark brown in rather
marked contrast, but most of the specimens from Cocos (Nuez) Island,
Costa Rica, Panama and Ecuador are chocolate brown or purple brown of
a more or less deep shade, rarely, however, as deep as in speciosus. One
young specimen (48 x 44 mm) from James Island, Galapagos, in 10 fms
is red purple, rather dull above but quite bright orally. The smallest speci-
men (5 mm), also from James Island, is a uniform light brown while the
next largest (8 mm), from Ecuador, has the upper surface red violet.
Distribution—The most typical specimens are from Ecuador,
Colombia and the Secas Islands, Panama, but very good specimens are
from Albemarle Island in the Galapagos. A large number of Clypeasters
dredged at Nuez Island, off Cocos Island, in 31-50 fms are best referred
to this species although the only adult from this locality is very dark
colored. It has, however, the proportions and general appearance of ochrus.
No specimens of ochrus were taken north of Costa Rica. Records from
other authors extend the known distribution as far north as La Paz, Lower
California.
Type.—M.C.Z. no. 4740.
Type locality—Acapulco, Mexico.
Depth.—Shore to 50 fms.
Specimens examined.—47 specimens from 15 stations.
298 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 63
Fig. 62. Agassizia scrobiculata Valenciennes, aboral view, x%, p. 339
Fig. 63. Agassizia scrobiculata Valenciennes, lateral view, x¥
PLATE 64
Fig. 64. Brisaster townsendi (A. Agassiz), aboral view, x34, p. 340
Fig. 65. Moira clotho (Michelin), aboral view, x53, p. 341
PLATE 65
Fig. 66. Brissopsis pacifica (A. Agassiz), aboral view, x4, p. 341
Fig. 67. Brissopsis pacifica (A. Agassiz), oral view, x4
PLATE 66
Fig. 68. Plagiobrissus pacificus H. L. Clark, aboral view, x1, p. 342
Fig. 69. Plagiobrissus pacificus H. L. Clark, oral view, x1
Fig. 70. Plagiobrissus pacificus H. L. Clark, lateral view, x1
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VOL. 8
ALLAN HANCOCK PACIFIC EXPEDITIONS
PL. 68
NO. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC PL. 69
VOL. 8
ALLAN HANCOCK PACIFIC EXPEDITIONS
pL. 70
NO. 5
Fig. 71.
Fig. 72.
Fig. 73.
Fig. 74.
Fig. 75.
Fig. 76.
Fig. 77.
Fig. 78.
Fig. 79.
Fig. 80.
Fig. 81.
CLARK: ECHINI OF WARMER EASTERN PACIFIC 307
PLATE 67
Ibiobryssus coelus H. L. Clark, aboral view, x54, p. 343
Idiobryssus coelus H. L. Clark, oral view, x84
Idiobryssus coelus H. L. Clark, lateral view, x24
Brissus latecarinatus (Leske), aboral view, x4, p. 343
PLATE 68
Brissus latecarinatus (Leske), aboral view with spines, x1,
p. 343
Meoma grandis Gray, aboral view, rough, x’, p. 344
Meoma grandis Gray, aboral view, smooth, x4
PLATE 69
Spatangus californicus H. L. Clark, aboral view, x%, p. 345
Spatangus californicus H. L. Clark, oral view, x1
PLATE 70
Gonimaretia laevis H. L. Clark, aboral view, x2, p. 347
Gonimaretia laevis H. L. Clark, oral view, x2
308 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Clypeaster speciosus Verrill
Plate 48, Fig. 29
Clypeaster speciosus Verrill, 1870a, p. 95.
H. L. Clark, 1914, p. 31, pl. 135, figs. 1, 2; pl.
136, fig. 5.
This is the common Clypeaster of the Gulf of California and the
Velero collection contains 143 examples of it, ranging in size from young
ones (6-20 mm long) whose specific identity may perhaps be debatable,
to full grown adults 120-130 mm long. The width is usually about 90 per
cent of the length, but shows considerable diversity ranging from 87 to
95 per cent. The color seems to be very constantly dark purple approach-
ing black; but very young specimens are reddish violet, and orally lighter
than on the upper side. The resemblance to the preceding species has been
discussed above so that little need be said here, but it may be again emphas-
sized that the line between the two species is often invisible.
Distribution —This Clypeaster is pre-eminently characteristic of the
Gulf of California and the Velero has not found it anywhere outside the
Gulf save at Socorro Island, where a fine series of specimens was secured,
and at Clarion Island, where two adults and a half a dozen young
Clypeasters were taken which may well be referred to speciosus for the
present. As far north in the Gulf as collecting was done this Clypeaster
occurs, but south of the Gulf, it is apparently replaced by the preceding
species.
Type.—Peabody Museum, Yale University. “None designated as
type.” (Stanley C. Ball)
Type locality—La Paz, Gulf of California.
Depth.—Shore to 50 fms.
Specimens examined.—143 specimens from 20 stations.
Clypeaster elongatus®, new species
Plate 48, Fig. 30; Plate 49, Fig. 31; Plate 50, Fig. 33
Length 111 mm, breadth 89 mm, height 27 mm; the breadth is thus
only .80 of the length, the height about .25. Mouth sunken 15 mm below
the sides of the test. Test rather evenly elevated from margin to apex,
perceptibly thinner posteriorly. Orally the test is flat near the margins but
soon begins to slope to the mouth. Petaloid area about 70 mm long by 60
mm wide. Tuberculation of test quite close and fine with 200 or more
tubercles to the square centimeter and 25 or so miliary tubercles to the
square millimeter ; the tuberculation is slightly more sparse orally than on
® elongatus=elongated in reference to the form of the test.
no. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 309
the upper side. Ridges between pore-pairs of unpaired petal (except
adapically) with 7 or 8 tubercles rather close together; the miliary
tubercles on these ridges scarcely more numerous. Madreporite rather
small, about 3 mm across; ocular pores minute, but evident; genital pores
distinct but not very large. Unpaired petal nearly 40 mm long, and about
17 mm wide, the interporiferous area some 9 mm, where widest, near
middle; there are about 60 pore-pairs on each side and they nearly meet
at tip so the petal is virtually closed. Anterior petals rather more than 30
mm long, and 17 mm wide, elliptical rather than oval, and only very
slightly open at tip. Posterior petals, similar but longer (36 mm) and
wider (19 mm), scarcely more open at tip. Periproct about 5 mm across,
scarcely longer than wide, some 3 mm from the test margin, covered with
numerous miliary-bearing plates.
Primary spines smooth and blunt, dorsally about 2 mm long but some-
what longer orally and near the mouth they may be 4 or 5 mm long. They
are blunt and seem fragile; they may be slightly flattened or thickened at
tip but are not noticeably widened. Miliary spines as long as the primaries
and excessively slender; with the larger spines they form a close dense coat
on the dorsal side; orally they are less numerous and hence the covering
of the oral side is notably less dense than dorsally. Pedicellariae numerous
and large; there is great diversity in the size but nearly all are tridentate
and none are characteristic.
Color of the dry specimen is vinaceous to Hay’s brown, aborally;
orally the vinaceous tint is more or less replaced by clay color. The oral
spines are clay color or lighter with the middle, and often most of the
distal half definitely dark vinaceous brown on the upper or outer side.
The unique holotype of this Clypeaster was taken December 13, 1934,
at Station 347-35 off North Seymour Island, in the Galapagos Islands, on
a bottom of sand, rock and shell, in 3 fms. It was first referred to rotundus,
later to ochrus or speciosus, but the long, narrow and high test, the un-
usual coloring and the shape of the petals, preclude reference to any of
these known species. Among the scores of Clypeasters in the Hancock
collection this individual stands out unique. It seems best to give it a name,
even if additional specimens are never taken.
Type.—Holotype, AHF no. 47.
T ype locality.—Station 347-35, off North Seymour Island, Galapagos
Islands, 3 fms, December 13, 1934.
Distribution—Type locality.
Depth.—3 fms.
Specimens examined—The type specimen only.
310 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Family Scutellidae
Dendraster excentricus Eschscholtz
Plate 49, Fig. 32
Scutella excentrica Eschscholtz, 1831, p. 19.
Dendraster excentricus L. Agassiz and Desor, 1847, p. 135.
Echinarachnius excentricus A. Agassiz, 1873, pl. 13a, figs. 1-4.
There are literally thousands of specimens of this common and char-
acteristic west coast sand-dollar in the Velero collection but as rather
more than eleven thousand are young individuals less than 10 mm in
diameter they offer no help in the almost hopeless attempt to distinguish
reliable specific and varietal lines in the mass of material at hand. Leaving
obviously very youthful specimens out of account, there are approximately
750 specimens to which it should be possible to attach varietal if not
specific names, for the alternative of calling them all excentricus is simply
an evasion of the problem. After long and critical study of all the usable
specimens which are past 30 mm in diameter, and of scores of smaller
specimens as well, it seems necessary to recognize 3 species and one named
variety in addition to the long known excentricus. It should be added at
once that this does not leave excentricus a constantly well-defined species.
Far from it! In the material here identified as excentricus sens. str. there
are many specimens which differ obviously from a typical individual,
though they come from the same region and even from the same station.
In the absence of still more abundant material, the only practicable course
is to list them as excentricus, pointing out the features in which they are
puzzling.
Normal specimens may reach a very large size, the largest coming
from the northern stations. Specimens from the Oregon coast are fre-
quently 75 mm long, 80 mm wide and 12-14 mm high (or thick) ; the
largest in the Velero collection is 87 mm long and 95 mm wide. Ordinary
specimens from Monterey and southward are usually less than this, a
typical specimen being 65 x 70 x 9 mm but there is a great deal of diversity
in shape and stoutness. Among the Channel Islands and south to Corona
del Mar, this sand-dollar is very common in shallow water, from low
water mark down to at least 40 fms. South of the Mexican line there
seems to be a tendency to a more elongate form and a variety elongatus
was described in 1935 (H. L. Clark, p. 122). The present collection con-
tains few specimens that can be referred to this variety and 8 of these are
young bare tests (bleached), 7 from the vicinity of Cedros Island, Lower
California and 1 from off San Nicolas Island, California. A fifth speci-
men is very different in appearance from these tests for although it is also
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 311
bare it is not at all bleached but is a fine shade of brown, with the dark
brown lines of the posterior half of the test (one of the supposed characters
of elongatus) very conspicuous. It is from Laguna Beach, California, and
measures 62 x 63 x 9 mm. One other specimen, also bare and not bleached,
measures 39 x 37 x 5 mm and is of the same color and texture as specimens
of the same size of the species of Dendraster described below as laevis.
The petaloid area shows clearly that it is not /aevis and hence it would
seem to be an odd elongatus.
Several other specimens show peculiarities that distinguish them as
not normal. ‘Iwo of these deserve a few words of description. The smaller,
although not truly circular measures 43 x 43 mm. The anterior petal is
long and narrow with a relatively uncrowded tuberculation, while the
anterior paired petals are notably long and open at the tip. The color is a
brown orange, particularly bright on the oral surface. It was taken north
of Anacapa Island, California, in 15 fms. The other peculiar specimen
measures 55 mm long by 61 mm in breadth. The anterior margin is nearly
straight for about 15 mm while the posterior is straight for 20 mm or more.
The apex of the test is about 30 mm from the anterior margin but the
madreporite and the adjoining petal-bases are only 18 mm from the pos-
terior margin. Associated with this, the lateral petals are unusually diverg-
ing, the tips of the posterior pair being 22 mm apart. The anterior pair
are also markedly diverging and very long, their tips 35 mm apart. The
color of the test where bare is dull purple. The dense coat of spines is
more or less orange or yellow but the color of the test is dominant on the
upper surface; hence the general effect of the upper side is dark, dull
brown orange; the lower side is predominantly brown orange.
Coloration in excentricus is generally dull but shows a considerable
diversity. The specimens from Oregon are essentially gray, the lighter
with a tendency to cream color or dull white, especially orally, the darker
ones to a dusky brown or even black. Most California specimens tend
towards either a dull violet black or a bright maroon brown; the latter in
some cases might be called dark red. Owing to the dense coat of spines
on the lower surface, the oral side is usually lighter or more brightly
colored than the upper. As Dendrasters often congregate in great numbers
on suitable bottoms, they tend to crowd each other into a more or less
vertically inclined position and thus lie tier upon tier on a rough bottom
over which the tidal currents flow back and forth. As a result of this the
anterior fourth (more or less) of the animal becomes bleached to some
degree while the not buried part remains dark. This peculiar bicoloration
may be quite striking.
312 ALLAN HANCOCK PACIFIC EXPEDITIONS VoL. 8
Distribution —The Velero has taken excentricus at 42 stations, of
which the northernmost was on the Oregon coast, 43° 41’ N Lat. and the
most southern near the tip of Lower California. No specimens have been
taken at the Galapagos Islands or near any of the off shore islands. The
great bulk of the material is from shallow water, out to about 20 fms, for
the most part, but a few specimens have been taken in 40-50 fms and one
specimen is labeled as from 113-127 fms, southeast of Marcial Point on
the east coast of Lower California. There is perhaps some mistake about
this label as no other specimens are known from the Gulf. Other authors
extend the range to Alaska.
T ype.—Unknown.
Type locality—‘“An der Kiiste der Insel Unalaschka, am Kamt-
schatischen Meere.” (Grant and Hertlein, 1938, p. 84)
Depth.—Shore to 50 fms.
Specimens examined.—Approximately 12,316 specimens from 42
stations.
Dendraster laevis’, new species
Plate 50, Figs. 34-36
Length 52 mm, breadth 57 mm, height 6 mm; the breadth is thus
markedly greater than the length; in young specimens, however, the
length may be greater, a specimen 30 mm long is only 28 mm wide; the
largest specimen is 68 mm long, 75 mm wide and 8 mm high. Lower
surface perfectly flat, the mouth not depressed at all. Both mouth and
periproct are remarkably small, the former 2 mm, the latter scarcely 1 mm
across. The petaloid area is remarkably small, only about 26 mm long by
29 mm across; the apex is 30 mm from the anterior margin. Anterior
petal is longest but the anterior paired petals are nearly as long, 15 to 15.5
mm; posterior petals about 11 mm. All the petals are about 6 mm wide.
The pore-pairs are so small and the tuberculation of the whole test is so
fine, there are no very evident distinctive features, but the pore-pairs make
a single narrow band along each side of each petal; the intermediate area
is quite uniformly covered with minute tubercles. The four genital pores
are quite evident but the ocular pores are indistinct. Both above and below
the outlines of the plates composing the test are clearly shown and the
pattern they make is striking. This feature is not at all evident in either
the larger specimens or in the bare tests but it is a very handsome feature
of the holotype. Primary spines short and slender, knobbed at the tip,
7 laevis=smooth, in reference to the remarkably close coat of delicate spines.
no. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 313
lying flat and close so that the surface of the test is relatively smooth, quite
unlike the condition in the other species of the genus. Pedicellariae very
small and not abundant or peculiar. At each mouth angle is a group of
8-10 slender, acute primary spines, lying almost horizontally over the
mouth. Color old gold orally, more nearly deep colonial buff aborally. In
the largest specimens the color is very dull, especially on the upper side;
orally the yellow-brown shade is very evident.
This very distinct species was taken by the Velero at 9 stations between
San Miguel Island, California, and the Coronados Islands, Mexico, in
4-30 fms. While most of the 118 specimens are young or very young, large
adults were taken at 5 stations. The young are commonly quite yellow but
the big adults are rather dull colored. A bare test when carefully cleaned
revealed well the extraordinarily small petaloid area. The very small size
of mouth and anus is another striking feature.
Type.—Holotype, AHF no. 48, 82 paratypes.
Type locality.—Station 976-39, north of Santa Barbara Island, Cali-
fornia, 15-20 fms, May 28, 1939.
Distribution Channel Islands, California to Coronados Islands,
Mexico.
Depth.—4-30 fms.
Specimens examined.—The type, paratypes and 35 specimens, from 9
stations.
Dendraster mexicanus®, new species
Plate 51, Figs. 37-39
Length 60 mm, breadth 60 mm, height 9 mm. Lower surface nearly
flat, the mouth a little sunken. Test rather evenly elevated from margin to
apex which is about 28 mm from posterior margin. Madreporite just pos-
terior to apex. Petaloid area 32 by 33 mm with the anterior petal 18 mm
and each of the anterior laterals about the same. Tuberculation of test
very close and fine near margin but distinctly coarser on the elevated part
of test, particularly within the petals; on oral surface the tuberculation is
coarser and more widely spaced. Posterior petals about 15 mm long and
8 mm wide; they are slightly curved and diverge markedly, the inner
margins being 15 mm apart at tips; poriferous areas very wide (about 3
mm), the ridges between the pore-pairs are very narrow but carry a
crowded series of 3-9 minute tubercles. Areas between the poriferous
zones carry fairly numerous, but not crowded tubercles of diverse sizes;
8 mexicanus, in reference to the southern habitat of the species.
314 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
PLATE 71
Fig. 82. Lovenia cordiformis A. Agassiz, aboral view, rough, x74, p. 347
Fig. 83. Lovenia cordiformis A. Agassiz, aboral view, smooth, x2
ECHINI OF WARMER
CLARK:
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 317
none are conspicuously large but many are larger than the tubercles of the
interradial areas. Anterior lateral petals similar in general tuberculation
but the great breadth of the poriferous zones is very striking; the petals
are 9 mm wide and the interporiferous area is only 3 mm. There are 8-10
(or even more) tubercles in each series across the poriferous area. Anterior
unpaired petal is similar to the other petals but is perhaps a trifle narrower.
Abactinal surface densely covered with spines of which those covering
the interporiferous part of each of the paired petals are 2 or 3 mm long.
This series of long spines forks at the distal end of the petal and continues
to the test margin. Between petals 1 and 2 (as well as between 4 and 5
and 1 and 5) there are 2 parallel series of long spines running to the disk
margin, where all spines are 3 or even 4 mm long. This belt of long spines
forms the outer margin of the disk. Orally, the primary spines are 3-5 mm
long, cinnamon buff to light purple drab in color and contrast decidedly
with the dark purple drab of the disk. Pedicellariae very numerous, chiefly
minute ophicephalous? or triphyllous, crowded among the spines. Color of
bare test dark purple drab; of the spine coat, light brownish drab.
There are 7 adult specimens of this Dendraster, of which 2 are more
or less cinnamon brown while the other 5 are of varying shades of purple
drab. The latter, including the holotype, are from San Rosario Bay, west
coast of Lower California, in 15 fms. The other 2 and 13 very young
individuals, 11-16 mm across, are from Lagoon Head Anchorage, west
coast of Lower California, in 7 fms. The rough appearance of the posterior
part of the dorsal surface, due to the long spines in radiating series (much
as in rugosus) is a notable feature of this species but is not as important
as the character and appearance of the petals on the cleaned test.
Type.—Holotype, AHF no. 49, 4 paratypes.
Type locality Station 610-37, Rosario Bay, Lower California, 15
fms, February 28, 1937.
Distribution —West coast of Lower California from Rosario Bay to
Lagoon Head Anchorage.
Depth.—7-15 fms.
Specimens examined.—The type, paratypes and 15 specimens from 2
stations.
9It is regrettable that Clark gives no information about these ophicephalous
pedicellariae otherwise found only in Echinodiscus and (young) Astriclypeus
among Scutellids. It would have been particularly interesting to learn whether
they are bivalved as the other pedicellariae, bivalved ophicephalous pedicellariae
being otherwise a very great rarity. In a couple of specimens of this species, a
young one and a fine adult specimen, sent me by Prof. McCulloch, I do not find a
single ophicephalous pedicellaria.
318 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Dendraster rugosus!°, new species
Plate 52, Figs. 40-41
Length 49 mm, breadth 47 mm, height 9 mm; the breadth is thus
very nearly equal to the length and it is probable that in fully grown indi-
viduals the equality is complete. Lower surface almost completely flat, the
oral area being very slightly depressed (when the animal is upside down).
Test rather evenly elevated from margin to apex, which is definitely
posterior in position, less than 20 mm from posterior margin. Petaloid
area 26 by 30 mm with the anterior petal the longest and narrowest.
Tuberculation of test very close and fine abactinally but with numerous
primary tubercles scattered irregularly about, mostly in the petals or near
the test margin; tuberculation of oral surface rather sparse except near
margin. Posterior petals about 11 mm long and 5.5 mm wide; they are
quite straight and diverge markedly, the inner margins being 12 mm apart
at tips; poriferous areas very narrow, about a millimeter wide, and the
ridges between pore-pairs are so narrow they carry no primary tubercles.
The areas between the poriferous zones carry longitudinal series of rela-
tively large tubercles, about 4 basally but only 2 or 1 near the tips; these
series are more or less irregular and incomplete but give a definite char-
acter to the petals. Anterior lateral petals essentially similar but larger,
14 by 6.5 mm; the poriferous zones are wider than in the posterior pair
but are still quite narrow. The anterior unpaired petal is about 16 mm
long but only 5 mm wide, for the poriferous zones are only a little curved
and are as narrow as possible. Interporiferous areas in all 3 anterior petals
show their primary tubercles rather conspicuously in 4 somewhat imperfect
but still evident longitudinal series. Whole abactinal surface closely cov-
ered with spines; the primary spines, borne by the large tubercles of the
petals and by similar tubercles scattered over the interambulacral areas,
are about 2 mm long, nearly white and rather sharp, the other spines are
scarcely half as long and are thickened at the tip and rather bluntly pointed.
Genital pores 4, fairly large; the 5 ocular pores much less distinct. Mouth
small, largely concealed by the long (4 or 5 mm) white spines crowding
around it. Anterior primary spines of oral surface, 2 or 3 mm long, curve
outwards to left and right on the anterior fifth of the test. Pedicellariae
very small and hard to detect, not distinctive. Color of bare test pale ecru
drab or pale vinaceous drab, spines whitish.
0 — .
; rugosus—rough, in reference to the appearance caused by the numerous
projecting primary spines.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 319
There are 4 specimens of this interesting Dendraster in the Velero col-
lection. The holotype is the largest, with the others 37-39 mm long; the
smallest 36 mm wide. They resemble the type in color and in every other
respect. [hey were taken in Bay San Sebastian, Vizcaino, west coast of
Lower California, in 17 fms on a sandy bottom. The delicate coloration,
the long white primary spines and the very narrow poriferous zones are
quite distinctive.
Type.—Holotype, AHF no. 50, 3 paratypes.
Type locality—Bay San Sebastian Vizcaino, Lower California, 17
fms, August 28, 1932.
Distribution—Bay San Sebastian Vizcaino, Lower California.
Depth.—17 fms.
Specimens examined.—The type and paratypes from 1 station.
Genus ENCOPE L. Agassiz
In establishing the genus Encopfe L. Agassiz (1840, Cat. syst. ectyp.
Echinod. Mus. Neocom. p. 6; 17) mentions only the species grandis. This
species accordingly is the genotype. To take a species not originally in-
cluded in the genus as the genotype is against the rules and against common
sense. In a letter to me of May 8th, 1947 (the last but one he wrote to
me) Clark writes: ‘A young palaeontologist here (viz. Dr. Durham)
who is working on a Monograph of fossil Encofes insists that grandis is the
type. As you of course know, I consider that emarginata is the type. If you
agree with him that grandis is the type, I'll have to make some changes
in my MS; but if you agree with me that emarginata is the type then I
shall be comfortably settled. At any rate grandis and emarginata are not
congeneric, so one of them must have a new name.”
It is clear that Clark thus leaves to me the decision of the question
which species is the genotype of Encope—and there cannot be the slightest
doubt but that grandis is the genotype. Further, I thoroughly disagree
with Clark in seeing two different generic types in grandis and emarginata.
The only noteworthy difference is that the edge of the test is thicker in
grandis than in the other species; but whether the edge is some 5 mm or
only 2-3 mm thick is certainly not a difference of generic value, and any
difference of morphological value does not exist between grandis and the
other species. Particularly it must be emphasized that the internal struc-
ture is exactly alike in both.
As said in the preface I think Clark is here making too many species;
but without having access to the collections on which he is basing his
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
a ENCOPE GRANDIS
B75 Cae Ee isa
8 E PERSPECTIVA
ela ee SER ORS =
c E. weTMoREI
320
LAEVIS
ae
E = MICROPORA
Ee in| lee ieS
la es cocosi
IRREGULARIS
a ee
ECUADORENSIS
—————
FRAGILIS
oe
INSULARIS
Text figure 1. Outline drawings at median longtitudinal axis of the
species of Encope illustrated on Plates 53-59.
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC
« E. cranois
a a A Ss
8’ E. perspectiva
ee E WETMORE!
eee
bp’ E. Laevis
e’ E. microrpora
F’ E. cocosi
GC = GALAPAGENSIS
H’ E. IRREGULARIS
’ E. Ecuaporensis
ee eR Dit
y’ E. Fraciuis
k’ E. insucaris
Text figure 2. Outlines of transverse sections of same specimens at
areas of greatest thickness or height.
321
322 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
——
various species I cannot go into a critical valuation of them, and I think it
therefore the only fair thing to publish this part of his manuscript as he
left it. Future investigations will have to decide about the value of all
these species.
One more species, Encope Stokesii L. Agassiz will have to be added to
the West American species of Encope. Clark gradually became convinced
that this species is only the young of micropora, but this is decidedly a mis-
take. The internal structure of the test is markedly different from that
typical of Encope, the buccal cavity being not closed as it is in Encope but
remaining in open connection with intestinal cavity. The subgenus Melli-
tella established by Duncan (1889) for this species is therefore fully
justified—perhaps it should rather form a genus of its own.
It is very improbable that this species, so fairly common in the warmer
region of West American seas, should not be represented in the Velero
collections. Specimens of this species have probably been identified by
Clark as Encope micropora. A re-examination of his material will be
needed for settling this matter.
Encope (Mellitella) Stokesii will be dealt with fully in the forthcom-
ing Vol. [V.2 of my Monograph of the Echinoidea. Th. Mortensen
The very large number (1,212) of Encopes taken by the Velero in her
voyages to the tropical Pacific has necessitated a very detailed and time
consuming comparative study of this well-defined and easily recognized
genus. It was soon evident that more species of Encope are living on the
western coast of tropical America than had been hitherto supposed and
that the specific lines drawn needed realignment. In July, 1946, Austin
Hobart Clark of the United States National Museum published a ‘“‘Re-
vision of the Pacific Species of the Genus Encope” with the description of a
well-marked new species and the recognition of two new subspecies and a
new variety. No statement as to the number of specimens available is made
but Mr. Clark tells me that both the subspecies are based on “bare white
tests devoid of spines.” There are very few bare tests in the Velero col-
lection and the identification of such tests is a difficult matter, with the
possibility of error being very large, unless specimens with spines from the
same locality accompany them. The hundreds of Encofes available from
the Gulf of California show that the Encopes there are either the well-
known form here listed as Encope grandis or micropora. It is fruitless to
try and recognize either borealis or californica as subspecies or varieties,
as they lack any constant association with any locality or any large group
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 323
of specimens. On the other hand what Mr. Clark calls micropora galapa-
gensis is a very well-marked species, not found elsewhere and apparently
the only Encope found at the Galapagos Islands. If the type perspectiva
jonesi really came from these islands, it is strange nothing like it occurs in
the Velero collection. Its being a bare test debars it from further considera-
tion in this report.
The following key to the eleven species of Encopfe here recognized
will require modification when more material from the Central and South
American coast is available.
A. Unpaired lunule in between the posterior petals for at least most
of its length.
B. Covering of test dorsally made up of rather slender spine-
lets the tips of which are swollen but not flat-topped, nor
inverted ; cones more or less considerably in contact .
A : ‘ : : 5 : : : : . perspectiva
BB. Covering of test dorsally made up of spinelets whose tips
are more or less evidently inverted cones.
Covering of test a fairly smooth pavement of cone-bases
loosely in contact; color dark olive-gray . : . wetmorei
Cone-bases larger and in close contact, making a very
smooth pavement; petals very large, straight, wide at tip;
color olive-brown or darker with marginal fringe of short
brown or reddish-brown spinelets; lower surface brown
: : : , ; : : : : : : . laevis
AA. Unpaired lunule not forward between posterior petals but more
or less in line with the posterior lunules or anterior to them.
C. Lunules usually more or less circular or oval and
relatively small; test usually less than 120 mm in
length, commonly about 90-100 mm. Spines of test
very slender with only slightly enlarged tips .
P : ‘ : ; : ; ‘ . micropora
Test covered with the very numerous slender spines,
each tipped with an oval ball, commonly white or
light colored . ; ; : ; : ; . COCOSi
CC. Lunules much longer than wide; test often wider
than long.
ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
D.
Lunules about twice as long as wide; test thick
and heavy, the margins a couple of millimeters
or more in thickness.
E. Test large, up to 150 mm h. d.; unpaired
lunule usually longer than posterior pair,
its anterior end a little in advance of the
other two; test high, flattened between
tip of anterior petal and the posterior
lunule, highest point may be determinable
as in middle or at either end ; normal color
light brown above, with disk and lunule
margins purple . : : . galapagensis
EE. Test smaller; unpaired lunule about the
same size and about in line with posterior
lunules. Color dark gray or blackish.
Test flattened on top, commonly irregu-
lar in outline, its greatest height about
one-sixth or one-seventh of length; lun-
ules relatively small and narrow, the pos-
terior three nearly in line . . irregularis
Test more curved on top, oval or broadly
oval; its height about one-tenth of length;
lunules long and narrow, the anterior trio
slightly smaller . ; ; . ecuadorensis
DD. Lunules 3 to 4 times as long as wide; test thin
and light with relatively thin edges.
Unpaired lunule wider than other lunules;
its anterior tip little or not at all in front of the
posterior pair . . : 3 ; . fragilis
Unpaired lunule long and narrow, its anterior
end well in advance of the posterior pair . ‘
; : : . insularis
In this key Clark, according to his views did not include the species
grandis. It is easily distinguished from the rest of the species by its margin
being thicker, some 5 mm or even more. Th. Mortensen
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 325
Encope grandis L. Agassiz
Plate 53, Fig. 42
Encope grandis L. Agassiz, 1841, p. 57, pl. 6.
This easily recognized sea-urchin, or key-hole urchin, is very abundant
in the Gulf of California and the Velero has brought home no fewer than
1,222 specimens, ranging in size from 40 x 40 mm to the big adults exceed-
ing 100 mm in length with a breadth usually somewhat less but often
somewhat greater. The largest specimen in the collection is 116 mm long
by 111 mm wide and 16 mm thick, while another individual from the
same station (689-37) is 111 mm long by 115 mm wide and 16 mm
thick. The percentage of the total weight which the skeleton makes is
very great but has not been worked out. Although the coarse and heavy
build makes the specimens unmistakable, there is extraordinary diversity
in the posterior unpaired lunule, which ranges from nearly circular to a
long and narrow slit. The most striking case is a specimen 100 x 90 mm
in which the lunule is circular and only 3 mm in diameter. This is such an
extreme case it is probably pathological. At the other extreme is an indi-
vidual with the lunule 20 mm long but only 6 mm wide. There is appar-
ently no correlation between size or weight of the individual and the size
of the lunule. Thus a specimen 100 x 105 mm has the lunule 17 x 10 mm
but another specimen from the same lot, 92 x 92 mm has the lunule 33 x 16
mm. The color is dull purple, approaching black, or some shade of brown,
ranging from fawn color in some young individuals to very deep blackish
brown in adults. The marginal area is often darker than the center and the
lower surface is lighter than the upper, often markedly so. But there is no
distinctive feature in the coloration.
Distribution —The Velero took grandis at only 10 fathoms, all in the
Gulf of California, mostly along shore, but occasionally it was dredged in
from 1 to 10 fms. The scarcity of young individuals or even small adults is
striking but may be associated with the fact that all the material was col-
lected between January 30 and March 23. Perhaps collections made in
summer or early fall would show a larger number of young.
Type. (Germany ?)
Type locality —“. . . . provient probablement des Antilles.”
Depth.—Shore to 10 fms.
Specimens taken.—1,222 specimens from 10 stations.
326 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Encope perspectiva L. Agassiz
Plate 53, Fig. 43
Encope perspectiva L. Agassiz, 1841, pp. 51, 146, pl. 10b, figs. 1-5.
A. H. Clark, 1946, pl. 3, upper fig.
It is a curious thing that this well characterized species is not ade-
quately represented in the Hancock collection. In fact the only Encopes
which can be referred to it with any confidence are 5 young ones and 2
fragments which were taken at Station 485-35 in Tenacatita Bay, Jalisco,
Mexico, in 5 fms on a bottom of sand and shells. They range from 37 x 39
mm to 56 x 58 mm and are exceedingly thin and flat. The unpaired lunule
is well forward between the hind pair of petals and is the largest of all. The
spine-covering of the dorsal side of the test is made up of countless spine-
lets whose tips are ovate balls—a distinguishing feature of perspectiva.
The color ranges from Lincoln green, through dusky olive green to dusky
yellow green. The lower surface is brown, with more or less gray on the
ambulacra. The Velero did not take an adult example of this well-marked
species at any other place, but in March, 1939, she dredged, at Chacahua
Bay, Mexico, some distance southeast of Tenacatita Bay, 14 specimens of
very young Encopes, which have notably long, narrow unpaired lunules
and are apparently young wetmorei or perspectiva, presumably the latter.
Distribution—Known range from other authors, Ballenas Bay,
Lower California to Costa Rica.
T ype-—Unknown.
Type locality—Unknown.
Depth.—5-15 fms.
Specimens examined.—21 specimens from 2 stations.
Encope wetmorei A. H. Clark
Plate 54, Fig. 44
Encope wetmorei A. H. Clark, 1946, p. 2, pl. 1s ple?) fies ke
This interesting new species, so recently described, may be distin-
guished at once by the anterior position of the unpaired lunule, which lies
well in between the posterior petals, and by the peculiar character of the
spinules which cover the dorsal surface. These spinules have inverted
conical tips, the bases of the cones forming a more or less smooth pavement
covering the upper side of the test. This pavement is not so continuous or
smooth as in the following species but is very faintly rough or “furry.” The
spines around the unpaired lunule are relatively long (3 mm or 4 mm), flat
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 327
and truncate. Around the other lunules they are smaller and more slender.
The bare test is a distinct gray but the spine covering is a very dark shade
of olive gray.
There are 4 specimens of wetmorei in the collection, each about
100 mm long with a width nearly or quite as great. One specimen is in-
jured posteriorly so that the unpaired lunule has never been cut off at the
rear. The other 3 specimens are symmetrical and uninjured. All of these
specimens were taken in 5-10 fms on the south side of Petatlan Bay,
Guerrero, Mexico, in March 1934. Besides these adults there are 32 very
young Encopes, 6-15 mm in diameter, taken in 25 fms south of White
Friars, near Petatlan Bay, which may be referred to wetmorei as they are
apparently not perspectiva, and the position of the posterior lunule indi-
cates they must be one of the two species.
Distribution —Mazatlan, Mexico, to Pearl Islands, Panama.
Type-—U.S.N.M. No. E. 6768.
Type locality —San José, Pearl Islands, Bay of Panama.
Depth.—5-25 fms.
Specimens examined.—36 specimens from 2 stations.
Encope laevis!!, new species
Plate 54, Fig. 45; Plate 55, Fig. 46
Length 117 mm, width 115 mm, height 12 mm. Test stout, relatively
heavy and quite flat dorsally; highest point only about 25 mm from
anterior margin, thence it slopes very gradually and slightly to the un-
paired lunule, the margins of which are a little elevated ; height of test
there about 10 mm; posterior margin of test slightly convex but in other
specimens it may be quite straight. Unpaired petal about 40 mm long,
21 mm wide some 10 mm from the tip, which is nearly but not quite
closed; poriferous zones very wide (6 mm) with the narrow, crowded
ridges each bearing some 25 minute tubercles, of about the same size as
those which densely cover the median area. Anterior lunule small, about
6 mm long by 2 or 3 mm wide, 8 mm from the margin. Anterior paired
petals about 40 mm from anterior margin only about 10 mm in front of
the madreporite which is very large (10 mm) and densely granulated ; the
petals are about 35 mm long and 20 mm wide near the blunt tip; their
lunules are only about 9 or 10 mm from test margin and only a little
larger than the unpaired one. Posterior petals 42 mm long and 20 mm
11 Jaevis—smooth, in reference to the remarkably smooth upper surface.
328 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
wide near the very blunt tip, their lunules are a little longer (about 8 mm)
but scarcely wider than the anterior 3. Unpaired lunule nearly 12 x 3 mm,
and almost wholly within the area limited by the posterior petals. Its
surrounding spines are relatively long (2-4 mm) flattened, particularly
at the somewhat truncated tips, which, however, are not chisel-shaped.
Entire upper surface of test densely covered with very low spines each of
which is an inverted cone; the bases of these cones form a very smooth
secondary surface for the upper side of the animal. Only around the
lunules and at the very margin of the disk are there really spinelets and
these are definitely blunt, though the tips of many are rounded at the
corners. Lower surface as usual in the genus, the spines long and slender
(4-5 mm) but lying more or less flat against the test, pointing towards the
mouth on interambulacra 1 and 4 and towards the margin on inter-
ambulacra 2 and 3; on interambulacrum 5 they all tend to point towards
the lunule, as they do on the five ambulacra.
No two of the 33 specimens are exactly alike in color, but in general
they may be called olive brown lighter towards the center. Many speci-
mens are darker than the holotype, appearing to be clove brown or even
bone brown. None, however, approach the olive gray (almost deep green)
of wetmoret. The marginal spines and the lower surface are definitely
brown lacking the olive tint. The cleaned test where bare and dry (but not
at all bleached) is a pale olive gray rather markedly in contrast with the
normal olive brown of the spine-coat. While this species resembles both
perspectiva and wetmorei in the position of the posterior lunule, it cannot
well be referred to either one. It seems best to regard it as a related but
distinct species. A specimen, for deposit in the National Museum, was sent
to Mr. Austin Clark. He, however, fails to see any essential difference
between it and wetmorei. Further careful study of the material at hand
compels me to maintain daevis as a well-marked species.
Distribution—All of the specimens of this species are from Station
962-39, 11 miles northwest of Corinto, Nicaragua, in 1-3 fms on a bottom
of sand and dead leaves.
Type-—AHF no. 51, 32 paratypes.
Type locality.—Station 962-39, 11 miles NW of Corinto, Nicaragua,
1-3 fms May 4, 1939.
Distribution.—Nicaragua.
Depth.—1-3 fms.
Specimens examined.—The type and paratypes, from 1 station.
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 329
Encope micropora L. Agassiz
Plate 55, Fig. 47
Encope micropora L. Agassiz, 1841, pp. 50, 146, pl. 10a, figs. 4-8.
Encope californica Verrill, 1871, p. 586.
Encope micropora var. borealis A. H. Clark, 1946, p. 6, pl. 4.
The very large series of Encopes from the Gulf of California is con-
vincing evidence that there is but one species of Encope in that Gulf [be-
sides E. grandis.| ‘The Velero collection included 712 specimens from 21
stations. They range in size from 6 to well over 100 mm in diameter. The
finest specimens are from south of Mangles Anchorage, on the west side
of the Gulf, in Lat. 26° 16’ 55” N, in 3-5 fms and from Puerto Refugio,
Angel de la Guardia Island, Lat. 29° 42’ 37” N, along a rocky shore. The
largest measures 127 mm long by 130 mm wide and 13 mm high. The only
microporas in the collection from outside the Gulf of California are a
handsome specimen, 105 x 104 mm and two very young ones (16 and 17
mm across), collected on the west coast of Lower California, at Lagoon
Head Anchorage (28° 12’ 25” N) Station 612-37, in 7 fms and a small
bare test from San Juanica Bay, (Station 615-37). The attempt to dis-
tinguish varieties borealis and californica fails completely in the light of
this abundant material, which is notably homogeneous in general appear-
ance even though the color ranges from a deep brown, almost a purple
brown, through chocolate brown, red brown or deep yellow brown to a
sort of fawn brown. The position of the apex of the test ranges from well
anterior to distinctly posterior but the character is not associated with any
other distinctive feature and seems to have no taxonomic value.
Distribution—According to other authors, micropora ranges from
Lower California to Peru and the Galapagos Islands. (The Galapagos
records are probably based on what is here treated as FE. galapagensis.)
Type.—Paris Museum.
Type locality —M. Stokes “mais il n’en est aucun dont l’origine me
soit connue.”
Depth.—Shore to 16 fms.
Specimens examined.—712 specimens from 21 stations.
330 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Encope cocosi?2, new species
Plate 56, Figs. 48-49
Test large, stout, with small lunules. Length 120 mm and width the
same; height, greatest about 35 mm from anterior margin, 12 mm; test
slopes very gradually to the interambulacral lunule and then rather sharply
to margin. Unpaired petal about 40 mm long by 16 mm wide, open by 3 or
4 mm, only 5 mm back of the lunule; poriferous areas about 4 mm wide,
the intermediate area 8 mm. Tuberculation of test very fine and close, the
petal-ridges with a single closely placed line of some 20 minute tubercles.
Anterior lunule small, oval, about 6x 4 mm, only 5 or 6 mm from test
margin. Anterior paired lunules rather far back, a line joining them would
be 50 mm from anterior margin of test; they are very small (6-8 mm
long, 4-5 mm wide) and only 6-8 mm from test margin. Adjoining petals
short and stout, about 33x 18 mm, ending about 10 mm from lunule.
Posterior petals markedly longer (40 mm) than anterior pair, about 18-
20 mm wide. Interambulacral lunule small (9x 4 mm), a little smaller
than the posterior pair (10 x5 mm) which virtually accompany it, as the
3 lie in a straight line parallel to the nearly straight posterior margin of
test. Lower surface relatively uneven as the ambulacral furrows are evi-
dent and the vicinity of the lunules depressed. Spines of abactinal surface
very numerous, short and slender, each one terminating in a conspicuously
swollen light brown tip, resulting in a relatively smooth surface; only
around the margin and the lunules are the spines elongated and somewhat
pointed. Color clove brown becoming warm sepia on the margin; the
lower surface is unusually evenly dark with little difference between the
ambulacral and interambulacral areas.
Besides the holotype there is only one other Excope in the collection
similar to this one in form and structural details and it is noticeably dif-
ferent in color as the swollen tips of the spines are very light cream color,
resulting in a very definite drab upper surface; the lower is a deep bone
brown. Both the brown and the drab specimens were taken in Wafer Bay,
Cocos Island, Costa Rica, in 2-4 fms. Although apparently a derivative
from the micropora stock, it is easily distinguished by the primary spines.
Type.—Holotype, AHF no. 52, 1 paratype.
Type locality —Station 108-33, Wafer Bay, Cocos Island, Costa Rica,
2-4 fms.
Distribution.—Cocos Island.
Depth.—2-4 fms.
Specimens examined.—The type and paratype, from 1 station.
*2 cocost, of Cocos Island, the only place where it has been taken.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 331
Encope galapagensis A. H. Clark
Plate 57, Figs. 50-51
Encope micropora galapagensis A. H. Clark, 1946, p. 7.
One of the most interesting results of the study of the large number of
Encopes in the Velero collection was the discovery that the Galapagos
form is easily recognizable as a distinct species. The character upon which
the subspecies was based is not perfectly constant, for while the test is as a
rule equally high both in front of and behind the middle, there is individual
diversity which results in many specimens being slightly highest anterior
to the middle. Very rarely a specimen is highest posteriorly. The most
obvious character is the color, commonly an antique or brussels brown
above with the margins of the lunules as well as of the test itself dusky
purple. The lower side is a dark purple drab, sometimes quite purple,
sometimes quite drab. The contrast between the upper and lower surfaces
is usually marked. About 10 per cent of the specimens tend to be either
grayish with a purple cast or even a rather evident purple. Such specimens
show little evidence of the antique brown of normal specimens. The
lunules are commonly elongated and relatively narrow, from 2 to 4 times
as long as wide, but the anterior lunules are frequently small and more
nearly circular. The unpaired lunule is longest and its anterior end is
commonly between the tips of the posterior petals, but in some individuals
a line between those tips would be entirely anterior to the unpaired lunule.
Commonly the test is nearly circular but it is usually a little wider than
long when full grown. The margin between the posterior petals is often
nearly or quite straight but is commonly more or less convex.
The smallest specimen at hand is 10 mm in diameter with only the
unpaired lunule evident. At 12-15 mm the posterior paired lunules appear
as notches in ambulacra 1 and 5, but the notches indicative of the 3
anterior lunules are not evident until the young Encofe is about 25 mm
across. These young are very pale gray brown but at 3 mm the color is
more brown and margin of the test is purple as in adults but not markedly
so. The largest individuals exceed 130 mm in diameter, the length and
breadth about equal or the width greater. At Tagus Cove, Albemarle
Island, fine specimens were secured, 152 x 151 mm, and at Academy Bay,
Indefatigable Island, specimens 150 x 155 mm were collected.
Distribution —This fine Encope was taken only at the Galapagos
Islands, from North and South Seymour Islands and Indefatigable,
Albemarle, Charles, Hood, and Chatham Islands.
Type.—U.S.N.M. No. E. 6817.
Type locality —Chatham Island, Galapagos.
Depth.—2-73 fms.
Specimens examined.—120 specimens from 22 stations.
332 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Encope irregularis‘*, new species
Plate 58, Figs. 52-53
Test stout and more or less irregular in shape. Of all the specimens at
hand, only one, a relatively young one, is symmetrical. ‘The one selected
for holotype is here used as basis of description but no other specimen is
like it in all details. Length 112 mm, width 116 mm, height 13 mm. Apex
of test only 25 mm back of anterior margin, from which point the test
slopes gradually to posterior lunule and thence rapidly to margin. Un-
paired petal and anterior pair of about equal length, 35 mm and width
18-20 mm; poriferous zones, 4 mm across with some 20 very small
tubercles in each series; interporiferous area 10 mm wide with very fine
tuberculation; tips of petal open, only 4 mm from anterior lunule, which
is only about 5 mm from margin and is 9 x 4 mm in size. Anterior paired
lunules about 40 mm from anterior end of test; each is about 8 mm long
by 3 mm wide; the right hand one is closed by about 7 mm of test but the
left hand one has never closed, though the lower margin has bent forward
as though to do its part. Posterior petals somewhat longer than anterior
but essentially similar. Interambulacral lunule small, only 8 mm long by
2.5 mm wide; its posterior end is 15 mm from rear end of test and its
anterior end is well within (3 or 4 mm) the line joining the tips of the
posterior petals. Posterior lunules short and narrow, 10x 2 mm, only
about 7 mm from test margin. Lower surface flat or barely concave, the
pattern formed by the ambulacra unusually distinct and striking. Spines
of abactinal surface relatively short and crowded, the tips swollen and flat
for the most part but around the lunules long, flat and pointed or truncate.
Color of abactinal surface very dark, approximately fuscous or fuscous
black of Ridgway ; the marginal fringe of short spines, the tips of the spines
around the lunules and the whole lower surface are brown of some shade,
usually dark and often with a purple cast. There is considerable diversity
in the shades of color, so that no two individuals are exactly alike. In one
large specimen there is a distinct hint of deep green, while in several dark
violet is indicated. In general one may say the color is dark and dull.
Besides the holotype there are 21 specimens of this Encope, but 4 of
these are so small (less than 25 mm) their identity is based wholly on the
fact that they are from the coast of Costa Rica. The remaining 17 are
remarkable for their asymmetry. The smallest, 85 mm in length, shows no
13; eS on . . .
irregularis=lacking regularity in form or character, in reference to the fact
that nearly all the specimens are imperfectly formed at least in some detail.
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 333
striking defect but all of the rest are surprisingly different from each
other. The least asymmetrical has a noticeable concave margin adjoining
the right posterior lunule; the holotype has the left anterior lunule still
unclosed ; in another the lunules, especially the anterior one, are small; in
several two or more lunules are unclosed ; and finally a specimen 135 x 130
mm, is the most distorted of all, having no two interradial margins alike,
only the anterior and the interambulacral lunules closed, and the whole
left margin irregularly distorted.
Distribution —The holotype and 13 paratypes are from Octavia Bay,
Colombia, in 2 fms. Two small adults (about 115 mm long) are from
Bahia Honda, Panama, one from shore, the other in 15-20 fms, and a
similar but smaller specimen is from the Secas Islands, in 3 fms. The very
irregular abnormal adult described above and 3 very young individuals
are from Salinas Bay, Costa Rica, in 2 fms. The other very small specimen
is from Cocos Bay, south of Port Culebra, Costa Rica, in 2 fms. Appar-
ently this is a southern species, reaching a northern limit in Costa Rica.
Type.—Holotype AHF no. 53, 13 paratypes.
Type locality.—Station 434-35, Octavia Bay, Colombia, 2 fms, Janu-
ary 28, 1935.
Distribution—Colombia to Costa Rica.
Depth.—2-20 fms.
Specinems examined.—The type, paratypes and 8 specimens from 6
stations.
Encope ecuadorensis"‘, new species
Plate 59, Figs. 54-55
Test rather stout, 139 mm long, 146 mm wide and 15 mm high;
greatest thickness just anterior to the paired petals, when it decreases
slowly until near the posterior lunules and then more rapidly to margin.
Unpaired petal 40 mm long and 15 mm wide, elongated elliptical ; porif-
erous zones narrow, less than one-fourth of petal width, the pore-pairs
rather crowded, the ridges between with a close series of 12-16 tubercles ;
closed tip of petal (which is damaged in the holotype) is about 10 mm
from the lunule, which is 12 mm long by 4 mm wide, and some 9 mm from
the margin. Anterior paired petals shorter and wider than the unpaired
one, the form being somewhat oval rather than elliptical ; they point to the
anterior lunules which are 13 x 5 mm, and 10 mm from margin. Posterior
14 ecuadorensis, in reference to the country where this is the local Encope.
334 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
petals a bit longer but not equal to the unpaired one, the lunules 18 x 5 mm
and 10 mm from margin. Interambulacral lunule relatively small, about
equal to one of the anterior pair; its posterior end about 10 mm from
margin. Lower surface of test flat and not peculiar. Spines of abactinal
surface very numerous, slender with abruptly bulbous tips, lying inclined
towards horizontal, forming a fairly close coat ; on the lower side the spines
are relatively long but lie nearly horizontal ; they may be blunt or pointed
but are not notably enlarged at tip. Color of upper surface Saccardo’s
umber of Ridgway’s Standard, but warm sepia and cinnamon brown on
ora! surface.
In addition to the holotype, there are but two other smaller adults and
a bare test. This test is 130 mm long, 125 mm wide and 13 mm high; aside
from its elongated form it is peculiar in having the anterior petal more
elongated than in the type and relatively narrower. One of the other speci-
mens, 120 mm long by 122 mm wide, is notable for its very dark color,
Chaetura drab above, and cinnamon drab orally on the interambulacra,
more drab on the ambulacral areas. The test itself is light olive gray. The
other small adult is about 107 mm each way and the color is intermediate
between the other two adults, a dull drab above, more cinnamon below.
Although resembling fragilis in some ways, the character of the more solid
test and the narrow poriferous zones of the anterior petal serve to distin-
guish this southern form.
Distribution —This Encope is the tropical continental species, quite
different from the Galapagos species and readily distinguished from the
Panamic or Mexican forms. In addition to the 4 adults, there are 92 young
Encopes, 5-30 mm in diameter which are too young for positive identifica-
tion but are here considered as ecuadorensis on geographical grounds. The
material was all taken at 5 stations on the Ecuadorean coast between
Santa Elena Bay, Lat. 2° 10’ 36” S, and San Francisco Bay, Lat. 0° 38’
40” N, in 2-12 fms.
T'ype-—Holotype, AHF no. 54, 2 paratypes.
Type locality—Station 205-34, Santa Elena Bay, off La Libertad,
Ecuador, 8-10 fms, February 8, 1934.
Distribution —Ecuador.
Depth.—2-12 fms.
8 pecimens examined.—The type, paratypes and 93 specimens, from 6
stations.
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 335
Encope fragilis15, new species
Plate 60, Figs. 56-57
Test rather fragile, 124 mm long, 130 mm wide and 10 mm high;
greatest thickness just a little back of tip of unpaired petal which is 30 mm
long and 14 mm wide; poriferous zones wide, the two equal in width to
the median area; pore-pairs crowded, the narrow ridges between them
covered by a single series of 20-24 minute tubercles ; closed tip of petal 5 or
6 mm from anterior lunule which is 8 mm long and about 1.5 mm wide;
its tip 8 mm from disk margin. Anterior paired petals about 26 mm long
and 13 mm wide; they point directly to anterior lunules which are about
10 mm from their tip and measure 12 mm long by 2 (or less) wide. Pos-
terior petals almost duplicates in size and form of the unpaired one.
Posterior lunules about 15 mm long, 2.5-4 mm wide. Interambulacral
lunule very large, 20 mm long by 8 mm wide, only about 8 mm from
margin. Lower surface of test perfectly flat but not peculiar. Spines of
abactinal surface very numerous, slender with abruptly bulbous tips lying
almost horizontal, forming a delicate smooth coat; spines around lunules
long and pointed, those of lower surface slender, pointed or blunt or evena
bit widened at tip. Light brownish olive above, brightest on the petaloid
area and on the lunule margins, darkest at disk margin; faintly outlining
many plates are dark, Andover green lines, forming an indefinite but more
or less evident pattern ; lower surface a tawny olive.
In addition to the symmetrical and handsome holotype there are 5
other Encopfes clearly belonging to this striking species. The smallest
measures 80 x 80 mm with a thickness of 8 mm; the anterior and left
posterior lunules are not completely closed in; a less symmetrical specimen
is 90 x 88 mm and has both the right hand lunules open; a very symmetri-
cal specimen, 90 x 92 mm has all the lunules fully closed; a very unsym-
metrical and irregular adult is approximately 135 mm long and 125 mm
wide; no lunule is closed in save the anterior which is imperfectly shut in.
The largest individual is 154 mm long, 152 mm wide and 11 mm thick
(35-40 mm back of the anterior margin) ; the dull greenish lines are
present but much less evident than in the holotype. Of the smaller speci-
mens only one gives any indication of them.
Distribution—Most of the material referable to this very striking
species was taken in or near Petatlan Bay, Mexico, at 5 stations. The
holotype is from Station 764-38, near White Friars Rocks, Mexico, in
15-20 fms; the largest specimen and the very irregular adult mentioned
above are from Station 963-39, nearby but in slightly deeper water (20-25
15 fragilis=easily broken, in reference to its relatively delicate structure.
336 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
fms), and the 3 young individuals also mentioned are from Station 265-
34 in Petatlan Bay, close by in 5-10 fms. In addition to these clearly
identifiable specimens there are 146 small Encopes, 4-15 mm across, which
are presumably young individuals of this fine species. Fifty-eight of these
are from Station 764-38 (Type station). The others are from Station
264-34, south of White Friars, in 25 fms; Station 265-34, Petatlan Bay,
5-10 fms; Station 267-34, west of Morro de Petatlan, 25 fms and Station
965-39, Tenacatita Bay, 8-15 fms.
Type.—Holotype, AHF no. 55, 5 paratypes.
Type locality—Station 764-38, North of White Friars, Mexico,
15-20 fms, January 8, 1939.
Distribution.—Petatlan Bay to Tenacatita Bay, Mexico.
Depth.—5-25 fms.
Specimens examined—The type, paratypes and 146 juvenile speci-
mens from 6 stations.
Encope insularis"°, new species
Plate 61, Figs. 58-59
Test large and flat, moderately thin at the margins, conspicuously flat
and thin in young specimens (under 100 mm in diameter). Length 135
mm, width 133 mm, height 14 mm; greatest thickness on a line connect-
ing the anterior lunules, whence test slopes evenly to the anterior margin
and somewhat more gradually to the posterior margin. Unpaired petal
about 30 mm long and 12 mm wide becoming somewhat pointed at tip
but not closed; the poriferous zones are narrow (3 mm) with the ridges
bearing about 10 tubercles; the interporiferous area is 6 mm wide, its tip
about 11 mm from the anterior lunule which is 9 mm long and nearly 4 mm
wide, located 8 or 9 mm from test margin. Anterior paired petals rather
far back, a little shorter and wider than the unpaired one. Lunules of
these petals very similar in size and form to that of the anterior area.
Posterior petals longer than the others, about 35 mm in length and 15 mm
wide, their lunules long and narrow, 20 mm by 5 mm only 8 mm from
margin. In all specimens except the holotype and the next largest, these
lunules remain open distally. Interambulacral lunule long and narrow,
about 20 by 4 mm, nearly 20 mm from margin, its anterior end 8 mm in
front of the paired lunules. Posterior margin of test nearly straight in the
type but less definitely so in other specimens. Lower surface of test flat and
not peculiar. Spines of abactinal surface very numerous, slender with more
16 j * . - :
Is! ——s of an island, in reference to its occurrence at Socorro and Clarion
Siands,
No. 5 CLARK: ECHINI OF WARMER EASTERN PACIFIC 337
or less markedly bulbous tips, lying inclined towards horizontal and form-
ing a fairly even coat ; in other specimens the bulbous tips are less uniformly
evident, and the coat is not so smooth. Color mummy brown, the lower
surface a brighter shade.
Besides the holotype, there are 4 other adults (100 to 120 mm in
greatest diameter) which undoubtedly represent this rather fragile species.
The largest closely resembles the type but is a somewhat brighter brown.
The other 3 are nearly black, the smallest in particular is very dark above
and light purplish gray on the oral side. A specimen 75 mm in diameter is a
deep brown while a test 53 mm across is more nearly black. There are 2
bleached fragments and 4 very young individuals which are labeled as
from Clarion or Socorro Islands.
Distribution —The holotype, which is the largest specimen, was taken
in Braithwaite Bay, Socorro Island, in 14-18 fms, in January, 1934, and
another adult and 8 young individuals were collected at the same time and
place. Five months later, 2 small adults, 2 very young specimens and a
water worn fragment of an adult were taken at or near the same spot.
Five years later one fine adult and 2 very young individuals were taken at
Sulphur Bay, Clarion Island, in 25-45 fms. Apparently the species is con-
fined to the vicinity of those two islands.
Type.—Holotype, AHF no. 56, 4 paratypes.
Type locality—Station 129-34, Braithwaite Bay, Socorro Island,
Mexico, 14-18 fms, January 3, 1934.
Distribution Socorro and Clarion Islands, Mexico.
Depth.—14-45 fms.
Specimens examined.—The type, paratypes and 13 specimens, from 5
stations.
Mellita longifissa Michelin
Plate 62, Fig. 60
Mellita longifissa Michelin, 1858, p. 360, pl. 8, fig. 1.
This striking and easily recognized sand-dollar seems to be either very
rare or very hard to collect for in all of the many years collecting by the
V elero, it has been met with but once, when 2 apparently dead tests were
taken at San Juanico Bay, well up on the western coast of Lower Cali-
fornia. They were evidently picked up on the beach as the label reads
simply “shore.”’ One is completely bare of spines and is a trifle water worn.
The other, which is slightly the larger, has rather more than half the upper
surface still fairly well covered with the short, curved, capitate, white
spines. As the epidermis of the test is apparently black or blackish, the
resulting effect is a “pepper-and-salt” gray. Apparently this was at least
338 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
approximately the color in life. This specimen is 70 mm long, 75 mm wide
and 9 mm high at the test apex, which is about 30 mm from anterior
margin. The unpaired lunule is 22 mm long and 1.5 mm wide. The rarity
of this well-marked species, more apparent perhaps than real, is probably
due to an unusually subterranean habit which defies ordinary trawling and
dredging.
Distribution Gulf of California to Panama.
Type.—Paris Museum?
Type locality“ Habite.-Localite inconnué.-Collection Michelin.”
Depth.—Shore to 30 fms.
Specimens examined.—2 specimens from 1 station.
Family Cassidulidae
Cassidulus pacificus (A. Agassiz)
Plate 62, Fig. 61
Pygorhynchus pacificus A. Agassiz, 1863, p. 27.
Cassidulus pacificus Grant and Hertlein, 1938, p. 108, pl. 13, fig. 6; pl. 29,
figs. 4, 5; pl. 30, fig. 6.
Mortensen, 1948, Monograph of the Echinoidea,
IVi, p. 210, pl. 2, figs. 1, 2, 11-13, 19; pl. 11, figs. 2,
11-13.17
This interesting Cassiduloid is represented by 19 specimens and frag-
ments of another. The fragments are the ventral surface of a large indi-
vidual about 50 mm long by 40 mm wide, and half a dozen pieces of the
upper part of the test. The chief interest in these pieces is the color, for
they have preserved the color of the living animal to a remarkable degree.
The ground color is cream color becoming quite yellow along the sides
and on the margins of the lower surface. This surface is quite white and
unspotted but the upper surface is covered with irregular blotches of
Andover green. Around the periproct, more particularly below it, is an
evident patch of vinaceous drab. With these fragments, a specimen
20 x 17 x 10 mm was taken which is undamaged; it is nearly white with
evidence of cream color and with irregular rather numerous small blotches
of purple. The other 18 specimens show considerable diversity; 5 speci-
mens are quite brown, the largest one conspicuously blotched at the an-
terior end, dorsally with a darker shade; several half grown individuals
are nearly white, with faint indications of small dusky blotches; some
small specimens are light gray without spots. These small individuals are
about 8 mm long.
a This work unknown to Clark, being published only half a year after his
van M.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 339
Distribution—The Velero found no Cassiduloids along the mainland
coast. All of the 20 specimens were taken at islands, 9 at Socorro, 9 at
Clarion, and one, with the adult fragments was dredged in 48-73 fms
near Barrington Island in the Galapagos Islands. The Socorro specimens
were found in 4-10 fms near Cape Rule and in deeper water (10-30 fms)
in Braithwaite Bay. The Clarion material was taken off the north side of
the island in 30-56 fms or off Sulphur Bay in 25-26 fms. Other records
show the distribution to include the Gulf of California to Panama.
Type—M.C.Z. no. 2719 (Cotype).
Type locality Acapulco, Mexico.
Depth—4-73 fms.
Specimens examined.—20 specimens from 7 stations.
Family Hemiasteridae
Agassizia scrobiculata Valenciennes
Plate 63, Figs. 62-63
A gassizia scrobiculata Valenciennes, 1846, pl. 1, figs. 2-2f.
Although this Spatangoid is reported from various stations between
Mazatlan, Mexico and Capon, Peru, the Yelero met with it very rarely. A
bare test, 16 x 13 x 10 mm was found on shore at Willards Island, Gulf of
California, far north of Mazatlan, and a large bare test (21x 19x16
mm) was picked up on shore at Cartago Bay, Albemarle Island, Galapagos
Islands. One of the prizes of the Velero’s visit to the Galapagos in 1938
is a remarkably fine test of this Spatangoid presented to Captain Hancock
by Mr. Osorio of Ritter’s Landing, Charles Island. It was picked up along
shore but is neither bleached nor damaged in any way, although the spine-
coat is entirely lacking. It measures 55 mm in length, 50 mm in breadth,
and 40 mm in height. The color is a dull light brown, the 4 genital and 5
ocular pores are easily seen and the fascicles are notably distinct. This is by
far the largest specimen yet recorded. Besides these bare tests, the Velero
dredged one living specimen, a very unusual capture. It is 16 x 14x 12 mm
with the surprisingly long white spines, 2 to 3 mm. The test is white but
the dried muscles at the bases of the spines are yellow brown, so the whole
effect is very pale brown. This interesting capture was made in the Gulf of
California, between Angel de la Guardia and Mejia Islands, in 6-11 fms.
Distribution.—Gulf of California to Peru; Galapagos Islands.
Type.—Paris Museum?
Type locality —Unknown.
Depth.—Shore to 11 fms.
Specimens examined.—4 specimens from 4 stations.
340 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Brisaster townsendi (A. Agassiz)
Plate 64, Fig. 64
Schizaster townsendi A. Agassiz, 1898, p. 82.
Schizaster (Brisaster) townsendi Mortensen, 1907, pt. 2, p. 123.
Brisaster townsendi H. L. Clark, 1917, p. 179, pl. 155, figs. 4, 6, 8.
A common Spatangoid of the Eastern Pacific between southern Alaska
and the Galapagos Islands, this species was taken by the Velero at 14 sta-
tions. The largest of the 120 specimens is badly damaged but measures 60
mm in width and must have been about 70 mm long and some 30 mm high.
The smallest is 7 x 5.5 mm. All are some shade of brown, ranging from a
very light shade, with fascicles conspicuously darker (Station 1163-40,
215-225 fms) to a very dark brown, the fascioles more or less conspicuous
(Station 1497-42, 60-74 fms).
The specimens from Station 1133-40, off Redondo Beach, California,
in 49-172 fms are remarkable for having the lateroanal fasciole developed
as a conspicuous band more or less closely surrounding the anus, not in any
sense lateral. At several stations both Brisaster and Brissopsis occur and it
seems probable that some of the puzzling specimens are hybrids. In his full
decription of this Brisaster, Agassiz (1904) refers to the shape of the
young when 10 mm in diameter. Some of the Velero specimens are smaller
than that and have a very striking pentagonal form. These young pen-
tagonal individuals have the sides of the pentagon measuring 5 or 6 mm.
The peripetalous fasciole follows the margin of the pentagon on the upper
side. Ventrally each interambulacrum is a convex bulge, the posterior
(subanal) one somewhat the largest. The mouth is but little anterior to
the center, and there is no definite sternum.
Distribution—The Velero took Brisaster at stations ranging from
Oregon (Station 1497-42, Lat. 44° 49’ 10” N) to southern California
(Station 1223-41, Lat. 33°, 27’ 10” N), at depths of 20-250 fms, but the
bulk of the specimens are from the vicinity of the Channel Islands in
depths of more than a hundred fathoms. The greatest depth was in San
Pedro Channel at 225-250 fms. Off Redondo Beach specimens were taken
at 49-172 fms and off Santa Cruz Island in 64-134 fms. The absence of
Brisaster from stations further south is no doubt due to the little dredging
the Velero did at greater depths than 150 fms, for the Albatross material
on which the species is based was collected at depths of 146-995 fms at
stations from the Gulf of California to Panama. Other records extend the
range north to southeastern Alaska.
T'ype-—M.C.Z. no. 2961 (Cotype).
Type locality —Gulf of Panama to Guaymas, Mexico.
Depth.—20-995 fms.
Specimens examined.—120 specimens from 14 stations.
NO. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 341
Moira clotho!’ ( Michelin)
Plate 64, Fig. 65
Moera clotho Michelin, 1885, p. 247.
Moira clotho A. Agassiz, 1872a, p. 147.
H. L. Clark, 1917, p. 196, pl. 156, figs. 5-7.
This odd, highly specialized Spatangoid is easily recognized by the
deeply sunken, narrow, paired petals and even more deeply sunken anterior
petal which presumably serves as a brood pouch for the young. The genus
includes several species and ranges from the southeastern coast of the
United States and the western coast of tropical America to Japan and
Australia, to Suez, and Zanzibar, but apparently does not occur in Euro-
pean seas or on the west coast of Africa. The Velero met with it at 4 widely
separated stations. There are 35 specimens altogether but more than half
are very small, 5-10 mm long, and none are very large. These small ones
are chiefly from Ecuador in 1 or 2 fms. The finest specimens are from the
upper end of the Gulf of California, near Consag Rock, on a bottom of
brown mud in 21 fms. The largest are about 26 mm long, by 22 mm wide
by 18 mm high. They are almost pure white in color with the lateroanal
fasciole yellow.
Distribution Gulf of California to Ecuador.
Type.—Paris Museum?
Type locality —Mazatlan, Mexico.
Depth.—1-21 fms.
Specimens examined.—35 specimens from 4 stations.
Family Spatangidae
Brissopsis pacifica (A. Agassiz)
Plate 65, Figs. 66-67
Toxobrissus pacificus A. Agassiz, 1898, p. 83.
Brissopsis pacifica Mortensen, 1907, p. 168.
H. L. Clark, 1917, p. 203, pl. 155, fig. 1.
This is apparently the commonest Spatangoid in the Eastern Pacific.
The Velero took 674 specimens at 65 stations. They range in size from
very young (6x5 mm) to big adults (57x 47x31 mm), but the very
great majority are less than 35 mm long. As both Brissopsis pacifica and
Brisaster townsendi were taken at 6 stations, it is not surprising that hy-
brids occur apparently with considerable frequency. While the two species
are alike in dull coloration, pacifica tends to lighter shades and the fascioles
are more conspicuous. There is some evidence that muddy or even foul
18In Grant and Hertlein, 1938, the specific name is consistently misspelled
“Clothro.> s1.E.C-
342 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
bottoms affect the coloration and appearance of the specimens but the
data are not adequate to warrant positive statements. The evidence of
hybridization with Brisaster has been presented under the discussion of
Brisaster townsendi.
Distribution.—The fact that Brissopsis was taken at 65 stations is evi-
dence of its general distribution, but an analysis of the data shows that
nearly all the material is from moderately deep water off the coast of
California, particularly in the vicinity of the Channel Islands. Speci-
mens were secured near Clarion Island, but the most notable extension
of the range is the taking of 2 small but indubitable specimens at Station
786-38, northeast of Indefatigable Island in the Galapagos Islands, in
392 fms.
Type-—M.C.Z. no. 3063 (Cotype).
Type locality—Albatross Station 3355, off Point Mala, Panama,
182 fms.
Depth.—5-39 fms.
Specimens examined.—674 specimens from 65 stations.
Plagiobrissus pacificus H. L. Clark
Plate 66, Figs. 68-70
Plagiobrissus pacificus H. L. Clark, 1940, p. 351, pl. 2, figs. 3, 4.
The occurrence of this rather striking Spatangoid in the Gulf of Cali-
fornia is of great interest as it was known hitherto only from the type
and paratype taken on Hannibal Bank, Panama, in 35 fms. In the Velero
collection there are 67 specimens of this interesting Spatangoid, of which
9 are large enough to be called adults, but the test is thin and fragile and
4 are more or less crushed. The smallest is 10 x 7.5 x 4.5 mm and is quite
undamaged. The largest is 56 x 41 x 20 mm, thus much larger than the
holotype. Several specimens have a gray tint to the test instead of brown
and there is a great deal of diversity as to the degree of brownness. While
the general tuberculation and the arrangement of fascioles agree well with
Metalia, the shape of the test is quite different from the short thickset
form of most of the known species of that genus, and the large tubercles
in interambulacra 1 and 5 show conclusively that it is not that genus.
Distribution.—All the material of Plagiobrissus was taken in the Gulf
of California at depths of 5 to 75 fms except 3 specimens, as follows: a
single fine bare test from Station 948-39, which is in Panama Bay in 30-35
fms; a very good half-grown specimen from Station 773-38, which is off
Nuez Island, Cocos Island, in 31-50 fms; and a damaged young individual,
14 mm long, whose identification is somewhat dubious, from Station 212-
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 343
34, off La Plata Island, Ecuador. Apparently this Spatangoid occurs
throughout the Panamic region, but further material is necessary to estab-
lish the limits of its distribution.
Type.—M.C.Z. no. 7625.
Type locality —Hannibal Bank, Panama, 35 fms.
Depth.—5-75 fms.
Specimens examined.—67 specimens from 19 stations.
Idiobryssus coelus H. L. Clark
Plate 67, Figs. 71-73
Idiobryssus coelus H. L. Clark, 1939, pp. 173-176, pl. 17.
The holotype of this odd little Spatangoid is the only specimen at hand.
It was taken, together with a smaller paratype, in 40-70 fms, in Darwin
Bay, Tower Island, Galapagos Islands, January 16, 1938. It is 12 mm
long by 10 mm wide and 4 or 5 mm high. The color in life was white, but
the dry specimen is more or less pale brown. The paratype is in the
Museum of Comparative Zoology, Cambridge.
Distribution —Tower Island, Galapagos Islands.
Holotype —AHF no. 2.
Type locality—Darwin Bay, Tower Island, Galapagos Islands.
Depth—40-70 fms.
Specimens examined.—The type.
Brissus latecarinatus (Leske)
Plate 67, Fig. 74; Plate 68, Fig. 75
Spatangus brissus var. latecarinatus Leske, 1778, pp. xx, 185.
Brissus carinatus Gray, 1825, p. 431.
Brissus latecarinatus H. L. Clark, 1917, p. 219.
This widespread Indo-Pacific Spatangoid has long been known from
the Panamic region but the Yelero has not secured many specimens nor
are any of those at hand even half grown. Moreover of the 21 specimens,
16 are bare tests, nearly all bleached, and the largest is only 56 x 44 x 30
mm. This is almost one-third of the size of the largest recorded
specimen, now in the collection of the California Academy of Sciences,
which Grant and Hertlein (1938, p. 130) describe as 166 mm long, 128
mm wide and 106 mm high. Of the Velero specimens having the spines on
and evidently alive when taken, the largest is 39 x 32 x 32 mm (Station
1079-40). The other Velero specimens, alive when taken, were found at
Secas Islands, Panama (Station 446-35) and at Pond Island, Gulf of
California (Station 1079-40). The bleached tests at hand, were found
along the east shore of Angel de la Guardia Island, March 6, 1936; at
344 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Willards Island, in January, 1940, and at Puerto Refugio, Angel de la
Guardia Island, also in January, 1940. All the specimens at hand were
taken along shore.
Distribution—Gulf of California to Panama; Indo-Pacific from
Hawaii, Australia to the Red Sea and Japan.
Type—Unknown.
Type locality—Unknown.
Depth.—Shore.
Specimens examined.—21 specimens from 6 stations.
Meoma grandis Gray
Plate 68, Figs. 76-77
Meoma grandis Gray, 1851, p. 132.
A. Agassiz, 1873, p. 603, pl. 34, figs. 1, 2.
This big dull-colored Spatangoid appears to be common in the tropical
Eastern Pacific, north of the equator, and the Velero collection contains
some 90 specimens from 25 stations. All, however, are adults as the
smallest is 80 x 72x 39 mm. The largest is nearly twice that but is at
present broken into two large bare pieces. It was in life at least 150 mm
long, by 137 mm wide and 75 mm high. No other specimen is nearly so
large, the biggest being 120 x 110 x 50 mm. Only 2 specimens, besides the
large fragments, are bare. The color of the naked tests is a light brown
gray or a deep brown with a violet cast. All of the other specimens are
brown, usually very dark but the fragments from Cocos Island are defi-
nitely yellow brown. Several specimens are nearly black. On the whole,
this J/eoma must be regarded as the dullest colored, most unattractive
echinoid of the west coast.
Distribution—Meoma ranges from Port Utria. Colombiafi some 6
degrees north of the equator, in 15-30 fms, to the upper end of the Gulf
of California (Angeles Channel, 28° 57’ N). At the outlying islands, the
Velero took good specimens in Braithwaite Bay, Socorro Island, and
fragments of at least two large ones in Chatham Bay, Cocos Island. At
the Galapagos Islands, the only specimen met with was the very large one,
whose bare fragments were dredged in 58-60 fms, east of the south end of
Albemarle Island. The bathymetric range of Meoza is not great, from
along shore in a fathom or so to something less than 60 fms.
Type.—British Museum (No number).
Type locality—Gray’s original locality, “Australia,” generally con-
sidered erroneous.
Depth.—Shore to 60 fms.
Specimens examined.—90 specimens from 25 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 34
wa
Spatangus californicus H. L. Clark
Plate 69, Figs. 78-79
Spatangus californicus H. L. Clark, 1917, p. 235, pl. 156, figs. 1-3; pl.
157, fig. 10.
This interestingly isolated Spatangus is represented in the Velero col-
lection by 240 specimens from 47 stations, ranging in size from
21x 18x12 mm to 86x 79 x 43 mm. The general form is ovoid, more or
less flattened both above and below, but some individuals are more ellip-
soidal though flattened on the oral side. Occasionally the breadth equals the
length and in such individuals the height may be less than half the breadth.
The color shows a considerable diversity and is difficult to describe.
Theoretically the test is a rich deep purple and the spines more or less
nearly white, but often the test is deep red brown or, on the other hand,
a clear gray without hint of red or purple. Occasionally the test is light
with dark blotches and such specimens have a very different appearance
and are much more nearly handsome than the dull purple one. The spines
may be cream color or yellow or orange brown but are usually quite in
contrast with the test.
The development of the very young into mature individuals affords
some very interesting and puzzling stages which are confusing because of
the resemblance to Palaeotropus. The striking feature of these stages is the
simple nature of the ambulacral pores which are single and not paired. In
some very young individuals only the pores at the apical end of the ambu-
lacra are clearly present and each is single. There is thus no well-marked
petal, only a double series of pores at the upper end of each ambulacrum
as in Palaeotropus. With increasing age and size the older pores elongate
horizontally and become two pores united by a short groove essentially
like the pore-pairs on each side of the petal in Spatangus and its allies. ‘This
alteration in the appearance of the petals does not take place at the same
stage of development in all young, but usually occurs very soon after the
assumption of the spatangoid form so that most individuals 8-10 mm long
have normal petals of 2 columns of double pores. Rarely, individuals as
much as 15-20 mm long, have only single pores in each side of the ambula-
crum, and the absence of petals in such specimens is very puzzling and may
cause them to be referred to Palaeotropus. (See Fig. 76.)
Distribution —The Velero did not take Spatangus until February,
1936, when specimens were secured in 100-120 fms, south of Coronados
Island in the Gulf of California. A little later specimens were dredged in
165 fms off San Francisquito Bay and in 100 fms a bit nearer the mouth
346 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
of the Gulf. It was over 2 years before further specimens were secured
and these were taken off Santa Catalina Island in 50 fms. The following
year Spatangus was found in deep water, 130-330 fms, off Santa Rosa
Island. In the fall of 1939, 5 stations between Catalina Island and San
Benito Island, Lower California, yielded numerous specimens. In the
months between February 1940 and October 1941, especially in the
summer of 1941, Spatangus was secured at many stations at depths of 35-
Text figure 3. Spatangus californicus x4.
225 fms. But nearly all the material was taken near or south of the Chan-
nel Islands and near San Benito and Cedros Islands. A few specimens
were secured near the end of Lower California but none were taken
further south nor near any of the outlying islands.
Type —U.S.N.M. No. 789.
Type locality—Albatross Station 2973, “off Southern California,
68 fms.”
Depth.—35-225 fms.
Specimens examined.—240 specimens from 47 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 347
Gonimaretia laevis H. L. Clark
Plate 70, Figs. 80-81
Gonimaretia laevis H. L. Clark, 1917, p. 244, pl. 161, figs. 5-7.
This interesting Spatangoid, originally described from 3 small speci-
mens (24-34 mm long) is well represented in the Velero collection by 57
specimens and the large adults are exceptionaly handsome for a Spatang-
oid. The largest at hand is 42 x 36 x 18 mm, about a third larger than the
holotype; the smallest, not quite 10 mm long, well shows the characteristic
features of both genus and species. The color of the largest specimens is a
light fawn brown but many of the smaller ones are quite a dark brown.
The primary spines are not conspicuous but are very slender and quite
appressed. The coat of secondary spines, in the best specimens, is very
dense and has a silky appearance and feeling. The large pedicels near the
mouth are a deep brown (almost black in dry specimens). Although lack-
ing the diversity of color and the numerous long primaries of Lovenia,
this Spatangoid rivals that notable genus in the general attractiveness ot
its appearance.
Distribution Although taken at 21 stations, this very striking “sea-
mouse” (if it may be called by the popular name long ago given to
Lovenia) was found chiefly among the Channel Islands, California, and
up in the Gulf at least to Angel de la Guardia Island. A single specimen
was taken at the San Benito Islands, Mexico. One lot is labeled 814-38, a
station near Hood Island, Galapagos Islands, but in the absence of other
specimens from south of the Gulf, this label must be regarded with sus-
picion. There is no doubt of the specimens being young Gonirnaretia for
they closely resemble others of the same size from the Gulf of California.
While occasionally taken in 3-5 fms, most specimens were dredged in
water of moderate depths, 20-165 fms.
Type.-—U.S.N.M. No. 805.
Type locality—A lbatross Station 2911, south of San Clemente
Island, California.
Depth.—3-165 fms.
Specimens examined.—57 specimens from 21 stations.
Lovenia cordiformis A. Agassiz
Plate 71, Figs. 82-83
Lovenia cordiformis A. Agassiz, 1872, p. 57.
H. L. Clark, 1917, p. 254, pl. 161, figs. 8-12.
Grant and Hertlein, 1938, p. 136, pl. 13, figs. 1, 2.
There can be little question that this relatively common Spatangoid is
one of the handsomest species of the warm Eastern Pacific. It is well repre-
sented in the Velero collection by 100 specimens from 44 stations, but
348 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
owing to the relatively delicate test most of these, unless very small, were
more or less damaged when taken. The very long primary spines are
exceedingly slender and fragile and are commonly broken off to a con-
siderable extent, and the test itself is more often broken than not. The
largest specimen in the collection is from the Galapagos Islands and
measures 75 x 51 x 20 mm. The anterior left corner and most of the large
primaries are gone. Another very large specimen is 55 mm across but it is
badly damaged at the posterior end so its living length cannot be deter-
mined. Many of the specimens are less than 10 mm in length, some less
than 5 mm, and these small ones are easily confused with other young
Spatangoids but if the condition of the specimen permits the internal
fasciole to be seen, there can be no confusion of Lovenia with other genera.
The color of these striking Spatangoids is diverse, ranging from a light
gray brown with the long primary spines pure white to a deep yellowish
brown or a gray brown with the primaries not essentially different. The
gray brown shades into lavender and purple, with the long spines con-
spicuously banded. The handsomest specimens are definitely light purple,
fading into light brown or dirty white orally. The long spines on the sides
and orally are unicolor, lavender or very pale brown, but dorsally they are
prettily banded with light brown, pale orange or dirty white, and lavender
or purple. The long spines of ambulacrum 3 are nearly white with widely
separated very narrow bands of dark purple. Bare tests are a light dingy
lavender or yellow brown, if not bleached, and the sunken primary
tubercles, large oddly shaped “petals” and conspicuous inner fasciole make
them unusually interesting curios.
Distribution —Lovenia is a striking feature of shore and shallow
water collecting at Newport and Corona del Mar. Further north it ranges
to San Pedro and the Channel Islands, where it has been taken in 17-75
fms. The northern limit of its range is apparently just above 34°. It is
fairly common in the Gulf of California, as far north as 29° 33’, in water
2-75 fms deep. Rather common on the west coast of Mexico, it extends its
range to the Secas and Jicarita Islands, Panama, in 12-30 fms. The Velero
took it twice at Braithwaite Bay, Socorro Island, twice at Cocos Island
and twice in the Galapagos Islands, at Chatham and Albemarle Islands
in 4 and 30 fms. It seems a little strange that more specimens were not
secured at the Galapagos Islands, for so conspicuous a shallow water sea-
urchin is not easily overlooked.
Type.—M.C.Z. no. 3188 (Cotype).
Type locality—San Diego, Guaymas.”
Depth.—Shore to 75 fms.
Specimens examined.—100 specimens from 44 stations.
No. 5 CLARK : ECHINI OF WARMER EASTERN PACIFIC 349
LITERATURE CITED
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Acassiz, A. AND H. L. CLark
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Acassiz, L.
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350 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
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352 ALLAN HANCOCK PACIFIC EXPEDITIONS VOL. 8
Taxonomic Index with page numbers of illustration in bold face.
Cidaroida, 229
Cidaridae, 229
Eucidaris Thouarsii (Valencien-
nes), 229, 239
Hesperocidaris panamensis (A.
Agassiz), 230, 239
perplexa (H. L. Clark), 231, 240
asteriscus H. L. Clark, 233, 240,
241
Centrocidaris Doederleini (A.
Agassiz), 234, 242
Centrechinoida, 235
Centrechinoidae, 235
Centrechinus mexicanus (A.
Agassiz), 235, 242
Astropyga pulvinata (Lamarck),
236, 251
Centrostephanus coronatus (Ver-
rill), 237, 251
Arbaciidae, 244
Arbacia incisa (Blainville; ? #
Gmelin), 244, 252
spatuligera (Valenciennes), 246,
252
Tetrapygus niger (Molina), 247,
252
Echinidae, 247
Lytechinus anamesus H. L. Clark,
247, 253
pictus (Verrill), 249, 253
semituberculatus (Valenciennes),
253, 261
Toxopneustes roseus (A. Agassiz),
254, 262
Tripneustes depressus A. Agassiz,
254, 263
Strongylocentrotidae, 265
Loxechinus albus (Molina), 255,
265
Caenocentrotus gibbosus (L.
Agassiz and Desor), 255, 265
Allocentrotus fragilis, 256, 276
Strongylocentrotus franciscanus (A.
Agassiz), 257, 278
purpuratus (Stimpson), 256, 279
Echinometridae, 281
Echinometra oblonga (Blainville)
257, 281
VanBrunti A. Agassiz, 258,
293
Exocycloida, 294
Clypeastridae, 294
Clypeaster europacificus H. L.
Clark, 258, 294
rotundus (A. Agassiz), 267,
296
ochrus H. L. Clark, 267, 297
speciosus Verrill, 268, 308
elongatus H. L. Clark, 268,
269, 270, 308
Scutellidae, 310
Dendraster excentricus Esch-
scholtz, 269, 310
laevis H. L. Clark, 270, 312
mexicanus H. L. Clark, 271,
313
rugosus H. L. Clark, 272, 318
Encope, 319
grandis L. Agassiz, 273, 320,
321, 325
perspectiva L. Agassiz, 273,
320, 321, 326
wetmorei A. H. Clark, 274,
320, 321, 326
laevis H. L. Clark, 274, 283,
320, 321, 327
micropora L. Agassiz, 283, 320,
321, 329
cocosi H. L. Clark, 284, 320,
321, 330
galapagensis A. H. Clark, 285,
320, 321, 331
irregularis H. L. Clark, 286,
320, 321, 332
ecuadorensis H. L. Clark, 287,
320, 321, 333
fragilis H. L. Clark, 288, 320,
321, 335
insularis H. L. Clark, 289, 320,
321, 336
Mellita longifissa Michelin, 290,
337
Cassidulidae, 338
Cassidulus pacificus (A.
Agassiz), 290, 338
Hemiasteridae, 339
Agassizia scrobiculata Valencien-
nes, 299, 339
Brisaster townsendi (A.
Agassiz), 300, 340
Moira clotho (Michelin), 300,
341
Spatangidae, 341
Brissopsis pacifica (A. Agassiz),
301, 341
Plagiobrissus pacificus H. L.
Clark, 302, 342
Idiobryssus coelus H. L. Clark,
303, 343
Brissus latecarinatus (Leske),
303, 304, 343
Meoma grandis Gray, 304, 344
Spatangus californicus H. L.
Clark, 305, 345, 346
Gonimaretia laevis H. L. Clark,
306, 347
Lovenia cordiformis A. Agassiz,
315, 347
INDEX
Allan Hancock Foundation Publications of The University of Southern California,
Volume 8. Titles of papers and plate illustrations are in bold face.
(Achelous) angustus, Portunus, 3
aculeatus, Phyllophorus,
64, 108, 133, 184
Agassizia scrobiculata, 226, 298, 299, 339
albomaculata, Ophionereis, 32
albus, Echinus, 265
Loxechinus, 255, 259, 265
Strongylocentrotus, 265
alexanderi, Ophiocoma, 19
Alexandria magnifica, 295
Allocentrotus fragilis, 256, 259, 276, 277
Amphichondrius, 22
granulosus, 23
laevis, 22, 24, 48
Amphicontus, 1
minutus, 2, 4
amphilogus, Ophiodesmus, 33, 35, 58
Amphiodia, 24
sculptilis, 24, 25, 50
Amphiophiura, 36, 40
irregularis, 36, 38, 39, 54
paucisquama, 38, 39, 40, 56
Amphioplus, 27, 28
philohelminthius, 27, 28, 50
Amphiuridae, 15
anamesus, Lytechinus, 247, 248, 249,
250, 253, 260
Anaperus, 77, 102, 120, 125
carolinus, 121
glasselli, 120, 122
peruana, 119, 120, 121, 130
peruanus, 120, 131
peruviana, 63, 120
anceps, Cercodemas, 98
A new genus of Brittle Stars,
Amphicontus, 1
anexigua, 91
Neopentamera, 63, 69, 90, 91, 160
angustus, Portunus (Achelous), 3
annulata, Ophionereis, 32, 34
anomalus, Leptychaster (Parastropec-
ten), 201
antarcticus, Psolus, 148
Apentamera, 77, 89, 91
lepra, 63, 68, 70, 89, 91, 92, 160
Arbacia, 246
incisa, 244, 245, 246, 250, 252
spatuligera, 246, 250, 252
stellata, 244, 245
Arbaciidae, 244
A report on the echini of the warmer
Eastern Pacific based on the collec-
tions of the Velero III, 225
Asteriidae, 210
asteriscus, Hesperocidaris, 233, 234, 238,
240, 241, 243
Astriclypeus, 317
Astropectinidae, 197
Astropyga pulvinata, 236, 250, 251
variety venusta, 236
Athyone, 77, 118, 120
glasselli, 63, 118, 119, 120
Athyonidium, 122, 123, 127
chilensis, 64, 70, 74, 127, 128, 129,
131
bairdi, Ophiacantha, 13
beebei, Pentamera, 63, 85, 86, 90, 158
(ae i 64, 65, 66, 137, 139,
190
Benthopectinidae, 201
benti, variety zaca, Thyone, 119
bidentata, Thyone, 63, 65, 66, 67, 69, 104,
105, 105, 170
Boletia rosea, 262
borealis, Encope, 322
briareus, Coronaster, 211
Thyone, 72, 74, 119, 120, 121, 122
Brisaster, 340, 342
townsendi, 298, 300, 340, 342
(Brisaster) Schizaster, townsendi, 340
Brissopsis, 340, 342
pacifica, 266, 298, 301, 341
Brissus carinatus, 343
latecarinatus, 303, 304, 307, 343
brissus, variety latecarinatus, Spatan-
gus, 343
bullatum, Psolidium, 144
Caenocentrotus, 276
gibbosus, 226, 255, 259, 265
calcigera, Pentamera, 74, 85
californica, Cucumaria, 63, 65, 66, 67,
68, 69, 70, 78, 79, 80, 82, 83, 154
Encope, 322, 329
Ophiocnida, 25, 27, 50
californicus, Luidiaster, 201, 205, 206,
216, 218
Spatangus, 305, 307, 345, 346
carinatus, Brissus, 343
carolinus, Anaperus, 121
Cassidulidae, 338
Cassidulus pacificus, 226, 290, 291, 338
Centrechinidae, 235
Centrechinoida, 235
Centrechinus, 235, 237, 244
mexicanus, 226, 235, 237, 242, 243
Centrocidaris, 234
Doederleini, 226, 234, 242, 243
[ 353 ]
354 INDEX
Centrostephanus, 237, 244
coronatus, 226, 237, 250, 251
Cercodemas, 98
anceps, 98
chierchia, Cucumaria, 86
Pentamera, 63, 64, 66, 67, 68, 69,
84, 85, 86, 89, 160
chilensis, Athyonidium, 64, 70, 74, 127,
128, 129, 131
Cucumaria, 63, 69, 78, 80, $1, 91,
156
Ophiophragmus, 19, 22
Thyone, 122, 128
Thyone (Stolus), 127
chiloensis, Cucumaria, 88
Pentamera, 63, 65, 69, 70, 85, 88, 89,
89, 160
Cidaridae, 229
Cidaris (Eucidaris) Thouarsii, 229
Thouarsii, 229
Cidarites pulvinata, 236
Cidaroida, 229
cigaro, Thyone, 120
clotho, Moera, 341
Moira, 298, 300, 341
clothro, Moira, 341
Clypeaster, 294, 296, 297, 308, 309
elongatus, 226, 266, 268, 269, 270,
308
europacificus, 226, 258, 259, 294,
295, 296
ochrus, 226, 266, 267, 297, 309
rotundus, 226, 266, 267, 296, 309
speciosus, 266, 268, 297, 308, 309
Clypeastridae, 294
cocosi, Encope, 282, 284, 320, 321, 323,
330
coelus, Idiobryssus, 227, 303, 307, 343
Colochirus, 98, 99
peruanus, 99
convergens, Psolidium, 142
quadrangularis, 98
cordiformis, Lovenia, 227, 314, 315, 347
Coronaster, 210, 212
briareus, 211
eclipes, 213
haliceps, 211, 213
marchenus, 210, 213, 220,
222
volsellatus, 211, 212, 213
coronata, Echinodiadema, 237
coronatus, Centrostephanus, 226, 237,
250, 251
crassa, Ophiolepis, 42
crax, Cucumaria, 63, 66, 78, 83, 154
crocea, Holothuria, 84
Cucumaria, 76,77, 78, 93,99, 103
californica, 63, 65, 66, 67, 68, 69,
70, 78, 79, 80, 82, 83, 154
chierchia, 86
chilensis, 63, 69, 78, 80, 81, 91, 156
chiloensis, 88
crax, 63, 66, 78, 83, 154
curata, 78
dubiosa, 63, 66, 67, 78, 81, 82, 84,
156
exigua, 80, $1, 91
fallax, 78, 79, 80
fisheri, 82
frondosa, 78, 80
godeffroyi, 63, 78, 83
japonica, 78
kollikeri, 82
leonina, 81, 82, 83
lubrica, 78, 81, 82
miniata, 78
pulcherrima, 88
pseudocurata, 78
salmini, 82, 83
tabulifera, 88
vegae, 78
Cucumariidae, 71, 73, 76
curata, Cucumaria, 78
dawsoni, Luidiaster, 205
debilis, Stenorhynchus, 3
Dendraster, 225, 227, 311, 317, 319
excentricus, 266, 269, 310, 311,
312
variety elongatus, 310, 311
laevis, 266, 270, 311, 312
mexicanus, 271, 275, 313
rugosus, 272, 275, 318
Dendrochirota, 61, 63, 71
depressus, Tripneustes, 226, 250, 254,
263, 264
destichadus, Phyllophorus, 135
Diadema, 235, 243
mexicanum, 235, 243
Diadematidae, 243
Diadematoidea, 243
dictyota, Ophionereis, 29, 30, 32, 52
diomedeae, Psolus 64, 65, 67, 68, 69,
70, 147, 149, 194
disacanthus, Ophiophragmus, 15, 16, 58
Doederleini, Centrocidaris, 226, 234,
242, 243
Goniocidaris, 234
doliolum, Pentacta, 99
Dorocidaris panamensis, 230
dorsipes, Psolidium, 64, 69, 70, 142, 143,
192
dubiosa, Cucumaria, 63, 66, 67, 78, 81,
82, 84, 156
duplicatus, Eucyclus, 122, 127
eat eae Psolidium, 64, 65, 143, 144,
Echinarachnius excentricus, 310
Echinidae, 247
INDEX
Echinocidaris incisa, 244, 245
Echinodiadema coronata, 237
Echinodiscus, 317
Echinometra, 281, 293, 294
insularis, 293
mathaei, 281, 292
mathaei oblonga, 281
oblonga, 226, 257, 259, 281, 292
Van Brunti, 226, 258, 259, 281,
292, 293
variety rupicola, 293
Echinometridae, 281
Echinostrephus, 292
Echinus (Agarites) spatuliger, 246
Echinus albus, 265
niger, 247
oblongus, 281
purpuratus, 279
stellatus, 245
Echinus (Psammechinus) semituber-
culatus, 261
Echinus (Toxopneustes) gibbosus, 265
eclipes, Coronaster, 213
ecuadorensis, Encope, 287, 291, 320
321, 324, 333, 334
ekmani, Psolidium, 64, 65, 70, 143, 145,
148, 149, 194
elongatus, Clypeaster, 226, 266, 268, 269,
270, 308, 310, 311
emarginata, Encope, xi, xii, 319
Encope, xi, 227, 319, 320, 322, 323, 329,
330, 331, 332, 333, 334, 335, 336
borealis, 322
californica, 322, 329
cocosi, 282, 284, 320, 321, 323,
330
ecuadorensis, 287, 291, 320, 321,
324, 333, 334
emarginata, xi, xii, 319
fragilis, 288, 291, 320, 321, 324,
334, 335
galapagensis, 226, 282, 285, 320,
321, 324, 329, 331
grandis, xi, 273, 275, 319, 320, 321,
322, 324, 325, 329
insularis, 289, 291, 320, 321, 324,
336
irregularis, 282, 286, 320, 321, 324,
332
laevis, 274, 275, 282, 283, 320,
321, 323, 327, 328
(Mellitella) Stokesii, 322
micropora, xii, 282, 283, 320, 321,
322, 323, 329, 330
variety borealis, 329
variety californica, 329
micropora galapagensis, 323, 331
perspectiva, 273, 275, 320, 321
322, 323, 326, 327, 328
355
perspectiva jonesi, 323
Stokesii, 322
wetmorei, 274, 275, 320, 321, 323,
326, 327, 328
ian Psolidium, 64, 65, 143, 144,
Eucidaris Thouarsii, 226, 229, 238, 239
Eucyclus, 122, 123, 127
duplicatus, 122, 127
Eupentacta pseudoquinquesemita, 90, 91
europacificus, Clypeaster, 226, 258, 259,
294, 295, 296
eurybrachiplax, Ophionereis, 32, 34
Euthyonidium, 122, 123, 124, 127
occidentalis, 124, 126
ovulum, 64, 65, 68, 70, 122, 124, 125,
126, 129, 130, 131, 182
seguroensis, 124
veleronis, 64, 67, 124, 126, 182
excentrica, Scutella, 310
excentricus, Dendraster, 266, 269, 310,
311, 312
variety elongatus, 310
Echinarachnius, 310
exigua, Cucumaria, 80, 81, 91
Exocycloida, 294
fallax, Cucumaria, 78, 79, 80
fisheri, Cucumaria, 82
floridae, Tamaria, 210
fragilis, Allocentrotus, 256, 259, 276,
217
Encope, 288, 291, 320, 321, 324,
334, 335
Strongylocentrotus, 276
franciscana, Toxocidaris, 278
franciscanus, Strongylocentrotus, 257,
259, 278, 279, 280, 281
frondosa, Cucumaria, 78, 80
fusus, Thyone, 103, 105, 120
galapagensis, Encope, 226, 282, 285, 320,
321, 324, 329, 331
gibber, Neothyone, 63, 64, 67, 70, 108
109, 110, 111, 112, 113, 174
Stolus, 109, 113
Thyone, 109, 110, 113
gibbosa, Neothyone, 63, 64, 67, 68, 70,
92, 109, 110, 111, 113, 114, 118
176, 178
gibbosus, Caenocentrotus, 226, 255, 259,
265
Echinus (Toxopneustes), 265
glasselli, Athyone, 63, 118, 119, 120, 122
Thyone, 119
godeffroyi, Cucumaria, 63, 78, 83
Gonimaretia, 347
laevis, 227, 306, 307, 347
Goniocidaris Doederleini, 234
356
grandis, Encope, xi, 273, 275, 319, 320,
321, 322, 324, 325, 329
Meoma, 227, 304, 307, 344
granulosus, Amphichondrius, 23
grisea, Leptopentacta, 93, 93, 94, 97, 162
halicepus, Coronaster, 211, 213
hancocki, Thyonepsolus, 64, 65, 68, 70,
138, 140, 188
Heliocidaris stenopora, 293
Hemiasteridae, 339
Hemieuryalidae, 9
Hesperocidaris asteriscus, 233, 234, 238,
240, 241, 243
panamensis, 226, 230, 231, 232,
238, 239
perplexa, 231, 232, 234, 238, 240
hispida, Ophiocnida, 27
Holothuria crocea, $4
peruana, 129, 131
peruviana, 120
horrida, Paulia, 3
horridus, Luidiaster, 205
Idiobryssus coelus, 227, 303, 307, 343
imbricata, Leptopentacta, 94, 96
Parocnus, 96
imbricatus, Ocnus, 93
Inachoides microrhynchus, 3
incisa, Arbacia, 244, 246, 250, 252
Echinocidaris, 244, 245
inermis, Leptychaster (Parastropecten),
200
insularis, Echinometra, 293
Encope, 289, 291, 320, 321, 324, 336
irregularis, Amphiophiura, 36, 38, 39, 54
Encope, 282, 286, 320, 321, 324, 332
japonica, Cucumaria, 78
japonicus, Leptopentacta, 93
Stichopus, 72, 74
javanica, Leptopentacta, 93, 93, 94, 98
javanicus, Ocnus, 93
kéllikeri, Cucumaria, 82
laevis, Amphichondrius, 22, 24, 48
Dendraster, 266, 270, 311, 312
Encope, 274, 275, 282, 283, 320,
321, 323, 327, 328
Gonimaretia, 227, 306, 307, 347
Sigsbeia, 9, 11, 44
latecarinatus, Brissus, 303, 304, 307, 343
lechleri, Thyone, 119
leonina, Cucumaria, 81, 82, 83
Sate 2 Pentanners, 63, 68, 70, 89, 91, 92,
Leptopentacta, 76, 92, 93
grisea, 93, 93, 94, 97, 162
imbricata, 94, 96
javanica, 93, 94, 98
nina, 63, 70, 93, 94, 97, 98, 164
nova, 63, 66, 93, 94, 95, 95, 96,
98, 162
INDEX
panamica, 63, 65, 66, 67, 93, 93,
94, 96, 162
panamensis, 98
typica, 93, 94, 96
Leptychaster, 197
(Parastropecten) stellatus, 197,
200, 201, 216
anomalus, 201
inermis, 200
leurum, Ophiomisidium, 35, 36, 54
Linckiidae, 206
lineata, Sigsbeia, 11
Lissothuria, 136
ornata, 139
lonchophorus, Ophiophragmus, 16,
18, 48
longifissa, Mellita, 290, 291, 337
Lovenia, 347, 348
cordiformis, 227, 314, 315, 347
Loxechinus albus, 255, 259, 265
lubrica, Cucumaria, 78, 81, 82
lugubris, Pachythyone, 63, 66, 115, 116,
117, 117, 118, 180
‘Thyone, 116
Luidiaster, 201
californicus, 201, 205, 206, 216, 218
dawsoni, 205
horridus, 205
Lytechinus, 249, 260
anamesus, 247, 248, 249, 250, 253,
260
panamensis, 260
pictus, 248, 249, 250, 253, 260
semituberculatus, 226, 250, 253,
261, 262
macrolepis, Psolus, 150
magnifica, Alexandria, 295
ee Coronaster, 210, 213, 220,
22
marginatus, Ophiophragmus, 19
mathaei, Echinometra, 281, 292
mathaei oblonga, Echinometra, 281
Mellita longifissa, 290, 291, 337
Mellitella, 322
(Mellitella) Stokesii, Encope, 322
Meoma, 344
grandis, 227, 304, 307, 344
mexicana, Diadema, 235, 243
Thyonacta, 63, 65, 68, 69, 101,
102, 168
mexicanus, Centrechinus, 226, 235, 237,
242, 243
Dendraster, 271, 275, 313
micropora, Encope, xii, 282, 283, 320,
321, 322, 323, 329, 330
variety borealis, 329
variety californica, 329
micropora galapagensis, Encope, 323,
331
INDEX
microrhynchus, Inachoides, 3
miniata, Cucumaria, 78
minutus, Amphicontus, 2, 4
mionactis, Phataria, 206, 208, 220
Moera clotho, 341
Moira clotho, 298, 300, 341
clothro, 341
molle, Thyonidium, 130
mollis, Pattalus, 64, 66, 67, 70, 120,
IZ25AZS AZO 1275 28,2990, 0S)
molpadioides, Ocnus, 93
multispinus, Ophiomyces, 13, 15, 46
murrhina, Sigsbeia, 11
neofusus, Thyone, 63, 65, 69, 103,
104, 105, 170
Neopentamera, 76, 89, 90
anexigua, 63, 69, 90, 91, 160
Neothyone, 77, 108, 109
gibber, 63, 64, 67, 70, 108, 109,
110, 111, 112, 113, 174
gibbosa, 63, 64, 67, 68, 70, 92, 109,
110, 111, 113, 114, 118, 176, 178
panamensis, 63, 68. 70, 109, 111,
112, 113, 176
Neothyonidium, 135
New Eastern Pacific sea stars, 197
New Ophiurans of the Allan Hancock
Pacific Expedition, 9
niger, Echinus, 247
Tetrapygus, 247, 250, 252
nina, Leptopentacta, 63, 70, 93, 94, 97,
98, 164
nova, Leptopentacta, 63, 66, 93, 94, 95,
95, 96, 98, 162
nuda, Ophionereis, 32
nutriens, Psolidium, 138
Thyonepsolus, 63, 64, 137, 138, 139
140, 186
oblonga, Echinometra, 226, 257, 259,
281, 292
oblongus, Echinus, 281
obstipa, T'amaria, 208, 210, 220,
222
occidentalis, Euthyonidium, 124, 126,
ochrus, Clypeaster, 226, 266, 267, 297,
309
Ocnus, 92, 93
imbricatus, 93
javanicus, 93
molpadioides, 93
panamensis, 89
pygmaeus, 93
typicus, 93
Ophiacantha, 11
bairdi, 13
phragma, 11, 13, 44
Ophiacanthidae, 11
Ophiactis, 33
savignyi, 19
357
ophiactoides, Ophiophragmus, 18, 19, 46
Ophiochitonidae, 29
Ophiocnida, 25
californica, 25, 27, 50
hispida, 27
Ophiocoma alexanderi, 19
Ophiodesmus, 33
amphilogus, 33, 35, 58
Ophiolepididae, 35
Ophiolepis, 41
crassa, 42
plateia, 41, 42, 56
variegata, 42
Ophiomisidum, 35
leurum, 35, 36, 54
Ophiomyces, 13
multispinus, 13, 15, 46
Ophionereis, 29, 30, 33, 35
albomaculata, 32
annulata, 32, 34
dictyota, 29, 30, 32, 52
eurybrachiplax, 32, 34
nuda, 32
perplexa, 30, 32, 54
reticulata, 30
Ophiophragmus, 15, 19
chilensis, 19, 22
disacanthus, 15, 16, 58
lonchophorus, 16, 18, 48
marginatus, 19
ophiactoides, 18, 19, 46
papillatus, 19, 20, 48
paucispinus, 19
stellatus, 21, 22, 52
tabogensis, 19
Ophiothrix tabogae, 25
violacea, 25
ornata, Lissothuria, 139
ovulum, Euthonidium, 64, 65, 68, 70,
122, 124, 125, 126, 129, 130, 131, 182
Stolus, 124, 125
Thyone, 119, 122, 124, 125, 126
Pachythone, 77, 114, 115
lugubris, 63, 66, 115, 116, 117, 117,
118, 178, 180
pseudolugubris, 63, 64, 66, 69, 114,
115, 116, 117, 117, 118, 178, 180
rubra, 63, 114, 115, 116, 118, 180
pacifica, Brissopsis, 298, 301, 341
pacificus, Cassidulus, 226, 290, 291, 338
Plagiobrissus, 298, 302, 342
Pygorhynchus, 338
Toxobrissus, 341
Palaetropus, 345
panamensis, Dorocidaris, 230
Hesperocidaris, 226, 230, 231, 232,
238, 239
Leptopentacta, 98
Lytechinus, 260
358 INDEX
Neothyone, 63, 68, 70, 109, 111, 112,
113, 176
Ocnus, 89
Pentamera, 63, 89
Thyone, 112
panamica, Leptopentacta, 63, 65, 66, 67,
93, 93, 94, 96, 162
papillatus, Ophiophragmus, 19, 20, 48
Paracaudina, 94
parafusus, Thyone, 63, 66, 104, 106, 170
Parastropecten, 197
Parocnus, 93, 96
imbricata, 96
patagonicus, Psolus, 63, 64, 145, 147,
148, 194
Pattalus, 123, 125, 129
mollis, 64, 66, 67, 70, 120, 122,
125, 126, 127, 128, 129, 130, 131
peruana, 125, 130
peruanus, 131
paucispinus, Ophiophragmus, 19
paucisquama, Amphiophiura, 38, 39,
40, 56
Paulia horrida, 3
pauper, Psolus, 148
Pentacta, 77, 91, 92, 96, 98, 99, 102
doliolum, 99
peruana, 63, 66, 99, 166
Pentamera, 76, 77, 84, 85, 86, 89, 90,
91, 103
beebei, 63, 85, 86, 90, 158
calcigera, 74, 85
chierchia, 63, 65, 66, 67, 68, 69, 84,
85, 86, 89, 160
chiloensis, 63, 65, 69, 70, 85, 88,
89, 89, 160
panamensis, 63, 89
pulcherrima, 84
zacae, 63, 84, 85, 90, 158
perplexa, Hesperocidaris, 231, 232, 234,
238, 240
Ophionereis, 30, 32, 54
Tretocidaris, 231
perspectiva, Encope, 273, 275, 320, 321
323, 326, 328
perspectiva jonesi, Encope, 323
peruana, Anaperus, 119, 120, 121, 130
Holothuria, 129, 130
Pattalus, 125, 130
Pentacta, 63, 66, 99, 166
Trepang, 120
Thyone, 121
peruanum, Thyonidium, 130
peruanus, Anaperus, 120, 131
Colochirus, 99
Pattalus, 131
peruviana, Anaperus, 63, 120
Holothuria, 120
peruvianus, Phyllophorus, 130
phantapus, Psolus, 147
Pharia, 207
Phataria, 206, 207
mionactis, 206, 208, 220
philohelminthius, Amphioplus, 27, 28,
50
phragma, Ophiacantha, 11, 13, 44
Phyllophoridae, 71, 73, 122, 123
Phyllophorus, 122, 123, 132
aculeatus, 64, 108, 133, 184
destichadus, 135
peruvianus, 130, 132
urna, 132
zacae, 64, 65, 67, 70, 133, 134, 182
pictus, Lytechinus, 248, 249, 250, 253,
260
Psammechinus, 249
pileolus, Toxopneustes, 262
Pinnaxoides, 265, 276
Plagiobrissus, 342
pacificus, 298, 302, 342
planum, Psolidium, 64, 70, 143, 145,
194
plateia, Ophiolepis, 41, 42, 56
Porites, 64
Portunus (Achelous) angustus, 3
Psammechinus pictus, 249
(Psammechinus) semituberculatus,
Echinus, 261
pseudocurata, Cucumaria, 78
pseudofusus, Thyone, 103, 107
pseudolugubris, Pachythyone, 63, 64,
66, 69, 114, 115, 116, 117, 117,
118, 178, 180
pseudoquinquesemita, Eupentacta, 90,
91
Psolidae, 71, 73, 135, 136
Psolidium, 135, 136, 139, 140, 141,
142
bullatum, 144
convergens, 142
dorsipes, 64, 69, 70, 142, 143, 192
ekmani, 64, 65, 70, 143, 145, 148,
149, 194
eubullatum, 64, 65, 143, 144, 190
nutriens, 138
planum, 64, 70, 143, 145, 194
Psolus, 135, 136, 142, 146,147
antarcticus, 148
diomedeae, 64, 65, 67, 68, 69, 70,
147, 149, 194
macrolepis, 150
patagonicus, 63, 64, 145, 147, 148,
194
pauper, 148
phantapus, 147
segregatus, 148
squamatus, 148
INDEX 359
variety segregatus, 64, 145, 147,
148, 194
pulcherrima, Cucumaria, 88
Pentamera, 84
pulvinata, Astropyga, 236, 250, 251
variety venusta, 236
Cidarites, 236
purpuratus, Echinus, 279
Strongylocentrotus, 249, 256, 259,
279, 280, 281
pygmaeus, Ocnus, 93
Pygorhynchus pacificus, 338
Pyromaia tuberculata, 3
quadrangularis, Colochirus, 98
reticulata, Ophionereis, 30
rosea, Boletia, 262
roseus, Toxopneustes, 226, 250, 254, 262
rotundus, Clypeaster, 226, 266, 267, 296,
309
Stolonoclypus, 296
rubra, Pachythyone, 63, 114, 115, 118,
180
Thyone, 115, 116
rugosus, Dendraster, 272, 275, 317, 318
sabanillensis, Thyonacta, 101
salmini, Cucumaria, 82, 83
savignyi, Ophiactis, 19
Schizaster townsendi, 340
(Brisaster) townsendi, 340
scleroderma, Tamaria, 210
scrobiculata, Agassizia, 226, 298, 299,
339
sculptilis, Amphiodia, 24, 25, 50
Scutella excentrica, 310
Scutellidae, 310
segregatus, Psolus, 148
seguroensis, Euthyonidium, 124
semituberculatus, Lytechinus, 226, 250,
253, 261, 262
semituberculatus, Echinus (Psamme-
chinus), 261
Lytechinus, 226, 250, 253, 261, 262
Sigsbeia, 9
laevis, 9, 11, 44
lineata, 11
murrhina, 11
similis, Thyone, 110, 111
Spatangidae, 341
Spatangus, 227, 266, 268, 345, 346
brissus variety latecarinatus, 343
californicus, 305, 307, 345, 346
spatuliger, Echinus (Agarites), 246
spatuligera, Arbacia, 246, 250, 252
speciosus, Clypeaster, 297, 308, 309
Sphaerothuria, 76
squamatus, Psolus, 148
squamatus variety segregatus, Psolus,
64, 145, 147, 148, 194
Staurocucumaria, 76
Stegophiura, 40
stellata, Arbacia, 244
stellatus, ? Echinus, 245
Leptychaster (Parastropecten),
197, 200, 201, 216
Ophiophragmus, 21, 22, 52
stenopora, Heliocidaris, 293
Stenorhynchus debilis, 3
Stichopus japonicus, 72, 74
Stokesii, Encope, 322
Stolonoclypus rotundus, 296
Stolus, 108
gibber, 109, 113
ovulum, 124, 125
(Stolus) chilensis, Thyone, 127
a Thyone, 63, 104, 107, 134,
17
Strongylocentrotidae, 265
Strongylocentrotus, 278
albus, 265
fragilis, 276
franciscanus, 257, 259, 278, 279, 280,
281
purpuratus, 249, 256, 259, 279, 280,
281
tabogae, Ophiothrix, 25
tabogensis, Ophiophragmus, 19
tabulifera, Cucumaria, 88
Tamaria, 208
floridae, 210
obstipa, 208, 210, 220,
222
scleroderma, 210
tenella, Thyone, 121
Tetrapygus niger, 247, 250, 252
The Holothurioidea collected by the
Velero III during the years 1932 to
1938, 61
Thouarsii, 229
Cidaris, 229
Cidaris (Eucidaris) , 229
Eucidaris, 226, 229, 238, 239
variety galapagensis, 229
Thyonacta, 77, 101
mexicana, 63, 65, 68, 69, 101, 102,
168
sabanillensis, 101
Thyone, 77, 91, 93, 101, 102, 103, 113,
119, 120, 121, 122, 123, 126, 131
benti, 119
variety zacae, 119
bidentata, 63, 65, 66, 67, 69, 104,
105, 105, 170
briareus, 72, 74, 119, 120, 121, 122
chilensis, 122, 128
cigaro, 120
fusus, 103, 105, 120
gibber, 109, 110, 113
glasselli, 119
360 INDEX
lechleri, 119
lugubris, 116
neofusus, 63, 65, 69, 103, 104, 105,
170
ovulum, 119, 122, 124, 125, 126
panamensis, 112
parafusus, 63, 66, 104, 106, 170
peruana, 121
pseudofusus, 103, 107
rubra, 115
similis, 110, 111
strangeri, 63, 104, 107, 134, 172
tenella, 121
Thyone (Stolus) chilensis, 127
Thyonepsolus, 135, 136, 137, 140, 142
beebei, 64, 65, 66, 137, 139, 190
hancocki, 64, 65, 68, 70, 138, 140,
188
nutriens, 63, 64, 137, 138, 139, 140,
186
veleronis, 64, 137, 140, 186, 188
Thyonidium, 122, 123, 127, 132, 133
molle, 130
peruanum, 130
townsendi, Brisaster, 298, 300, 340, 342
Schizaster, 340
Schizaster (Brisaster), 340
Toxobrissus pacificus, 341
Toxocidaris franciscana, 278
Toxopneustes, 262, 263
pileolus, 262
roseus, 226, 250, 254, 262
(Toxopneustes) gibbosus, Echinus, 265
Trepang peruana, 120
Tretocidaris perplexa, 231
Tripneustes, 264, 265, 278
depressus, 226, 250, 254, 263, 264
tuberculata, Pyromaia, 3
typica, Leptopentacta, 93, 94, 96
typicus, Ocnus, 93
urna, Phyllophorus, 132
Van Brunti, Echinometra, 226, 258,
259, 281, 292, 293
variety rupicola, 293
variegata, Ophiolepis, 42
vegae, Cucumaria, 78
bees Euthyonidium, 64, 67, 126,
182
Ee 64, 137, 140, 186,
1
violacea, Ophiothrix, 25
volsellatus, Coronaster, 211, 212, 213
wetmorei, Encope, 274, 275, 320, 321,
323, 326, 327, 328
zacae, Pentamera, 63, 84, 85, 90, 158
Phyllophorus, 64, 65, 67, 70, 133,
134, 182
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ALLAN HANCOCK PACIFIC EXPEDITIONS
VOLUME 8 tS vv NUMBER 1
A NEW GENUS OF BRITTLE STARS,
AMPHICONTUS
(PLATE 1)
BY
ALEX HILL
RESEARCH ASSOCIATE
ALLAN HANCOCK FOUNDATION
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VOLUME 8 NUMBER 2
NEW OPHIURANS OF THE ALLAN HANCOCK
PACIFIC EXPEDITIONS
(PLATEs 2-9)
BY
FRED C. ZIESENHENNE
RESEARCH ASSOCIATE
ALLAN HANCOCK FOUNDATION
THE UNIVERSITY OF SOUTHERN CALIFORNIA
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1940
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ALLAN HANCOCK PACIFIC EXPEDITIONS
VOLUME 8 NUMBER 3
THE HOLOTHURIOIDEA COLLECTED BY
THE VELERO III DURING THE
YEARS 1932 to 1938
ParT I, DENDROCHIROTA
(Piates 10-30)
BY
ELISABETH DEICHMANN
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1941
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ALLAN HANCOCK PACIFIC EXPEDITIONS
VOLUME 8 NUMBER 4
NEW EASTERN PACIFIC SEA STARS
(PLATEs 31-34)
BY
FRED C. ZIESENHENNE
RESEARCH ASSISTANT
ALLAN HANCOCK FOUNDATION
THE UNIVERSITY OF SOUTHERN CALIFORNIA
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1942
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ALLAN HANCOCK PACIFIC EXPEDITIONS
VOLUME 8 NUMBER 5
A REPORT ON THE ECHINI
OF THE WARMER. EASTERN PACIFIC,
BASED ON THE COLLECTIONS OF THE
VELERO III
(PiateEs 35-71, TExT Ficures 1-3)
BY
THE UNIVERSITY OF SOUTHERN CALIFORNIA PRESS
LOS ANGELES, CALIFORNIA
1948
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